Leaf water potentials measured with a pressure chamber.
Boyer, J S
1967-01-01
Leaf water potentials were estimated from the sum of the balancing pressure measured with a pressure chamber and the osmotic potential of the xylem sap in leafy shoots or leaves. When leaf water potentials in yew, rhododendron, and sunflower were compared with those measured with a thermocouple psychrometer known to indicate accurate values of leaf water potential, determinations were within +/- 2 bars of the psychrometer measurements with sunflower and yew. In rhododendron. water potentials measured with the pressure chamber plus xylem sap were 2.5 bars less negative to 4 bars more negative than psychrometer measurements.The discrepancies in the rhododendron measurements could be attributed, at least in part, to the filling of tissues other than xylem with xylem sap during measurements with the pressure chamber. It was concluded that, although stem characteristics may affect the measurements, pressure chamber determinations were sufficiently close to psychrometer measurements that the pressure chamber may be used for relative measurements of leaf water potentials, especially in sunflower and yew. For accurate determinations of leaf water potential, however, pressure chamber measurements must be calibrated with a thermocouple psychrometer.
Leaf Water Potentials Measured with a Pressure Chamber
Boyer, J. S.
1967-01-01
Leaf water potentials were estimated from the sum of the balancing pressure measured with a pressure chamber and the osmotic potential of the xylem sap in leafy shoots or leaves. When leaf water potentials in yew, rhododendron, and sunflower were compared with those measured with a thermocouple psychrometer known to indicate accurate values of leaf water potential, determinations were within ± 2 bars of the psychrometer measurements with sunflower and yew. In rhododendron. water potentials measured with the pressure chamber plus xylem sap were 2.5 bars less negative to 4 bars more negative than psychrometer measurements. The discrepancies in the rhododendron measurements could be attributed, at least in part, to the filling of tissues other than xylem with xylem sap during measurements with the pressure chamber. It was concluded that, although stem characteristics may affect the measurements, pressure chamber determinations were sufficiently close to psychrometer measurements that the pressure chamber may be used for relative measurements of leaf water potentials, especially in sunflower and yew. For accurate determinations of leaf water potential, however, pressure chamber measurements must be calibrated with a thermocouple psychrometer. PMID:16656476
Use of Positive Pressures to Establish Vulnerability Curves 1
Cochard, Hervé; Cruiziat, Pierre; Tyree, Melvin T.
1992-01-01
Loss of hydraulic conductivity occurs in stems when the water in xylem conduits is subjected to sufficiently negative pressure. According to the air-seeding hypothesis, this loss of conductivity occurs when air bubbles are sucked into water-filled conduits through micropores adjacent to air spaces in the stem. Results in this study showed that loss of hydraulic conductivity occurred in stem segments pressurized in a pressure chamber while the xylem water was under positive pressure. Vulnerability curves can be defined as a plot of percentage loss of hydraulic conductivity versus the pressure difference between xylem water and the outside air inducing the loss of conductivity. Vulnerability curves were similar whether loss of conductivity was induced by lowering the xylem water pressure or by raising the external air pressure. These results are consistent with the air-seeding hypothesis of how embolisms are nucleated, but not with the nucleation of embolisms at hydrophobic cracks because the latter requires negative xylem water pressure. The results also call into question some basic underlying assumptions used in the determination of components of tissue water potential using “pressure-volume” analysis. PMID:16652947
Applications of the compensating pressure theory of water transport.
Canny, M
1998-07-01
Some predictions of the recently proposed theory of long-distance water transport in plants (the Compensating Pressure Theory) have been verified experimentally in sunflower leaves. The xylem sap cavitates early in the day under quite small water stress, and the compensating pressure P (applied as the tissue pressure of turgid cells) pushes water into embolized vessels, refilling them during active transpiration. The water potential, as measured by the pressure chamber or psychrometer, is not a measure of the pressure in the xylem, but (as predicted by the theory) a measure of the compensating pressure P. As transpiration increases, P is increased to provide more rapid embolism repair. In many leaf petioles this increase in P is achieved by the hydrolysis of starch in the starch sheath to soluble sugars. At night P falls as starch is reformed. A hypothesis is proposed to explain these observations by pressure-driven reverse osmosis of water from the ground parenchyma of the petiole. Similar processes occur in roots and are manifested as root pressure. The theory requires a pump to transfer water from the soil into the root xylem. A mechanism is proposed by which this pump may function, in which the endodermis acts as a one-way valve and a pressure-confining barrier. Rays and xylem parenchyma of wood act like the xylem parenchyma of petioles and roots to repair embolisms in trees. The postulated root pump permits a re-appraisal of the work done by evaporation during transpiration, leading to the proposal that in tall trees there is no hydrostatic gradient to be overcome in lifting water. Some published observations are re-interpreted in terms of the theory: doubt is cast on the validity of measurements of hydraulic conductance of wood; vulnerability curves are found not to measure the cavitation threshold of water in the xylem, but the osmotic pressure of the xylem parenchyma; if measures of xylem pressure and of hydraulic conductance are both suspect, the accepted view of the hydraulic architecture of trees needs drastic revision; observations that xylem feeding insects feed faster as the water potential becomes more negative are in accord with the theory; tyloses, which have been shown to form in vessels especially vulnerable to cavitation, are seen as necessary for the maintenance of P, and to conserve the supplementary refilling water. Far from being a metastable system on the edge of disaster, the water transport system of the xylem is ultrastable: robust and self-sustaining in response to many kinds of stress.
Diurnal patterns in Scots pine stem oleoresin pressure in a boreal forest.
Rissanen, K; Hölttä, T; Vanhatalo, A; Aalto, J; Nikinmaa, E; Rita, H; Bäck, J
2016-03-01
Coniferous tree stems contain large amounts of oleoresin under positive pressure in the resin ducts. Studies in North-American pines indicated that the stem oleoresin exudation pressure (OEP) correlates negatively with transpiration rate and soil water content. However, it is not known how the OEP changes affect the emissions of volatile vapours from the trees. We measured the OEP, xylem diameter changes indicating changes in xylem water potential and monoterpene emissions under field conditions in mature Scots pine (Pinus sylvestris L.) trees in southern Finland. Contrary to earlier reports, the diurnal OEP changes were positively correlated with temperature and transpiration rate. OEP was lowest at the top part of the stem, where water potentials were also more negative, and often closely linked to ambient temperature and stem monoterpene emissions. However, occasionally OEP was affected by sudden changes in vapour pressure deficit (VPD), indicating the importance of xylem water potential on OEP as well. We conclude that the oleoresin storage pools in tree stems are in a dynamic relationship with ambient temperature and xylem water potential, and that the canopy monoterpene emission rates may therefore be also regulated by whole tree processes and not only by the conditions prevailing in the upper canopy. © 2015 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.
What are the driving forces for water lifting in the xylem conduit?
Zimmermann, Ulrich; Schneider, Heike; Wegner, Lars H; Wagner, Hans-Jürgen; Szimtenings, Michael; Haase, Axel; Bentrup, Friedrich-Wilhelm
2002-03-01
After Renner had shown convincingly in 1925 that the transpirational water loss generates tensions larger than 0.1 MPa (i.e. negative pressures) in the xylem of cut leafy twigs the Cohesion Theory proposed by Böhm, Askenasy, Dixon and Joly at the end of the 19th century was immediately accepted by plant physiologists. Introduction of the pressure chamber technique by Scholander et al. in 1965 enforced the general belief that tension is the only driving force for water lifting although substantial criticism regarding the technique and/or the Cohesion Theory was published by several authors. As typical for scientific disciplines, the advent of minimal- and non-invasive techniques in the last decade as well as the development of a new, reliable method for xylem sap sampling have challenged this view. Today, xylem pressure gradients, potentials, ion concentrations and volume flows as well as cell turgor pressure gradients can be monitored online in intact transpiring higher plants, and within a given physiological context by using the pressure probe technique and high-resolution NMR imaging techniques, respectively. Application of the pressure probe technique to transpiring plants has shown that negative absolute pressures (down to - 0.6 MPa) and pressure gradients can exist temporarily in the xylem conduit, but that the magnitude and (occasionally) direction of gradients contrasts frequently the belief that tension is the only driving force. This seems to be particularly the case for plants faced with problems of height, drought, freezing and salinity as well as with cavitation of the tensile water. Reviewing the current data base shows that other forces come into operation when exclusively tension fails to lift water against gravity due to environmental conditions. Possible candidates are longitudinal cellular and xylem osmotic pressure gradients, axial potential gradients in the vessels as well as gel- and gas bubble-supported interfacial gradients. The multiforce theory overcomes the problem of the Cohesion Theory that life on earth depends on water being in a highly metastable state.
Hydraulic properties of rice and the response of gas exchange to water stress.
Stiller, Volker; Lafitte, H Renee; Sperry, John S
2003-07-01
We investigated the role of xylem cavitation, plant hydraulic conductance, and root pressure in the response of rice (Oryza sativa) gas exchange to water stress. In the field (Philippines), the percentage loss of xylem conductivity (PLC) from cavitation exceeded 60% in leaves even in watered controls. The PLC versus leaf water potential relationship indicated diurnal refilling of cavitated xylem. The leaf water potential causing 50 PLC (P(50)) was -1.6 MPa and did not differ between upland versus lowland rice varieties. Greenhouse-grown varieties (Utah) were more resistant to cavitation with a 50 PLC of -1.9 MPa but also showed no difference between varieties. Six-day droughts caused concomitant reductions in leaf-specific photosynthetic rate, leaf diffusive conductance, and soil-leaf hydraulic conductance that were associated with cavitation-inducing water potentials and the disappearance of nightly root pressure. The return of root pressure after drought was associated with the complete recovery of leaf diffusive conductance, leaf-specific photosynthetic rate, and soil-leaf hydraulic conductance. Root pressure after the 6-d drought (61.2 +/- 8.8 kPa) was stimulated 7-fold compared with well-watered plants before drought (8.5 +/- 3.8 kPa). The results indicate: (a) that xylem cavitation plays a major role in the reduction of plant hydraulic conductance during drought, and (b) that rice can readily reverse cavitation, possibly aided by nocturnal root pressure.
Wei, Chunfang; Tyree, Melvin T.; Steudle, Ernst
1999-01-01
The water relations of maize (Zea mays L. cv Helix) were documented in terms of hydraulic architecture and xylem pressure. A high-pressure flowmeter was used to characterize the hydraulic resistances of the root, stalk, and leaves. Xylem pressure measurements were made with a Scholander-Hammel pressure bomb and with a cell pressure probe. Evaporation rates were measured by gas exchange and by gravimetric measurements. Xylem pressure was altered by changing the light intensity, by controlling irrigation, or by gas pressure applied to the soil mass (using a root pressure bomb). Xylem pressure measured by the cell pressure probe and by the pressure bomb agreed over the entire measured range of 0 to −0.7 MPa. Experiments were consistent with the cohesion-tension theory. Xylem pressure changed rapidly and reversibly with changes in light intensity and root-bomb pressure. Increasing the root-bomb pressure increased the evaporation rate slightly when xylem pressure was negative and increased water flow rate through the shoots dramatically when xylem pressure was positive and guttation was observed. The hydraulic architecture model could predict all observed changes in water flow rate and xylem. We measured the cavitation threshold for oil- and water-filled pressure probes and provide some suggestions for improvement. PMID:10594106
Direct Demonstration of a Growth-Induced Water Potential Gradient.
Nonami, H.; Boyer, J. S.
1993-05-01
When transpiration is negligible, water potentials in growing tissues are less than those in mature tissues and have been predicted to form gradients that move water into the enlarging cells. To determine directly whether the gradients exist, we measured water potentials along the radius of stems of intact soybean (Glycine max [L.] Merr.) seedlings growing in vermiculite in a water-saturated atmosphere. The measurements were made in individual cells by first determining the turgor with a miniature pressure probe, then determining the osmotic potential of solution from the same cell, and finally summing the two potentials. The osmotic potentials were corrected for sample mixing in the probe. The measurements were checked with a thermocouple psychrometer that gave average tissue water potentials. In the elongating region, the water potential was highest near the xylem and lowest near the epidermis and in the center of the pith. In the basal, more mature region of the same stems, water potentials were near zero next to the xylem and throughout the tissue. These basal potentials reflected mostly the potential of the xylem, which extended into the elongating tissues. Thus, the high basal potential confirmed the high potential near the xylem in the elongating tissues. The psychrometer measurements for each tissue gave average potentials that agreed with the average of the cell potentials from the pressure probe. We conclude that a radial gradient was present in the elongating region that formed a water potential field in three dimensions around the xylem and that confirmed the predictions of Molz and Boyer (F.J. Molz and J.S. Boyer [1978] Plant Physiol 62: 423-429).
Direct Demonstration of a Growth-Induced Water Potential Gradient.
Nonami, H.; Boyer, J. S.
1993-01-01
When transpiration is negligible, water potentials in growing tissues are less than those in mature tissues and have been predicted to form gradients that move water into the enlarging cells. To determine directly whether the gradients exist, we measured water potentials along the radius of stems of intact soybean (Glycine max [L.] Merr.) seedlings growing in vermiculite in a water-saturated atmosphere. The measurements were made in individual cells by first determining the turgor with a miniature pressure probe, then determining the osmotic potential of solution from the same cell, and finally summing the two potentials. The osmotic potentials were corrected for sample mixing in the probe. The measurements were checked with a thermocouple psychrometer that gave average tissue water potentials. In the elongating region, the water potential was highest near the xylem and lowest near the epidermis and in the center of the pith. In the basal, more mature region of the same stems, water potentials were near zero next to the xylem and throughout the tissue. These basal potentials reflected mostly the potential of the xylem, which extended into the elongating tissues. Thus, the high basal potential confirmed the high potential near the xylem in the elongating tissues. The psychrometer measurements for each tissue gave average potentials that agreed with the average of the cell potentials from the pressure probe. We conclude that a radial gradient was present in the elongating region that formed a water potential field in three dimensions around the xylem and that confirmed the predictions of Molz and Boyer (F.J. Molz and J.S. Boyer [1978] Plant Physiol 62: 423-429). PMID:12231794
Seasonal variation in xylem pressure of walnut trees: root and stem pressures.
Ewers, F W; Améglio, T; Cochard, H; Beaujard, F; Martignac, M; Vandame, M; Bodet, C; Cruiziat, P
2001-09-01
Measurements of air and soil temperatures and xylem pressure were made on 17-year-old orchard trees and on 5-year-old potted trees of walnut (Juglans regia L.). Cooling chambers were used to determine the relationships between temperature and sugar concentration ([glucose] + [fructose] + [sucrose], GFS) and seasonal changes in xylem pressure development. Pressure transducers were attached to twigs of intact plants, root stumps and excised shoots while the potted trees were subjected to various temperature regimes in autumn, winter and spring. Osmolarity and GFS of the xylem sap (apoplast) were measured before and after cooling or warming treatments. In autumn and spring, xylem pressures of up to 160 kPa were closely correlated with soil temperature but were not correlated with GFS in xylem sap. High root pressures were associated with uptake of mineral nutrients from soil, especially nitrate. In autumn and spring, xylem pressures were detected in root stumps as well as in intact plants, but not in excised stems. In contrast, in winter, 83% of the xylem sap osmolarity in both excised stems and intact plants could be accounted for by GFS, and both GFS and osmolarity were inversely proportional to temperature. Plants kept at 1.5 degrees C developed positive xylem pressures up to 35 kPa, xylem sap osmolarities up to 260 mosmol l(-1) and GFS concentrations up to 70 g l(-1). Autumn and spring xylem pressures, which appeared to be of root origin, were about 55% of the theoretical pressures predicted by osmolarity of the xylem sap. In contrast, winter pressures appeared to be of stem origin and were only 7% of the theoretical pressures, perhaps because of a lower stem water content during winter.
Nonami, H; Schulze, E D
1989-01-01
Water potential, osmotic potential and turgor measurements obtained by using a cell pressure probe together with a nanoliter osmometer were compared with measurements obtained with an isopiestic psychrometer. Both types of measurements were conducted in the mature region of Tradescantia virginiana L. leaves under non-transpiring conditions in the dark, and gave similar values of all potentials. This finding indicates that the pressure probe and the osmometer provide accurate measurements of turgor, osmotic potentials and water potentials. Because the pressure probe does not require long equilibration times and can measure turgor of single cells in intact plants, the pressure probe together with the osmometer was used to determine in-situ cell water potentials, osmotic potentials and turgor of epidermal and mesophyll cells of transpiring leaves as functions of stomatal aperture and xylem water potential. When the xylem water potential was-0.1 MPa, the stomatal aperture was at its maximum, but turgor of both epidermal and mesophyll cells was relatively low. As the xylem water potential decreased, the stomatal aperture became gradually smaller, whereas turgor of both epidermal and mesophyll cells first increased and afterward decreased. Water potentials of the mesophyll cells were always lower than those of the epidermal cells. These findings indicate that evaporation of water is mainly occurring from mesophyll cells and that peristomatal transpiration could be less important than it has been proposed previously, although peristomatal transpiration may be directly related to regulation of turgor in the guard cells.
Water potential gradient in a tall sequoiadendron.
Tobiessen, P
1971-09-01
With an elevator installed in a 90-meter tall Sequoiadendron to collect the samples, xylem pressure potential measurements were made approximately every 15 meters along 60 meters of the tree's height. The measured gradient was about -0.8 bar per 10 meters of height, i.e., less than the hydrostatic gradient. Correction of the xylem pressure potential data by calibration against a thermocouple psychrometer confirmed this result. Similar gradients are described in the literature in tall conifers at times of low transpiration, although a different sampling technique was used. If the data in the present study and those supporting it are typical, they imply a re-evaluation of either the use of the pressure chamber to estimate water potential or the present theories describing water transport in tall trees.
Water Potential Gradient in a Tall Sequoiadendron
Tobiessen, Peter; Rundel, Philip W.; Stecker, R. E.
1971-01-01
With an elevator installed in a 90-meter tall Sequoiadendron to collect the samples, xylem pressure potential measurements were made approximately every 15 meters along 60 meters of the tree's height. The measured gradient was about −0.8 bar per 10 meters of height, i.e., less than the hydrostatic gradient. Correction of the xylem pressure potential data by calibration against a thermocouple psychrometer confirmed this result. Similar gradients are described in the literature in tall conifers at times of low transpiration, although a different sampling technique was used. If the data in the present study and those supporting it are typical, they imply a re-evaluation of either the use of the pressure chamber to estimate water potential or the present theories describing water transport in tall trees. PMID:16657786
Refilling of a Hydraulically Isolated Embolized Xylem Vessel: Model Calculations
VESALA, TIMO; HÖLTTÄ, TEEMU; PERÄMÄKI, MARTTI; NIKINMAA, EERO
2003-01-01
When they are hydraulically isolated, embolized xylem vessels can be refilled, while adjacent vessels remain under tension. This implies that the pressure of water in the refilling vessel must be equal to the bubble gas pressure, which sets physical constraints for recovery. A model of water exudation into the cylindrical vessel and of bubble dissolution based on the assumption of hydraulic isolation is developed. Refilling is made possible by the turgor of the living cells adjacent to the refilling vessel, and by a reflection coefficient below 1 for the exchange of solutes across the interface between the vessel and the adjacent cells. No active transport of solutes is assumed. Living cells are also capable of importing water from the water‐conducting vessels. The most limiting factors were found to be the osmotic potential of living cells and the ratio of the volume of the adjacent living cells to that of the embolized vessel. With values for these of 1·5 MPa and 1, respectively, refilling times were in the order of hours for a broad range of possible values of water conductivity coefficients and effective diffusion distances for dissolved air, when the xylem water tension was below 0·6 MPa and constant. Inclusion of the daily pattern for xylem tension improved the simulations. The simulated gas pressure within the refilling vessel was in accordance with recent experimental results. The study shows that the refilling process is physically possible under hydraulic isolation, while water in surrounding vessels is under negative pressure. However, the osmotic potentials in the refilling vessel tend to be large (in the order of 1 MPa). Only if the xylem water tension is, at most, twice atmospheric pressure, the reflection coefficient remains close to 1 (0·95) and the ratio of the volume of the adjacent living cells to that of the embolized vessel is about 2, does the osmotic potential stay below 0·4 MPa. PMID:12588721
Photosynthesis and leaf water relations in four American sycamore clones
DOE Office of Scientific and Technical Information (OSTI.GOV)
Tang, Z.; Land, S.B. Jr.
1995-11-01
Photosynthesis, transpiration, stomatal conductance, and xylem pressure potential were studied to examine clonal variation and clone-by-season interactions in rooted cuttings of four sycamore clones (Platanus occidentalis L.). These physiological parameters were measured during June through November of the second and third growing seasons in the field. Stomatal conductance, xylem pressure potential, and photosynthesis were higher in June-July than in August-November. The four clones did not differ significantly in yearly average photosynthetic rates, but clone 11 tended to have higher rates early in each growing season (June-July) than did the other three clones. Dry periods during August-September of the second seasonmore » and during October of the third season apparently caused clone 11 to close its stomata more than clone 17, as indicated by significant clone-by-season interactions for reductions in stomatal conductance and transpiration late in the morning. Clone 17 was generally able to maintain high xylem pressure potential, stomatal conductance, and transpiration throughout the growing season, probably because of its large root system. 36 refs., 2 figs., 5 tabs.« less
Water relations in silver birch during springtime: How is sap pressurised?
Hölttä, T; Dominguez Carrasco, M D R; Salmon, Y; Aalto, J; Vanhatalo, A; Bäck, J; Lintunen, A
2018-05-06
Positive sap pressures are produced in the xylem of birch trees in boreal conditions during the time between the thawing of the soil and bud break. During this period, xylem embolisms accumulated during wintertime are refilled with water. The mechanism for xylem sap pressurization and its environmental drivers are not well known. We measured xylem sap flow, xylem sap pressure, xylem sap osmotic concentration, xylem and whole stem diameter changes, and stem and root non-structural carbohydrate concentrations, along with meteorological conditions at two sites in Finland during and after the sap pressurisation period. The diurnal dynamics of xylem sap pressure and sap flow during the sap pressurisation period varied, but were more often opposite to the diurnal pattern after bud burst, i.e. sap pressure increased and sap flow rate mostly decreased when temperature increased. Net conversion of soluble sugars to starch in the stem and roots occurred during the sap pressurisation period. Xylem sap osmotic pressure was small in comparison to total sap pressure, and it did not follow changes in environmental conditions or tree water relations. Based on these findings, we suggest that xylem sap pressurisation and embolism refilling occur gradually over a few weeks through water transfer from parenchyma cells to xylem vessels during daytime, and then the parenchyma are refilled mostly during nighttime by water uptake from soil. Possible drivers for water transfer from parenchyma cells to vessels are discussed. Also the functioning of thermal dissipation probes in conditions of changing stem water content is discussed. © 2018 German Society for Plant Sciences and The Royal Botanical Society of the Netherlands.
Zwieniecki, Maciej A; Melcher, Peter J; Feild, Taylor S; Holbrook, N Michele
2004-08-01
We investigated phloem-xylem interactions in Acer rubrum L. and Acer saccharum Marsh. Our experimental method allowed us to determine xylem conductance of an intact branch by measuring the flow rate of water supplied at two delivery pressures to the cut end of a small side branch. We found that removal of bark tissue (phloem girdling) upstream of the point at which deionized water was delivered to the branch resulted in a decrease (24% for A. rubrum and 15% for A. saccharum) in branch xylem hydraulic conductance. Declines in hydraulic conductance with girdling were accompanied by a decrease in the osmotic concentration of xylem sap. The decrease in xylem sap concentration following phloem girdling suggests that ion redistribution from the phloem was responsible for the observed decline in hydraulic conductance. When the same measurements were made on branches perfused with KCl solution (approximately 140 mOsm kg(-1)), phloem girdling had no effect on xylem hydraulic conductance. These results suggest a functional link between phloem and xylem hydraulic systems that is mediated by changes in the ionic content of the cell sap.
Melcher, P J; Goldstein, G; Meinzer, F C; Yount, D E; Jones, T J; Holbrook, N M; Huang, C X
2001-01-01
Physiological traits related to water transport were studied in Rhizophora mangle (red mangrove) growing in coastal and estuarine sites in Hawaii. The magnitude of xylem pressure potential (P x ), the vulnerability of xylem to cavitation, the frequency of embolized vessels in situ, and the capacity of R. mangle to repair embolized vessels were evaluated with conventional and recently developed techniques. The osmotic potential of the interstitial soil water (π sw ) surrounding the roots of R. mangle was c. -2.6±5.52×10 -3 and -0.4±6.13×10 -3 MPa in the coastal and estuarine sites, respectively. Midday covered (non-transpiring) leaf water potentials (Ψ L ) determined with a pressure chamber were 0.6-0.8 MPa more positive than those of exposed, freely-transpiring leaves, and osmotic potential of the xylem sap (π x ) ranged from -0.1 to -0.3 MPa. Consequently, estimated midday values of P x (calculated by subtracting π x from covered Ψ L ) were about 1 MPa more positive than Ψ L determined on freely transpiring leaves. The differences in Ψ L between covered and transpiring leaves were linearly related to the transpiration rates. The slope of this relationship was steeper for the coastal site, suggesting that the hydraulic resistance was larger in leaves of coastal R. mangle plants. This was confirmed by both hydraulic conductivity measurements on stem segments and high-pressure flowmeter studies made on excised leafy twigs. Based on two independent criteria, loss of hydraulic conductivity and proportions of gas- and liquid-filled vessels in cryo-scanning electron microscope (cryo-SEM) images, the xylem of R. mangle plants growing at the estuarine site was found to be more vulnerable to cavitation than that of plants growing at the coastal site. However, the cryo-SEM analyses suggested that cavitation occurred more readily in intact plants than in excised branches that were air-dried in the laboratory. Cryo-SEM analyses also revealed that, in both sites, the proportion of gas-filled vessels was 20-30% greater at midday than at dawn or during the late afternoon. Refilling of cavitated vessels thus occurred during the late afternoon when considerable tension was present in neighboring vessels. These results and results from pressure-volume relationships suggest that R. mangle adjusts hydraulic properties of the water-transport system, as well as the leaf osmotic potential, in concert with the environmental growing conditions.
Three types of cavitation caused by air seeding.
Shen, Fanyi; Wang, Yuansheng; Cheng, Yanxia; Zhang, Li
2012-11-01
There are different opinions of the dynamics of an air bubble entering a xylem conduit. In this paper, we present a thorough mechanical analysis and conclude that there are three types of cavitation caused by air seeding. After an air seed enters a conduit at high xylem pressure P'(1), along with the drop of the water potential, it will expand gradually to a long-shaped bubble and extend continually. This is the first type of air seeding, or the type of expanding gradually. When the xylem pressure is moderate, right after an air seed enters a conduit, it will expand first. Then, as soon as the pressure reaches a threshold the bubble will blow up to form a bubble in long shape, accompanied by acoustic (or ultra-acoustic) emission. It will extend further as xylem pressure decreases continually. This is the second type of air seeding, or the type of expanding-exploding, becoming a long-shaped bubble-lengthening by degrees. In the range of P'(1) ≤ - 3P(o) (P(o) is atmospheric pressure), soon after an air seed is sucked into a conduit it will explode immediately and the conduit will be full of the gas of the bubble instantly. This is the third type of air seeding, or the type of sudden exploding and filling conduit instantly. The third type is the frequent event in daily life of plant.
Kröber, Wenzel; Zhang, Shouren; Ehmig, Merten; Bruelheide, Helge
2014-01-01
While the fundamental trade-off in leaf traits related to carbon capture as described by the leaf economics spectrum is well-established among plant species, the relationship of the leaf economics spectrum to stem hydraulics is much less known. Since carbon capture and transpiration are coupled, a close connection between leaf traits and stem hydraulics should be expected. We thus asked whether xylem traits that describe drought tolerance and vulnerability to cavitation are linked to particular leaf traits. We assessed xylem vulnerability, using the pressure sleeve technique, and anatomical xylem characteristics in 39 subtropical tree species grown under common garden conditions in the BEF-China experiment and tested for correlations with traits related to the leaf economics spectrum as well as to stomatal control, including maximum stomatal conductance, vapor pressure deficit at maximum stomatal conductance and vapor pressure deficit at which stomatal conductance is down-regulated. Our results revealed that specific xylem hydraulic conductivity and cavitation resistance were closely linked to traits represented in the leaf economic spectrum, in particular to leaf nitrogen concentration, as well as to log leaf area and leaf carbon to nitrogen ratio but not to any parameter of stomatal conductance. The study highlights the potential use of well-known leaf traits from the leaf economics spectrum to predict plant species' drought resistance. PMID:25423316
Kröber, Wenzel; Zhang, Shouren; Ehmig, Merten; Bruelheide, Helge
2014-01-01
While the fundamental trade-off in leaf traits related to carbon capture as described by the leaf economics spectrum is well-established among plant species, the relationship of the leaf economics spectrum to stem hydraulics is much less known. Since carbon capture and transpiration are coupled, a close connection between leaf traits and stem hydraulics should be expected. We thus asked whether xylem traits that describe drought tolerance and vulnerability to cavitation are linked to particular leaf traits. We assessed xylem vulnerability, using the pressure sleeve technique, and anatomical xylem characteristics in 39 subtropical tree species grown under common garden conditions in the BEF-China experiment and tested for correlations with traits related to the leaf economics spectrum as well as to stomatal control, including maximum stomatal conductance, vapor pressure deficit at maximum stomatal conductance and vapor pressure deficit at which stomatal conductance is down-regulated. Our results revealed that specific xylem hydraulic conductivity and cavitation resistance were closely linked to traits represented in the leaf economic spectrum, in particular to leaf nitrogen concentration, as well as to log leaf area and leaf carbon to nitrogen ratio but not to any parameter of stomatal conductance. The study highlights the potential use of well-known leaf traits from the leaf economics spectrum to predict plant species' drought resistance.
Dynamic control of osmolality and ionic composition of the xylem sap in two mangrove species.
López-Portillo, Jorge; Ewers, Frank W; Méndez-Alonzo, Rodrigo; Paredes López, Claudia L; Angeles, Guillermo; Alarcón Jiménez, Ana Luisa; Lara-Domínguez, Ana Laura; Torres Barrera, María Del Carmen
2014-06-01
• Premise of the study: Xylem sap osmolality and salinity is a critical unresolved issue in plant function with impacts on transport efficiency, pressure gradients, and living cell turgor pressure, especially for halophytes such as mangrove trees.• Methods: We collected successive xylem vessel sap samples from stems and shoots of Avicennia germinans and Laguncularia racemosa using vacuum and pressure extraction and measured their osmolality. Following a series of extractions with the pressure chamber, we depressurized the shoot and pressurized again after various equilibration periods (minutes to hours) to test for dynamic control of osmolality. Transpiration and final sap osmolality were measured in shoots perfused with deionized water or different seawater dilutions.• Key results: For both species, the sap osmolality values of consecutive samples collected by vacuum extraction were stable and matched those of the initial samples extracted with the pressure chamber. Further extraction of samples with the pressure chamber decreased sap osmolality, suggesting reverse osmosis occurred. However, sap osmolalities increased when longer equilibration periods after sap extraction were allowed. Analysis of expressed sap with HPLC indicated a 1:1 relation between measured osmolality and the osmolality of the inorganic ions in the sap (mainly Na + , K + , and Cl - ), suggesting no contamination by organic compounds. In stems perfused with deionized water, the sap osmolality increased to mimic the native sap osmolality.• Conclusions: Xylem sap osmolality and ionic contents are dynamically adjusted by mangroves and may help modulate turgor pressure, hydraulic conductivity, and water potential, thus being important for mangrove physiology, survival, and distribution. © 2014 Botanical Society of America, Inc.
Water filtration using plant xylem.
Boutilier, Michael S H; Lee, Jongho; Chambers, Valerie; Venkatesh, Varsha; Karnik, Rohit
2014-01-01
Effective point-of-use devices for providing safe drinking water are urgently needed to reduce the global burden of waterborne disease. Here we show that plant xylem from the sapwood of coniferous trees--a readily available, inexpensive, biodegradable, and disposable material--can remove bacteria from water by simple pressure-driven filtration. Approximately 3 cm(3) of sapwood can filter water at the rate of several liters per day, sufficient to meet the clean drinking water needs of one person. The results demonstrate the potential of plant xylem to address the need for pathogen-free drinking water in developing countries and resource-limited settings.
Water Filtration Using Plant Xylem
Chambers, Valerie; Venkatesh, Varsha; Karnik, Rohit
2014-01-01
Effective point-of-use devices for providing safe drinking water are urgently needed to reduce the global burden of waterborne disease. Here we show that plant xylem from the sapwood of coniferous trees – a readily available, inexpensive, biodegradable, and disposable material – can remove bacteria from water by simple pressure-driven filtration. Approximately 3 cm3 of sapwood can filter water at the rate of several liters per day, sufficient to meet the clean drinking water needs of one person. The results demonstrate the potential of plant xylem to address the need for pathogen-free drinking water in developing countries and resource-limited settings. PMID:24587134
NASA Technical Reports Server (NTRS)
Stahlberg, R.; Cosgrove, D. J.
1995-01-01
Excision of a growing stem causes local wound responses, such as membrane depolarization and growth inhibition, as well as effects at larger distances from the cut. In this study, cucumber hypocotyls were excised 100 mm below the hook, so that the growing region was beyond the reach of the wound-induced depolarization (up to 40 mm). Even at such a distance, the cut still caused a considerable and rapid drop in the hypocotyl growth rate. This growth response is not a direct wound response because it does not result from the cut-induced depolarization and because it can be simulated by root pressure manipulation (using a pressure chamber). The results indicate that the growth response resulted from the rapid release of the xylem pressure upon excision. To test this conclusion we measured the xylem pressure by connecting a pressure probe to the cut surface of the stem. Xylem pressure (Px) was found to be +10 to +40 kPa in cucumber hypocotyls and -5 to -10 kPa or lower in pea epicotyls. Excision of the cucumber hypocotyl base led to a rapid drop in Px to negative values, whereas excision in pea led to a rapid rise in Px to ambient (zero) pressure. These fast and opposite Px changes parallel the excision-induced changes in growth rate (GR): a decrease in cucumber and a rise in pea. The sign of the endogenous xylem pressure also determined whether excision induced a propagating depolarization in the form of a slow wave potential (SWP). Under normal circumstances pea seedlings generated an SWP upon excision whereas cucumber seedlings failed to do so. When the Px in cucumber hypocotyls was experimentally inverted to negative values by incubating the cumber roots in solutions of NaCN or n-ethylmaleimide, excision caused a propagating depolarization (SWP). The experiment shows that only hydraulic signals in the form of positive Px steps are converted into propagating electric SWP signals. These propagating depolarizations might be causally linked to systemic 'wound' responses, which occur independently of the short-distance or direct wound responses.
Cun-Yang Niu; Frederick C. Meinzer; Guang-You Hao
2017-01-01
1. In temperate ecosystems, freeze-thaw events are an important environmental stress that can induce severe xylem embolism (i.e. clogging of conduits by air bubbles) in overwintering organs of trees. However, no comparative studies of different adaptive strategies among sympatric tree species for coping with winter embolism have examined the potential role of the...
Zhenmin Tang; Jim L. Chambers; Suresh Guddanti; James P. Barnett
1999-01-01
Summary To examine physiological responses to thinning, fertilization, and crown position, we measured net photosynthesis (Pn), transpiration (E), vapor pressure difference (VPD), stomatal conductance (gs), and xylem pressure potential (Yl) between 0930 and 1130 h under ambient conditions in the...
Glassy-winged sharpshooter feeding does not cause air embolisms in xylem of well-watered plants.
USDA-ARS?s Scientific Manuscript database
Plant xylem vessels are under negative hydrostatic pressure (tension) as evapotranspiration of water from the leaf surface pulls the column of water in xylem upwards. When xylem fluid flux is under extreme tension, any puncture or breakage of the xylem vessel wall can cause formation of air embolis...
Physical analysis of the process of cavitation in xylem sap.
Shen, Fanyi; Gao, Rongfu; Liu, Wenji; Zhang, Wenjie
2002-06-01
Recent studies have confirmed that cavitation in xylem is caused by air bubbles. We analyzed expansion of a preexistent bubble adhering to a crack in a conduit wall and a bubble formed by the passage of air through a pore of a pit membrane, a process known as air seeding. We consider that there are two equilibrium states for a very small air bubble in the xylem: one is temporarily stable with a bubble radius r1 at point s1 on the curve P(r) relating pressure within the bubble (P) with bubble radius (r); the other is unstable with a bubble radius r2 at point s2 on Pr (where r1 < r2). In each equilibrium state, the bubble collapse pressure (2sigma/r, where sigma is surface tension of water) is balanced by the pressure difference across its surface. In the case of a bubble from a crack in a conduit wall, which is initially at point s1, expansion will occur steadily as water potential decreases. The bubble will burst only if the xylem pressure drops below a threshold value. A formula giving the threshold pressure for bubble bursting is proposed. In the case of an air seed entering a xylem conduit through a pore in a pit membrane, its initial radius may be r2 (i.e., the radius of the pore by which the air seed entered the vessel) at point s2 on Pr. Because the bubble is in an unstable equilibrium when entering the conduit, it can either expand or contract to point s1. As water vaporizes into the air bubble at s2, P rises until it exceeds the gas pressure that keeps the bubble in equilibrium, at which point the bubble will burst and induce a cavitation event in accordance with the air-seeding hypothesis. However, other possible perturbations could make the air-seeded bubble contract to s1, in which case the bubble will burst at a threshold pressure proposed for a bubble expanding from a crack in a conduit wall. For this reason some cavitation events may take place at a xylem threshold pressure (Pl'*) other than that determined by the formula, Plp'* = -2sigma/rp, proposed by Sperry and Tyree (1988), which is applicable only to air-seeded bubbles at s2. The more general formula we propose for calculating the threshold pressure for bubble breaking is consistent with the results of published experiments.
Frederick C. Meinzer; Daniel M. Johnson; Barbara Lachenbruch; Katherine A. McCulloh; David R. Woodruff
2009-01-01
The xylem pressure inducing 50% loss of hydraulic conductivity due to embolism (P50) is widely used for comparisons of xylem vulnerability among species and across aridity gradients. However, despite its utility as an index of resistance to catastrophic xylem failure under extreme drought, P50 may have no special...
Evidence for Hydraulic Vulnerability Segmentation and Lack of Xylem Refilling under Tension.
Charrier, Guillaume; Torres-Ruiz, José M; Badel, Eric; Burlett, Regis; Choat, Brendan; Cochard, Herve; Delmas, Chloe E L; Domec, Jean-Christophe; Jansen, Steven; King, Andrew; Lenoir, Nicolas; Martin-StPaul, Nicolas; Gambetta, Gregory Alan; Delzon, Sylvain
2016-11-01
The vascular system of grapevine (Vitis spp.) has been reported as being highly vulnerable, even though grapevine regularly experiences seasonal drought. Consequently, stomata would remain open below water potentials that would generate a high loss of stem hydraulic conductivity via xylem embolism. This situation would necessitate daily cycles of embolism repair to restore hydraulic function. However, a more parsimonious explanation is that some hydraulic techniques are prone to artifacts in species with long vessels, leading to the overestimation of vulnerability. The aim of this study was to provide an unbiased assessment of (1) the vulnerability to drought-induced embolism in perennial and annual organs and (2) the ability to refill embolized vessels in two Vitis species X-ray micro-computed tomography observations of intact plants indicated that both Vitis vinifera and Vitis riparia were relatively vulnerable, with the pressure inducing 50% loss of stem hydraulic conductivity = -1.7 and -1.3 MPa, respectively. In V. vinifera, both the stem and petiole had similar sigmoidal vulnerability curves but differed in pressure inducing 50% loss of hydraulic conductivity (-1.7 and -1 MPa for stem and petiole, respectively). Refilling was not observed as long as bulk xylem pressure remained negative (e.g. at the apical part of the plants; -0.11 ± 0.02 MPa) and change in percentage loss of conductivity was 0.02% ± 0.01%. However, positive xylem pressure was observed at the basal part of the plant (0.04 ± 0.01 MPa), leading to a recovery of conductance (change in percentage loss of conductivity = -0.24% ± 0.12%). Our findings provide evidence that grapevine is unable to repair embolized xylem vessels under negative pressure, but its hydraulic vulnerability segmentation provides significant protection of the perennial stem. © 2016 American Society of Plant Biologists. All Rights Reserved.
Evidence for Hydraulic Vulnerability Segmentation and Lack of Xylem Refilling under Tension1[OPEN
Charrier, Guillaume; Choat, Brendan; Delmas, Chloe E. L.; Domec, Jean-Christophe; King, Andrew; Lenoir, Nicolas
2016-01-01
The vascular system of grapevine (Vitis spp.) has been reported as being highly vulnerable, even though grapevine regularly experiences seasonal drought. Consequently, stomata would remain open below water potentials that would generate a high loss of stem hydraulic conductivity via xylem embolism. This situation would necessitate daily cycles of embolism repair to restore hydraulic function. However, a more parsimonious explanation is that some hydraulic techniques are prone to artifacts in species with long vessels, leading to the overestimation of vulnerability. The aim of this study was to provide an unbiased assessment of (1) the vulnerability to drought-induced embolism in perennial and annual organs and (2) the ability to refill embolized vessels in two Vitis species X-ray micro-computed tomography observations of intact plants indicated that both Vitis vinifera and Vitis riparia were relatively vulnerable, with the pressure inducing 50% loss of stem hydraulic conductivity = −1.7 and −1.3 MPa, respectively. In V. vinifera, both the stem and petiole had similar sigmoidal vulnerability curves but differed in pressure inducing 50% loss of hydraulic conductivity (−1.7 and −1 MPa for stem and petiole, respectively). Refilling was not observed as long as bulk xylem pressure remained negative (e.g. at the apical part of the plants; −0.11 ± 0.02 MPa) and change in percentage loss of conductivity was 0.02% ± 0.01%. However, positive xylem pressure was observed at the basal part of the plant (0.04 ± 0.01 MPa), leading to a recovery of conductance (change in percentage loss of conductivity = −0.24% ± 0.12%). Our findings provide evidence that grapevine is unable to repair embolized xylem vessels under negative pressure, but its hydraulic vulnerability segmentation provides significant protection of the perennial stem. PMID:27613852
NASA Astrophysics Data System (ADS)
Mrad, A.; Domec, J. C.; Huang, C. W.; Katul, G. G.
2017-12-01
Xylem tissues are specialized in offering the least possible resistance to water flow. However, this is not guaranteed when ascending sap reaches large negative pressures during periods of water stress when embolism within the xylem occurs, an inevitable step toward potential drought-induced mortality. Ongoing research into changing forest patterns and plant survival due to droughts rarely dispute the significance of Vulnerability Curves (VCs), plots that feature loss in relative conductance with declining liquid pressure (ψ). While Earth-Systems models routinely employ various VC functions, the theoretical underpinnings describing their shape remains lacking. VCs are the outcome of microscopic phenomena describing embolism formation, bubble-scale xylem properties allowing embolism spread, and hydraulic processes that dictate the water potential along the flow path. The work here explores how the upscaled version of these gives rise to popular mathematical shapes used to describe VC measurements: The Logistic and Weibull exceedance equations. Each of these two captures a distinct type of embolism spread: The Logistic VC arises when the probability that embolized vessels interact with intact ones describe embolism spread as water pressure decreases (labeled as a similarity law in botanic [1]). The Weibull VC arises when the aforementioned description includes the effects of ψ explicitly. Variability in xylem properties along the flow path is explored analytically using novel approaches borrowed from `super-statistics' and numerical simulations. The numerical simulations intend to single out which xylem network property is significant in describing the VC shape. The model results corroborate previous research (experimental and 3-dimensional high-resolution simulations) on the effect of vessel size and network topology. It is shown that (i) initial embolism locations alter air-seeding pressure and VC slope; (ii) redundancy and size variations decrease bubble propagation probability with decreasing ψ; and (iii) varying network properties spatially along the flow path delays total loss of conductance. The implications of these findings to linking VC shape to plant traits and the on-going debate about `r' vs `s' shape VCs are highlighted.
Characterization of Optimum Physiological Responses of Field-Grown Loblolly Pine
Zhenmin Tang; Jim L. Chambers; James P. Barnett
1999-01-01
Photosynthetic photon flux density (PPFD), air temperature (Ta), needle net photosynthesis (Pn), vapor pressure difference (VPD), stomata1 conductance (gw), transpiration (E), and predawn and daytime xylem pressure potentials (XPP) were measured in a loblolly pine (Pinus taeda L.) plantation in 1995 and 1996. Boundary-line analyses were conducted...
Hultine, K R; Koepke, D F; Pockman, W T; Fravolini, A; Sperry, J S; Williams, D G
2006-03-01
We investigated hydraulic constraints on water uptake by velvet mesquite (Prosopis velutina Woot.) at a site with sandy-loam soil and at a site with loamy-clay soil in southeastern Arizona, USA. We predicted that trees on sandy-loam soil have less negative xylem and soil water potentials during drought and a lower resistance to xylem cavitation, and reach E(crit) (the maximum steady-state transpiration rate without hydraulic failure) at higher soil water potentials than trees on loamy-clay soil. However, minimum predawn leaf xylem water potentials measured during the height of summer drought were significantly lower at the sandy-loam site (-3.5 +/- 0.1 MPa; all errors are 95% confidence limits) than at the loamy-clay site (-2.9 +/- 0.1 MPa). Minimum midday xylem water potentials also were lower at the sandy-loam site (-4.5 +/- 0.1 MPa) than at the loamy-clay site (-4.0 +/- 0.1 MPa). Despite the differences in leaf water potentials, there were no significant differences in either root or stem xylem embolism, mean cavitation pressure or Psi(95) (xylem water potential causing 95% cavitation) between trees at the two sites. A soil-plant hydraulic model parameterized with the field data predicted that E(crit) approaches zero at a substantially higher bulk soil water potential (Psi(s)) on sandy-loam soil than on loamy-clay soil, because of limiting rhizosphere conductance. The model predicted that transpiration at the sandy-loam site is limited by E(crit) and is tightly coupled to Psi(s) over much of the growing season, suggesting that seasonal transpiration fluxes at the sandy-loam site are strongly linked to intra-annual precipitation pulses. Conversely, the model predicted that trees on loamy-clay soil operate below E(crit) throughout the growing season, suggesting that fluxes on fine-textured soils are closely coupled to inter-annual changes in precipitation. Information on the combined importance of xylem and rhizosphere constraints to leaf water supply across soil texture gradients provides insight into processes controlling plant water balance and larger scale hydrologic processes.
Melcher, Peter J.; Zwieniecki, Maciej A.; Holbrook, N. Michele
2003-01-01
The relation between xylem vessel age and vulnerability to cavitation of sugar maple (Acer saccharum Marsh.) was quantified by measuring the pressure required to force air across bordered pit membranes separating individual xylem vessels. We found that the bordered pit membranes of vessels located in current year xylem could withstand greater applied gas pressures (3.8 MPa) compared with bordered pit membranes in vessels located in older annular rings (2.0 MPa). A longitudinal transect along 6-year-old branches indicated that the pressure required to push gas across bordered pit membranes of current year xylem did not vary with distance from the growing tip. To understand the contribution of age-related changes in vulnerability to the overall resistance to cavitation, we combined data on the pressure thresholds of individual xylem vessels with measurements of the relative flow rate through each annual ring. The annual ring of the current year contributed only 16% of the total flow measured on 10-cm-long segments cut from 6-year-old branches, but it contributed more than 70% of the total flow when measured through 6-year-old branches to the point of leaf attachment. The vulnerability curve calculated using relative flow rates measured on branch segments were similar to vulnerability curves measured on 6-year-old branches (pressure that reduces hydraulic conductance by 50% = 1.6–2.4 MPa), whereas the vulnerability curve calculated using relative flow rates measured on 6-year-old branches were similar to ones measured on the extension growth of the current year (pressure that reduces hydraulic conductance by 50% = 3.8 MPa). These data suggest that, in sugar maple, the xylem of the current year can withstand larger xylem tensions than older wood and dominates water delivery to leaves. PMID:12692336
Martin-StPaul, N K; Longepierre, D; Huc, R; Delzon, S; Burlett, R; Joffre, R; Rambal, S; Cochard, H
2014-08-01
Three methods are in widespread use to build vulnerability curves (VCs) to cavitation. The bench drying (BD) method is considered as a reference because embolism and xylem pressure are measured on large branches dehydrating in the air, in conditions similar to what happens in nature. Two other methods of embolism induction have been increasingly used. While the Cavitron (CA) uses centrifugal force to induce embolism, in the air injection (AI) method embolism is induced by forcing pressurized air to enter a stem segment. Recent studies have suggested that the AI and CA methods are inappropriate in long-vesselled species because they produce a very high-threshold xylem pressure for embolism (e.g., P50) compared with what is expected from (i) their ecophysiology in the field (native embolism, water potential and stomatal response to xylem pressure) and (ii) the P50 obtained with the BD method. However, other authors have argued that the CA and AI methods may be valid because they produce VCs similar to the BD method. In order to clarify this issue, we assessed VCs with the three above-mentioned methods on the long-vesselled Quercus ilex L. We showed that the BD VC yielded threshold xylem pressure for embolism consistent with in situ measurements of native embolism, minimal water potential and stomatal conductance. We therefore concluded that the BD method provides a reliable estimate of the VC for this species. The CA method produced a very high P50 (i.e., less negative) compared with the BD method, which is consistent with an artifact related to the vessel length. The VCs obtained with the AI method were highly variable, producing P50 ranging from -2 to -8.2 MPa. This wide variability was more related to differences in base diameter among samples than to differences in the length of samples. We concluded that this method is probably subject to an artifact linked to the distribution of vessel lengths within the sample. Overall, our results indicate that the CA and the AI should be used with extreme caution on long-vesselled species. Our results also highlight that several criteria may be helpful to assess the validity of a VC. © The Author 2014. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.
Kröber, W; Heklau, H; Bruelheide, H
2015-03-01
We explored potential of morphological and anatomical leaf traits for predicting ecophysiological key functions in subtropical trees. We asked whether the ecophysiological parameters stomatal conductance and xylem cavitation vulnerability could be predicted from microscopy leaf traits. We investigated 21 deciduous and 19 evergreen subtropical tree species, using individuals of the same age and from the same environment in the Biodiversity-Ecosystem Functioning experiment at Jiangxi (BEF-China). Information-theoretic linear model selection was used to identify the best combination of morphological and anatomical predictors for ecophysiological functions. Leaf anatomy and morphology strongly depended on leaf habit. Evergreen species tended to have thicker leaves, thicker spongy and palisade mesophyll, more palisade mesophyll layers and a thicker subepidermis. Over 50% of all evergreen species had leaves with multi-layered palisade parenchyma, while only one deciduous species (Koelreuteria bipinnata) had this. Interactions with leaf habit were also included in best multi-predictor models for stomatal conductance (gs ) and xylem cavitation vulnerability. In addition, maximum gs was positively related to log ratio of palisade to spongy mesophyll thickness. Vapour pressure deficit (vpd) for maximum gs increased with the log ratio of palisade to spongy mesophyll thickness in species having leaves with papillae. In contrast, maximum specific hydraulic conductivity and xylem pressure at which 50% loss of maximum specific xylem hydraulic conductivity occurred (Ψ50 ) were best predicted by leaf habit and density of spongy parenchyma. Evergreen species had lower Ψ50 values and lower maximum xylem hydraulic conductivities. As hydraulic leaf and wood characteristics were reflected in structural leaf traits, there is high potential for identifying further linkages between morphological and anatomical leaf traits and ecophysiological responses. © 2014 German Botanical Society and The Royal Botanical Society of the Netherlands.
Tree shoot bending generates hydraulic pressure pulses: a new long-distance signal?
Lopez, Rosana; Badel, Eric
2014-01-01
When tree stems are mechanically stimulated, a rapid long-distance signal is induced that slows down primary growth. An investigation was carried out to determine whether the signal might be borne by a mechanically induced pressure pulse in the xylem. Coupling xylem flow meters and pressure sensors with a mechanical testing device, the hydraulic effects of mechanical deformation of tree stem and branches were measured. Organs of several tree species were studied, including gymnosperms and angiosperms with different wood densities and anatomies. Bending had a negligible effect on xylem conductivity, even when deformations were sustained or were larger than would be encountered in nature. It was found that bending caused transient variation in the hydraulic pressure within the xylem of branch segments. This local transient increase in pressure in the xylem was rapidly propagated along the vascular system in planta to the upper and lower regions of the stem. It was shown that this hydraulic pulse originates from the apoplast. Water that was mobilized in the hydraulic pulses came from the saturated porous material of the conduits and their walls, suggesting that the poroelastic behaviour of xylem might be a key factor. Although likely to be a generic mechanical response, quantitative differences in the hydraulic pulse were found in different species, possibly related to differences in xylem anatomy. Importantly the hydraulic pulse was proportional to the strained volume, similar to known thigmomorphogenetic responses. It is hypothesized that the hydraulic pulse may be the signal that rapidly transmits mechanobiological information to leaves, roots, and apices. PMID:24558073
Tree shoot bending generates hydraulic pressure pulses: a new long-distance signal?
Lopez, Rosana; Badel, Eric; Peraudeau, Sebastien; Leblanc-Fournier, Nathalie; Beaujard, François; Julien, Jean-Louis; Cochard, Hervé; Moulia, Bruno
2014-05-01
When tree stems are mechanically stimulated, a rapid long-distance signal is induced that slows down primary growth. An investigation was carried out to determine whether the signal might be borne by a mechanically induced pressure pulse in the xylem. Coupling xylem flow meters and pressure sensors with a mechanical testing device, the hydraulic effects of mechanical deformation of tree stem and branches were measured. Organs of several tree species were studied, including gymnosperms and angiosperms with different wood densities and anatomies. Bending had a negligible effect on xylem conductivity, even when deformations were sustained or were larger than would be encountered in nature. It was found that bending caused transient variation in the hydraulic pressure within the xylem of branch segments. This local transient increase in pressure in the xylem was rapidly propagated along the vascular system in planta to the upper and lower regions of the stem. It was shown that this hydraulic pulse originates from the apoplast. Water that was mobilized in the hydraulic pulses came from the saturated porous material of the conduits and their walls, suggesting that the poroelastic behaviour of xylem might be a key factor. Although likely to be a generic mechanical response, quantitative differences in the hydraulic pulse were found in different species, possibly related to differences in xylem anatomy. Importantly the hydraulic pulse was proportional to the strained volume, similar to known thigmomorphogenetic responses. It is hypothesized that the hydraulic pulse may be the signal that rapidly transmits mechanobiological information to leaves, roots, and apices.
Xylem resistance to embolism: presenting a simple diagnostic test for the open vessel artefact.
Torres-Ruiz, José M; Cochard, Hervé; Choat, Brendan; Jansen, Steven; López, Rosana; Tomášková, Ivana; Padilla-Díaz, Carmen M; Badel, Eric; Burlett, Regis; King, Andrew; Lenoir, Nicolas; Martin-StPaul, Nicolas K; Delzon, Sylvain
2017-07-01
Xylem vulnerability to embolism represents an essential trait for the evaluation of the impact of hydraulics in plant function and ecology. The standard centrifuge technique is widely used for the construction of vulnerability curves, although its accuracy when applied to species with long vessels remains under debate. We developed a simple diagnostic test to determine whether the open-vessel artefact influences centrifuge estimates of embolism resistance. Xylem samples from three species with differing vessel lengths were exposed to less negative xylem pressures via centrifugation than the minimum pressure the sample had previously experienced. Additional calibration was obtained from non-invasive measurement of embolism on intact olive plants by X-ray microtomography. Results showed artefactual decreases in hydraulic conductance (k) for samples with open vessels when exposed to a less negative xylem pressure than the minimum pressure they had previously experienced. X-Ray microtomography indicated that most of the embolism formation in olive occurs at xylem pressures below -4.0 MPa, reaching 50% loss of hydraulic conductivity at -5.3 MPa. The artefactual reductions in k induced by centrifugation underestimate embolism resistance data of species with long vessels. A simple test is suggested to avoid this open vessel artefact and to ensure the reliability of this technique in future studies. © 2017 The Authors. New Phytologist © 2017 New Phytologist Trust.
Espino, Susana; Schenk, H. Jochen
2011-01-01
The maximum specific hydraulic conductivity (kmax) of a plant sample is a measure of the ability of a plants’ vascular system to transport water and dissolved nutrients under optimum conditions. Precise measurements of kmax are needed in comparative studies of hydraulic conductivity, as well as for measuring the formation and repair of xylem embolisms. Unstable measurements of kmax are a common problem when measuring woody plant samples and it is commonly observed that kmax declines from initially high values, especially when positive water pressure is used to flush out embolisms. This study was designed to test five hypotheses that could potentially explain declines in kmax under positive pressure: (i) non-steady-state flow; (ii) swelling of pectin hydrogels in inter-vessel pit membranes; (iii) nucleation and coalescence of bubbles at constrictions in the xylem; (iv) physiological wounding responses; and (v) passive wounding responses, such as clogging of the xylem by debris. Prehydrated woody stems from Laurus nobilis (Lauraceae) and Encelia farinosa (Asteraceae) collected from plants grown in the Fullerton Arboretum in Southern California, were used to test these hypotheses using a xylem embolism meter (XYL'EM). Treatments included simultaneous measurements of stem inflow and outflow, enzyme inhibitors, stem-debarking, low water temperatures, different water degassing techniques, and varied concentrations of calcium, potassium, magnesium, and copper salts in aqueous measurement solutions. Stable measurements of kmax were observed at concentrations of calcium, potassium, and magnesium salts high enough to suppress bubble coalescence, as well as with deionized water that was degassed using a membrane contactor under strong vacuum. Bubble formation and coalescence under positive pressure in the xylem therefore appear to be the main cause for declining kmax values. Our findings suggest that degassing of water is essential for achieving stable and precise measurements of kmax through woody plant samples. For complete rehydration of woody samples, incubation in water under vacuum for 24 h is suggested as a reliable technique that avoids bubble problems associated with flushing under high positive pressure. PMID:21147811
Do xylem fibers affect vessel cavitation resistance?
Jacobsen, Anna L; Ewers, Frank W; Pratt, R Brandon; Paddock, William A; Davis, Stephen D
2005-09-01
Possible mechanical and hydraulic costs to increased cavitation resistance were examined among six co-occurring species of chaparral shrubs in southern California. We measured cavitation resistance (xylem pressure at 50% loss of hydraulic conductivity), seasonal low pressure potential (P(min)), xylem conductive efficiency (specific conductivity), mechanical strength of stems (modulus of elasticity and modulus of rupture), and xylem density. At the cellular level, we measured vessel and fiber wall thickness and lumen diameter, transverse fiber wall and total lumen area, and estimated vessel implosion resistance using (t/b)(h)(2), where t is the thickness of adjoining vessel walls and b is the vessel lumen diameter. Increased cavitation resistance was correlated with increased mechanical strength (r(2) = 0.74 and 0.76 for modulus of elasticity and modulus of rupture, respectively), xylem density (r(2) = 0.88), and P(min) (r(2) = 0.96). In contrast, cavitation resistance and P(min) were not correlated with decreased specific conductivity, suggesting no tradeoff between these traits. At the cellular level, increased cavitation resistance was correlated with increased (t/b)(h)(2) (r(2) = 0.95), increased transverse fiber wall area (r(2) = 0.89), and decreased fiber lumen area (r(2) = 0.76). To our knowledge, the correlation between cavitation resistance and fiber wall area has not been shown previously and suggests a mechanical role for fibers in cavitation resistance. Fiber efficacy in prevention of vessel implosion, defined as inward bending or collapse of vessels, is discussed.
Zimmermann, D; Westhoff, M; Zimmermann, G; Gessner, P; Gessner, A; Wegner, L H; Rokitta, M; Ache, P; Schneider, H; Vásquez, J A; Kruck, W; Shirley, S; Jakob, P; Hedrich, R; Bentrup, F-W; Bamberg, E; Zimmermann, U
2007-01-01
The water supply to leaves of 25 to 60 m tall trees (including high-salinity-tolerant ones) was studied. The filling status of the xylem vessels was determined by xylem sap extraction (using jet-discharge, gravity-discharge, and centrifugation) and by (1)H nuclear magnetic resonance imaging of wood pieces. Simultaneously, pressure bomb experiments were performed along the entire trunk of the trees up to a height of 57 m. Clear-cut evidence was found that the balancing pressure (P(b)) values of leafy twigs were dictated by the ambient relative humidity rather than by height. Refilling of xylem vessels of apical leaves (branches) obviously mainly occurred via moisture uptake from the atmosphere. These findings could be traced back to the hydration and rehydration of mucilage layers on the leaf surfaces and/or of epistomatal mucilage plugs. Xylem vessels also contained mucilage. Mucilage formation was apparently enforced by water stress. The observed mucilage-based foliar water uptake and humidity dependency of the P(b) values are at variance with the cohesion-tension theory and with the hypothesis that P(b) measurements yield information about the relationships between xylem pressure gradients and height.
Post-fire Tree Mortality: Heating Increases Vulnerability to Cavitation in Longleaf Pine Branches
NASA Astrophysics Data System (ADS)
Lodge, A.; Kavanagh, K.; Dickinson, M. B.
2016-12-01
Tree mortality following wild and prescribed fires is of interest to both researchers and land managers. While some models exist that can predict mortality following fires, process-based models that incorporate physiological mechanisms of mortality are still being developed and improved. Delayed post-fire tree mortality has recently received increased attention, in part due to an increased use of prescribed fire as a restoration and management tool. One hypothesized mechanism of delayed mortality in trees is disruption of water transport in xylem due to exposure to the heat plume of a fire. This heat plume rapidly increases the vapor pressure deficit in the tree canopy, quickly increasing the tension on the water held in the xylem and leaves, potentially leading to cavitation. Cavitated xylem conduits can no longer transport water, eventually leading to tree death. We conducted a laboratory experiment examining whether heating stems increases their vulnerability to cavitation. We placed longleaf pine (Pinus palustris) branches in a water bath at sub-lethal temperatures (<60°C) and applied pressure in a cavitation chamber to simulate a range of xylem tension levels that may occur during fire. Percent loss of conductivity was measured following cavitation induced by various levels of applied pressure. When we compared the resulting vulnerability curves of heated branches to those of branches pressurized at room temperature, we observed increased vulnerability to cavitation in the heated samples especially at lower pressures. P50, or the pressure at which 50% of conductivity has been lost, decreased by 18% on branches heated to approximately 54°C. This suggests that stems heated during fires may be more vulnerable to cavitation, and provides some support for hydraulic disruption as a mechanism for post-fire tree mortality. Continued advancement in understanding of the mechanisms leading to delayed mortality will improve models predicting tree mortality.
Xylem Surfactants Introduce a New Element to the Cohesion-Tension Theory1[OPEN
Espino, Susana; Nima, Neda; Do, Aissa Y.T.; Michaud, Joseph M.; Papahadjopoulos-Sternberg, Brigitte; Yang, Jinlong; Steppe, Kathy
2017-01-01
Vascular plants transport water under negative pressure without constantly creating gas bubbles that would disable their hydraulic systems. Attempts to replicate this feat in artificial systems almost invariably result in bubble formation, except under highly controlled conditions with pure water and only hydrophilic surfaces present. In theory, conditions in the xylem should favor bubble nucleation even more: there are millions of conduits with at least some hydrophobic surfaces, and xylem sap is saturated or sometimes supersaturated with atmospheric gas and may contain surface-active molecules that can lower surface tension. So how do plants transport water under negative pressure? Here, we show that angiosperm xylem contains abundant hydrophobic surfaces as well as insoluble lipid surfactants, including phospholipids, and proteins, a composition similar to pulmonary surfactants. Lipid surfactants were found in xylem sap and as nanoparticles under transmission electron microscopy in pores of intervessel pit membranes and deposited on vessel wall surfaces. Nanoparticles observed in xylem sap via nanoparticle-tracking analysis included surfactant-coated nanobubbles when examined by freeze-fracture electron microscopy. Based on their fracture behavior, this technique is able to distinguish between dense-core particles, liquid-filled, bilayer-coated vesicles/liposomes, and gas-filled bubbles. Xylem surfactants showed strong surface activity that reduces surface tension to low values when concentrated as they are in pit membrane pores. We hypothesize that xylem surfactants support water transport under negative pressure as explained by the cohesion-tension theory by coating hydrophobic surfaces and nanobubbles, thereby keeping the latter below the critical size at which bubbles would expand to form embolisms. PMID:27927981
Xylem Surfactants Introduce a New Element to the Cohesion-Tension Theory.
Schenk, H Jochen; Espino, Susana; Romo, David M; Nima, Neda; Do, Aissa Y T; Michaud, Joseph M; Papahadjopoulos-Sternberg, Brigitte; Yang, Jinlong; Zuo, Yi Y; Steppe, Kathy; Jansen, Steven
2017-02-01
Vascular plants transport water under negative pressure without constantly creating gas bubbles that would disable their hydraulic systems. Attempts to replicate this feat in artificial systems almost invariably result in bubble formation, except under highly controlled conditions with pure water and only hydrophilic surfaces present. In theory, conditions in the xylem should favor bubble nucleation even more: there are millions of conduits with at least some hydrophobic surfaces, and xylem sap is saturated or sometimes supersaturated with atmospheric gas and may contain surface-active molecules that can lower surface tension. So how do plants transport water under negative pressure? Here, we show that angiosperm xylem contains abundant hydrophobic surfaces as well as insoluble lipid surfactants, including phospholipids, and proteins, a composition similar to pulmonary surfactants. Lipid surfactants were found in xylem sap and as nanoparticles under transmission electron microscopy in pores of intervessel pit membranes and deposited on vessel wall surfaces. Nanoparticles observed in xylem sap via nanoparticle-tracking analysis included surfactant-coated nanobubbles when examined by freeze-fracture electron microscopy. Based on their fracture behavior, this technique is able to distinguish between dense-core particles, liquid-filled, bilayer-coated vesicles/liposomes, and gas-filled bubbles. Xylem surfactants showed strong surface activity that reduces surface tension to low values when concentrated as they are in pit membrane pores. We hypothesize that xylem surfactants support water transport under negative pressure as explained by the cohesion-tension theory by coating hydrophobic surfaces and nanobubbles, thereby keeping the latter below the critical size at which bubbles would expand to form embolisms. © 2017 American Society of Plant Biologists. All Rights Reserved.
Knipfer, Thorsten; Fei, Jiong; Gambetta, Gregory A; McElrone, Andrew J; Shackel, Kenneth A; Matthews, Mark A
2015-08-01
Xylem flow of water into fruits declines during fruit development, and the literature indicates a corresponding increase in hydraulic resistance in the pedicel. However, it is unknown how pedicel hydraulics change developmentally in relation to xylem anatomy and function. In this study on grape (Vitis vinifera), we determined pedicel hydraulic conductivity (kh) from pressure-flow relationships using hydrostatic and osmotic forces and investigated xylem anatomy and function using fluorescent light microscopy and x-ray computed microtomography. Hydrostatic kh (xylem pathway) was consistently 4 orders of magnitude greater than osmotic kh (intracellular pathway), but both declined before veraison by approximately 40% and substantially over fruit development. Hydrostatic kh declined most gradually for low (less than 0.08 MPa) pressures and for water inflow and outflow conditions. Specific kh (per xylem area) decreased in a similar fashion to kh despite substantial increases in xylem area. X-ray computed microtomography images provided direct evidence that losses in pedicel kh were associated with blockages in vessel elements, whereas air embolisms were negligible. However, vessel elements were interconnected and some remained continuous postveraison, suggesting that across the grape pedicel, a xylem pathway of reduced kh remains functional late into berry ripening. © 2015 American Society of Plant Biologists. All Rights Reserved.
Fei, Jiong; McElrone, Andrew J.; Shackel, Kenneth A.; Matthews, Mark A.
2015-01-01
Xylem flow of water into fruits declines during fruit development, and the literature indicates a corresponding increase in hydraulic resistance in the pedicel. However, it is unknown how pedicel hydraulics change developmentally in relation to xylem anatomy and function. In this study on grape (Vitis vinifera), we determined pedicel hydraulic conductivity (kh) from pressure-flow relationships using hydrostatic and osmotic forces and investigated xylem anatomy and function using fluorescent light microscopy and x-ray computed microtomography. Hydrostatic kh (xylem pathway) was consistently 4 orders of magnitude greater than osmotic kh (intracellular pathway), but both declined before veraison by approximately 40% and substantially over fruit development. Hydrostatic kh declined most gradually for low (less than 0.08 MPa) pressures and for water inflow and outflow conditions. Specific kh (per xylem area) decreased in a similar fashion to kh despite substantial increases in xylem area. X-ray computed microtomography images provided direct evidence that losses in pedicel kh were associated with blockages in vessel elements, whereas air embolisms were negligible. However, vessel elements were interconnected and some remained continuous postveraison, suggesting that across the grape pedicel, a xylem pathway of reduced kh remains functional late into berry ripening. PMID:26077763
Davis, Stephen D; Ewers, Frank W; Sperry, John S; Portwood, Kimberly A; Crocker, Michelle C; Adams, Gerard C
2002-05-01
Progressive diebacks of outer canopy branchlets of Ceanothus crassifolius were repeatedly observed after rainless periods up to 9 mo in duration in the Santa Monica Mountains of southern California. Mean xylem pressures of branchlets near the end of drought were as low as -11.2 MPa (N = 22) with a mean of about 60 dead branchlets per shrub. Inoculation (N = 15) with three species of fungi previously isolated from the same population of C. crassifolius did not promote dieback, suggesting that the observed decline was not fungal induced, as had been proposed. Further, at least 50% of healthy-appearing twigs, without symptoms of dieback, contained isolatible endophytic fungi. We used a centrifugal force method to determine the range of xylem pressure causing cavitation (vulnerability curves) for branchlets (N = 12) and roots (N = 16). We combined vulnerability curves with soil texture data (N = 6) into a water transport model that estimated the critical values (P(Lcrit)) of leaf xylem pressure associated with the loss of water from soil to foliage. Maximum P(Lcrit) was between -10 and -11 MPa and within the range of minimum measured xylem pressures of branchlets during drought and dieback. Branchlet dieback correlated with seasonal declines in xylem pressure in concert with declining safety margins from hydraulic failure. Symptoms of dieback were duplicated in the field by partially severing stem xylem that normally supplied branchlets with water. Taken together, these results indicate that loss of hydraulic conductance to foliage was the probable cause of the observed dieback in C. crassifolius. Partial dieback of peripheral branchlets, and its attendant reduction in evaporative surface area, may be a last-resort mechanism for whole-plant water conservation and drought survival in this species.
Wegner, Lars H; Stefano, Giovanni; Shabala, Lana; Rossi, Marika; Mancuso, Stefano; Shabala, Sergey
2011-05-01
Early events in NaCl-induced root ion and water transport were investigated in maize (Zea mays L) roots using a range of microelectrode and imaging techniques. Addition of 100 mm NaCl to the bath resulted in an exponential drop in root xylem pressure, rapid depolarization of trans-root potential and a transient drop in xylem K(+) activity (A(K+) ) within ∼1 min after stress onset. At this time, no detectable amounts of Na(+) were released into the xylem vessels. The observed drop in A(K+) was unexpected, given the fact that application of the physiologically relevant concentrations of Na(+) to isolated stele has caused rapid plasma membrane depolarization and a subsequent K(+) efflux from the stelar tissues. This controversy was explained by the difference in kinetics of NaCl-induced depolarization between cortical and stelar cells. As root cortical cells are first to be depolarized and lose K(+) to the environment, this is associated with some K(+) shift from the stelar symplast to the cortex, resulting in K(+) being transiently removed from the xylem. Once Na(+) is loaded into the xylem (between 1 and 5 min of root exposure to NaCl), stelar cells become more depolarized, and a gradual recovery in A(K+) occurs. © 2011 Blackwell Publishing Ltd.
Arabidopsis thaliana as a model species for xylem hydraulics: does size matter?
Tixier, Aude; Cochard, Hervé; Badel, Eric; Dusotoit-Coucaud, Anaïs; Jansen, Steven; Herbette, Stéphane
2013-01-01
While Arabidopsis thaliana has been proposed as a model species for wood development, the potential of this tiny herb for studying xylem hydraulics remains unexplored and anticipated by scepticism. Inflorescence stems of A. thaliana were used to measure hydraulic conductivity and cavitation resistance, whereas light and electron microscopy allowed observations of vessels. In wild-type plants, measured and theoretical conductivity showed a significant correlation (R 2 = 0.80, P < 0.01). Moreover, scaling of vessel dimensions and intervessel pit structure of A. thaliana were consistent with structure–function relationships of woody plants. The reliability and resolution of the hydraulic methods applied to measure vulnerability to cavitation were addressed by comparing plants grown under different photoperiods or different mutant lines. Sigmoid vulnerability curves of A. thaliana indicated a pressure corresponding to 50% loss of hydraulic conductance (P 50) between –3 and –2.5MPa for short-day and long-day plants, respectively. Polygalacturonase mutants showed a higher P 50 value (–2.25MPa), suggesting a role for pectins in vulnerability to cavitation. The application of A. thaliana as a model species for xylem hydraulics provides exciting possibilities for (1) exploring the molecular basis of xylem anatomical features and (2) understanding genetic mechanisms behind xylem functional traits such as cavitation resistance. Compared to perennial woody species, however, the lesser amount of xylem in A. thaliana has its limitations. PMID:23547109
Arabidopsis thaliana as a model species for xylem hydraulics: does size matter?
Tixier, Aude; Cochard, Hervé; Badel, Eric; Dusotoit-Coucaud, Anaïs; Jansen, Steven; Herbette, Stéphane
2013-05-01
While Arabidopsis thaliana has been proposed as a model species for wood development, the potential of this tiny herb for studying xylem hydraulics remains unexplored and anticipated by scepticism. Inflorescence stems of A. thaliana were used to measure hydraulic conductivity and cavitation resistance, whereas light and electron microscopy allowed observations of vessels. In wild-type plants, measured and theoretical conductivity showed a significant correlation (R (2) = 0.80, P < 0.01). Moreover, scaling of vessel dimensions and intervessel pit structure of A. thaliana were consistent with structure-function relationships of woody plants. The reliability and resolution of the hydraulic methods applied to measure vulnerability to cavitation were addressed by comparing plants grown under different photoperiods or different mutant lines. Sigmoid vulnerability curves of A. thaliana indicated a pressure corresponding to 50% loss of hydraulic conductance (P 50) between -3 and -2.5MPa for short-day and long-day plants, respectively. Polygalacturonase mutants showed a higher P 50 value (-2.25MPa), suggesting a role for pectins in vulnerability to cavitation. The application of A. thaliana as a model species for xylem hydraulics provides exciting possibilities for (1) exploring the molecular basis of xylem anatomical features and (2) understanding genetic mechanisms behind xylem functional traits such as cavitation resistance. Compared to perennial woody species, however, the lesser amount of xylem in A. thaliana has its limitations.
Root pressure and beyond: energetically uphill water transport into xylem vessels?
Wegner, Lars H
2014-02-01
The thermodynamics of root pressure remains an enigma up to the present day. Water is transported radially into xylem vessels, under some conditions even when the xylem sap is more dilute than the ambient medium (soil solution). It is suggested here that water secretion across the plasma membrane of xylem parenchyma cells is driven by a co-transport of water and solutes as previously shown for mammalian epithelia (Zeuthen T. 2010. Water-transporting proteins. Journal of Membrane Biology 234, 57-73.). This process could drive volume flow 'energetically uphill', against the free energy gradient of water. According to the model, solutes released by xylem parenchyma cells are subsequently retrieved from the sap at the expense of metabolic energy to maintain the concentration gradient that drives the water secretion. Transporters of the CCC type known to mediate water secretion in mammalian cells have also been found in Arabidopsis and in rice. The mechanism proposed here for root pressure could also explain refilling of embolized vessels. Moreover, it could contribute to long-distance water transport in trees when the cohesion-tension mechanism of water ascent fails. This is discussed with respect to the old and the more recent literature on these subjects.
Dissolved atmospheric gas in xylem sap measured with membrane inlet mass spectrometry.
Schenk, H Jochen; Espino, Susana; Visser, Ate; Esser, Bradley K
2016-04-01
A new method is described for measuring dissolved gas concentrations in small volumes of xylem sap using membrane inlet mass spectrometry. The technique can be used to determine concentrations of atmospheric gases, such as argon, as reported here, or for any dissolved gases and their isotopes for a variety of applications, such as rapid detection of trace gases from groundwater only hours after they were taken up by trees and rooting depth estimation. Atmospheric gas content in xylem sap directly affects the conditions and mechanisms that allow for gas removal from xylem embolisms, because gas can dissolve into saturated or supersaturated sap only under gas pressure that is above atmospheric pressure. The method was tested for red trumpet vine, Distictis buccinatoria (Bignoniaceae), by measuring atmospheric gas concentrations in sap collected at times of minimum and maximum daily temperature and during temperature increase and decline. Mean argon concentration in xylem sap did not differ significantly from saturation levels for the temperature and pressure conditions at any time of collection, but more than 40% of all samples were supersaturated, especially during the warm parts of day. There was no significant diurnal pattern, due to high variability between samples. © 2015 John Wiley & Sons Ltd.
Nikinmaa, Eero; Sievänen, Risto; Hölttä, Teemu
2014-09-01
Tree models simulate productivity using general gas exchange responses and structural relationships, but they rarely check whether leaf gas exchange and resulting water and assimilate transport and driving pressure gradients remain within acceptable physical boundaries. This study presents an implementation of the cohesion-tension theory of xylem transport and the Münch hypothesis of phloem transport in a realistic 3-D tree structure and assesses the gas exchange and transport dynamics. A mechanistic model of xylem and phloem transport was used, together with a tested leaf assimilation and transpiration model in a realistic tree architecture to simulate leaf gas exchange and water and carbohydrate transport within an 8-year-old Scots pine tree. The model solved the dynamics of the amounts of water and sucrose solute in the xylem, cambium and phloem using a fine-grained mesh with a system of coupled ordinary differential equations. The simulations predicted the observed patterns of pressure gradients and sugar concentration. Diurnal variation of environmental conditions influenced tree-level gradients in turgor pressure and sugar concentration, which are important drivers of carbon allocation. The results and between-shoot variation were sensitive to structural and functional parameters such as tree-level scaling of conduit size and phloem unloading. Linking whole-tree-level water and assimilate transport, gas exchange and sink activity opens a new avenue for plant studies, as features that are difficult to measure can be studied dynamically with the model. Tree-level responses to local and external conditions can be tested, thus making the approach described here a good test-bench for studies of whole-tree physiology.
Tobin, M F; Pratt, R B; Jacobsen, A L; De Guzman, M E
2013-05-01
Vulnerability to cavitation curves describe the decrease in xylem hydraulic conductivity as xylem pressure declines. Several techniques for constructing vulnerability curves use centrifugal force to induce negative xylem pressure in stem or root segments. Centrifuge vulnerability curves constructed for long-vesselled species have been hypothesised to overestimate xylem vulnerability to cavitation due to increased vulnerability of vessels cut open at stem ends that extend to the middle or entirely through segments. We tested two key predictions of this hypothesis: (i) centrifugation induces greater embolism than dehydration in long-vesselled species, and (ii) the proportion of open vessels changes centrifuge vulnerability curves. Centrifuge and dehydration vulnerability curves were compared for a long- and short-vesselled species. The effect of open vessels was tested in four species by comparing centrifuge vulnerability curves for stems of two lengths. Centrifuge and dehydration vulnerability curves agreed well for the long- and short-vesselled species. Centrifuge vulnerability curves constructed using two stem lengths were similar. Also, the distribution of embolism along the length of centrifuged stems matched the theoretical pressure profile induced by centrifugation. We conclude that vulnerability to cavitation can be accurately characterised with vulnerability curves constructed using a centrifuge technique, even in long-vesselled species. © 2012 German Botanical Society and The Royal Botanical Society of the Netherlands.
Conservation of element concentration in xylem sap of red spruce
Kevin T. Smith; Walter C. Shortle
2001-01-01
We investigated the chemistry of xylem sap as a marker of red spruce metabolism and soil chemistry at three locations in northern New England. A Scholander pressure chamber was used to extract xylem sap from roots and branches cut from mature trees in early June and September. Root sap contained significantly greater concentrations of K, Ca, Mg, Mn, and A1 than branch...
Garvin, Naho; Doucette, William J; White, Jason C
2015-07-01
A pressure chamber method was used to examine differences in the root to shoot transfer and xylem sap solubility of caffeine (log Kow=-0.07), triclocarban (log Kow=3.5-4.2) and endosulfan (log Kow=3.8-4.8) for zucchini (cucurbita pepo ssp pepo), squash (cucurbita pepo ssp ovifera), and soybean (glycine max L.). Transpiration stream concentration factors (TSCF) for caffeine (TSCF=0.8) were statistically equivalent for all plant species. However, for the more hydrophobic endosulfan and triclocarban, the TSCF values for zucchini (TSCF=0.6 and 0.4, respectively) were 3 and 10 times greater than the soybean and squash (TSCF=0.2 and 0.05, respectively). The difference in TSCF values was examined by comparing the measured solubilities of caffeine, endosulfan and triclocarban in deionized water to those in soybean and zucchini xylem saps using a modified shake flask method. The measured solubility of organic contaminants in xylem sap has not previously been reported. Caffeine solubilities in the xylem saps of soybean and zucchini were statistically equal to deionized water (21500mgL(-1)) while endosulfan and triclocarban solubilities in the zucchini xylem sap were significantly greater (0.43 and 0.21mgL(-1), respectively) than that of the soybean xylem sap (0.31 and 0.11mgL(-1), respectively) and deionized water (0.34 and 0.11mgL(-1), respectively). This suggests that the enhanced root to shoot transfer of hydrophobic organics reported for zucchini is partly due to increased solubility in the xylem sap. Further xylem sap characterization is needed to determine the mechanism of solubility enhancement. Copyright © 2014 Elsevier Ltd. All rights reserved.
Riparian Plant Water Relations Along the North Fork Kings River, California
Janet L. Nachlinger; Stanley D. Smith; Roland J. Risser
1989-01-01
Plant water relations of five obligate riparian species were studied along California's North Fork Kings River. Diurnal stomatal conductance, transpiration, and xylem pressure potentials were measured throughout the 1986 growing season and in mid-season in 1987. Patterns were similar for all species although absolute values varied considerably. Maximum stomatal...
Brodersen, Craig R; Rico, Christopher; Guenni, Orlando; Pittermann, Jarmila
2016-02-01
Xylem network structure and function have been characterized for many woody plants, but less is known about fern xylem, particularly in species endemic to climates where water is a limiting resource for months at a time. We characterized seasonal variability in soil moisture and frond water status in a common perennial fern in the redwood understory of a costal California, and then investigated the consequences of drought-induced embolism on vascular function. Seasonal variability in air temperature and soil water content was minimal, and frond water potential declined slowly over the observational period. Our data show that Polystichum munitum was protected from significant drought-induced hydraulic dysfunction during this growing season because of a combination of cavitation resistant conduits (Air-seeding threshold (ASP) = -1.53 MPa; xylem pressure inducing 50% loss of hydraulic conductivity (P50 ) = -3.02 MPa) and a soil with low moisture variability. High resolution micro-computed tomography (MicroCT) imaging revealed patterns of embolism formation in vivo for the first time in ferns providing insight into the functional status of the xylem network under drought conditions. Together with stomatal conductance measurements, these data suggest that P. munitum is adapted to tolerate drier conditions than what was observed during the growing season. © 2015 John Wiley & Sons Ltd.
Sugar demand of ripening grape berries leads to recycling of surplus phloem water via the xylem.
Keller, Markus; Zhang, Yun; Shrestha, Pradeep M; Biondi, Marco; Bondada, Bhaskar R
2015-06-01
We tested the common assumption that fleshy fruits become dependent on phloem water supply because xylem inflow declines at the onset of ripening. Using two distinct grape genotypes exposed to drought stress, we found that a sink-driven rise in phloem inflow at the beginning of ripening was sufficient to reverse drought-induced berry shrinkage. Rewatering accelerated berry growth and sugar accumulation concurrently with leaf photosynthetic recovery. Interrupting phloem flow through the peduncle prevented the increase in berry growth after rewatering, but interrupting xylem flow did not. Nevertheless, xylem flow in ripening berries, but not berry size, remained responsive to root or shoot pressurization. A mass balance analysis on ripening berries sampled in the field suggested that phloem water inflow may exceed growth and transpiration water demands. Collecting apoplastic sap from ripening berries showed that osmotic pressure increased at distinct rates in berry vacuoles and apoplast. Our results indicate that the decrease in xylem inflow at the onset of ripening may be a consequence of the sink-driven increase in phloem inflow. We propose a conceptual model in which surplus phloem water bypasses the fruit cells and partly evaporates from the berry surface and partly moves apoplastically to the xylem for outflow. © 2014 John Wiley & Sons Ltd.
Couvreur, Valentin; Ledder, Glenn; Manzoni, Stefano; Way, Danielle A; Muller, Erik B; Russo, Sabrina E
2018-05-08
Trees grow by vertically extending their stems, so accurate stem hydraulic models are fundamental to understanding the hydraulic challenges faced by tall trees. Using a literature survey, we showed that many tree species exhibit continuous vertical variation in hydraulic traits. To examine the effects of this variation on hydraulic function, we developed a spatially-explicit, analytical water transport model for stems. Our model allows Huber ratio, stem-saturated conductivity, pressure at 50% loss of conductivity, leaf area, and transpiration rate to vary continuously along the hydraulic path. Predictions from our model differ from a matric flux potential model parameterized with uniform traits. Analyses show that cavitation is a whole-stem emergent property resulting from nonlinear pressure-conductivity feedbacks that, with gravity, cause impaired water transport to accumulate along the path. Because of the compounding effects of vertical trait variation on hydraulic function, growing proportionally more sapwood and building tapered xylem with height, as well as reducing xylem vulnerability only at branch tips while maintaining transport capacity at the stem base, can compensate for these effects. We therefore conclude that the adaptive significance of vertical variation in stem hydraulic traits is to allow trees to grow tall and tolerate operating near their hydraulic limits. This article is protected by copyright. All rights reserved.
Meitern, Annika; Õunapuu-Pikas, Eele; Sellin, Arne
2017-06-01
Physiological processes taking place in plants are subject to diverse circadian patterns but some of them are poorly documented in natural conditions. The daily dynamics of physico-chemical properties of xylem sap and their covariation with tree hydraulic traits were investigated in hybrid aspen (Populus tremula L.×P. tremuloides Michx) in field conditions in order to clarify which environmental drivers govern the daily variation in these parameters. K + concentration ([K + ]), electrical conductivity (σ sap ), osmolality (Osm) and pH of the xylem sap, as well as branch hydraulic traits, were measured in the field over 24-h cycles. All studied xylem sap properties and hydraulic characteristics including whole-branch (K wb ), leaf blade (K lb ) and petiole hydraulic conductances (K P ) showed clear daily dynamics. Air temperature (T A ) and photosynthetic photon flux density (PPFD), but also water vapour pressure deficit (VPD) and relative humidity (RH), had significant impacts on K wb K lb , K P , [K + ] and σ sap . Osm varied only with light intensity, while K B varied depending on atmospheric evaporative demand expressed as T A , VPD or RH. Xylem sap pH depended inversely on soil water potential (Ψ S ) and during daylight also on VPD. Although soil water content was close to saturation during the study period, Ψ S influenced also [K + ] and σ sap . The present study presents evidence of coupling between circadian patterns of xylem sap properties and plant hydraulic conductance providing adequate water supply to foliage under environmental conditions characterised by diurnal variation. Copyright © 2017 Elsevier GmbH. All rights reserved.
Ooeda, Hiroki; Terashima, Ichiro; Taneda, Haruhiko
2017-02-01
Two hypotheses have been proposed to explain the mechanism preventing the refilling vessel water from being drained to the neighboring functional vessels under negative pressure. The pit membrane osmosis hypothesis proposes that the xylem parenchyma cells release polysaccharides that are impermeable to the intervessel pit membranes into the refilling vessel; this osmotically counteracts the negative pressure, thereby allowing the vessel to refill. The pit valve hypothesis proposes that gas trapped within intervessel bordered pits isolates the refilling vessel water from the surrounding functional vessels. Here, using the single-vessel method, we assessed these hypotheses in shoots of mulberry (Morus australis Poir.). First, we confirmed the occurrence of xylem refilling under negative pressure in the potted mulberry saplings. To examine the pit membrane osmosis hypothesis, we estimated the semi-permeability of pit membranes for molecules of various sizes and found that the pit membranes were not semi-permeable to polyethylene glycol of molecular mass <20,000. For the pit valve hypothesis, we formed pit valves in the intervessel pits in the short stem segments and measured the maximum liquid pressure up to which gases in bordered pits were retained. The threshold pressure ranged from 0.025 to 0.10 MPa. These values matched the theoretical value calculated from the geometry of the pit chamber (0.0692-0.101 MPa). Our results suggest that gas in the pits is retained by surface tension, even under substantial positive pressure to resolve gases in the refilling vessel, whereas the molecule size required for the pit membrane osmosis mechanism in mulberry would be unrealistically large. © The Author 2016. Published by Oxford University Press on behalf of Japanese Society of Plant Physiologists. All rights reserved. For permissions, please email: journals.permissions@oup.com.
Physiological changes in red spruce seedlings during a simulated winter thaw
P.G. Schaberg; J.B. Shane; G.J. Hawley; G.R. Strimbeck; D.H. DeHayes; P.F. Cali; J.R. Donnelly
1996-01-01
We evaluated net photosynthesis, respiration, leaf conductance, xylem pressure potential (XPP) and cold hardiness in red spruce (Picea rubens Sarg.) seedlings exposed to either a continuous thaw (CT) or a daytime thaw with freezing nights (FN) for 8 days during mid-winter. Physiological differences between CT and FN seedlings were evident for all...
P.F. Murakami; P.G. Schaberg; J.B. Shane
2008-01-01
To better understand the effects of sugar accumulation on red color development of foliage during autumn, we compared carbohydrate concentration, anthocyanin expression and xylem pressure potential of foliage on girdled versus non-girdled (control) branches of 12 mature, open-grown sugar maple (Acer saccharum Marsh.) trees. Half of the study trees...
Effect of vegetative competition on the moisture and nutrient status of loblolly pine
G.A. Carter; J.H. Miller; D.E. Davis; R.M. Patterson
1984-01-01
A field study examined the effects of competing vegetation on the moisture and nutrient status of 5-year-old loblolly pines (Pinus taeda L.). Similar experiments were conducted on a Piedmont site and a Coastal Plain site using individual pines as experimental units. Predawn measurements of xylem pressure potential were made using detached needle...
Yang, Dongmei; Pan, Shaoan; Tyree, Melvin T
2016-08-01
Pressure-volume (PV) curve analysis is the most common and accurate way of estimating all components of the water relationships in leaves (water potential isotherms) as summarized in the Höfler diagram. PV curve analysis yields values of osmotic pressure, turgor pressure, and elastic modulus of cell walls as a function of relative water content. It allows the computation of symplasmic/apoplastic water content partitioning. For about 20 years, cavitation in xylem has been postulated as a possible source of error when estimating the above parameters, but, to the best of the authors' knowledge, no one has ever previously quantified its influence. Results in this paper provide independent estimates of osmotic pressure by PV curve analysis and by thermocouple psychrometer measurement. An anatomical evaluation was also used for the first time to compare apoplastic water fraction estimates from PV analysis with anatomical values. Conclusions include: (i) PV curve values of osmotic pressure are underestimated prior to correcting osmotic pressure for water loss by cavitation in Metasequoia glyptostroboides; (ii) psychrometer estimates of osmotic pressure obtained in tissues killed by freezing or heating agreed with PV values before correction for apoplastic water dilution; (iii) after correction for dilution effects, a solute concentration enhancement (0.27MPa or 0.11 osmolal) was revealed. The possible sources of solute enhancement were starch hydrolysis and release of ions from the Donnan free space of needle cell walls. © The Author 2016. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.
Fichot, Régis; Chamaillard, Sylvain; Depardieu, Claire; Le Thiec, Didier; Cochard, Hervé; Barigah, Têtè S; Brignolas, Franck
2011-03-01
Tests were carried out to determine whether variations in the hydraulic architecture of eight Populus deltoides×Populus nigra genotypes could be related to variations in leaf function and growth performance. Measurements were performed in a coppice plantation on 1-year-old shoots under optimal irrigation. Hydraulic architecture was characterized through estimates of hydraulic efficiency (the ratio of conducting sapwood area to leaf area, A(X):A(L); leaf- and xylem-specific hydraulic conductance of defoliated shoots, k(SL) and k(SS), respectively; apparent whole-plant leaf-specific hydraulic conductance, k(plant)) and xylem safety (water potential inducing 50% loss in hydraulic conductance). The eight genotypes spanned a significant range of k(SL) from 2.63 kg s(-1) m(-2) MPa(-1) to 4.18 kg s(-1) m(-2) MPa(-1), variations being mostly driven by k(SS) rather than A(X):A(L). There was a strong trade-off between hydraulic efficiency and xylem safety. Values of k(SL) correlated positively with k(plant), indicating that high-pressure flowmeter (HPFM) measurements of stem hydraulic efficiency accurately reflected whole-plant water transport efficiency of field-grown plants at maximum transpiration rate. No clear relationship could be found between hydraulic efficiency and either net CO(2) assimilation rates, water-use efficiency estimates (intrinsic water-use efficiency and carbon isotope discrimination against (13)C), or stomatal characteristics (stomatal density and stomatal pore area index). Estimates of hydraulic efficiency were negatively associated with relative growth rate. This unusual pattern, combined with the trade-off observed between hydraulic efficiency and xylem safety, provides the rationale for the positive link already reported between relative growth rate and xylem safety among the same eight P. deltoides×P. nigra genotypes.
Katherine J. Elliott; James M. Vose
1994-01-01
We measured net photosynthesis,leaf conductance, xylem water potential, and growth of Pinus strbus L. seedlings two years after planting on two clear-cut and burned sites in the southern Appalachians. Multiple regression analysis was used to relate seedling net pholosynthesis to vapor pressure deficit, seedling crown temperature, photosynthetically active radiation (...
Loss of water transport capacity due to xylem cavitation in roots of two CAM succulents.
Linton, M J; Nobel, P S
1999-11-01
Loss of axial hydraulic conductance as a result of xylem cavitation was examined for roots of the Crassulacean acid metabolism (CAM) succulents Agave deserti and Opuntia ficus-indica. Vulnerability to cavitation was not correlated with either root size or vessel diameter. Agave deserti had a mean cavitation pressure of -0.93 ± 0.08 MPa by both an air-injection and a centrifugal method compared to -0.70 ± 0.02 MPa by the centrifugal method for O. ficus-indica, reflecting the greater tolerance of the former species to low water potentials in its native habitat. Substantial xylem cavitation would occur at a soil water potential of -0.25 MPa, resulting in a predicted 22% loss of conductance for A. deserti and 32% for O. ficus-indica. For an extended drought of 3 mo, further cavitation could cause a 69% loss of conductance for A. deserti and 62% for O. ficus-indica. A model of axial hydraulic flow based upon the cavitation response of these species predicted that water uptake rates are far below the maximum possible, owing to the high root water potentials of these desert succulents. Despite various shoot adaptations to aridity, roots of A. deserti and O. ficus-indica are highly vulnerable to cavitation, which partially limits water uptake in a wet soil but helps reduce water loss to a drying soil.
Wason, Jay W; Anstreicher, Katherine S; Stephansky, Nathan; Huggett, Brett A; Brodersen, Craig R
2018-04-16
During drought, xylem sap pressures can approach or exceed critical thresholds where gas embolisms form and propagate through the xylem network, leading to systemic hydraulic dysfunction. The vulnerability segmentation hypothesis (VSH) predicts that low-investment organs (e.g. leaf petioles) should be more vulnerable to embolism spread compared to high-investment, perennial organs (e.g. trunks, stems), as a means of mitigating embolism spread and excessive negative pressures in the perennial organs. We tested this hypothesis by measuring air-seeding thresholds using the single-vessel air-injection method and calculating hydraulic safety margins in four northern hardwood tree species of the northeastern United States, in both saplings and canopy height trees, and at five points along the soil-plant-atmosphere continuum. Acer rubrum was the most resistant to air-seeding and generally supported the VSH. However, Fagus grandifolia, Fraxinus americana and Quercus rubra showed little to no variation in air-seeding thresholds across organ types within each species. Leaf-petiole xylem operated at water potentials close to or exceeding their hydraulic safety margins in all species, whereas roots, trunks and stems of A. rubrum, F. grandifolia and Q. rubra operated within their safety margins, even during the third-driest summer in the last 100 yr. © 2018 The Authors. New Phytologist © 2018 New Phytologist Trust.
Ogasa, Mayumi; Miki, Naoko H; Murakami, Yuki; Yoshikawa, Ken
2013-04-01
Woody species hydraulically vulnerable to xylem cavitation may experience daily xylem embolism. How such species cope with the possibility of accumulated embolism is unclear. In this study, we examined seven temperate woody species to assess the hypothesis that low cavitation resistance (high vulnerability to cavitation) is compensated by high recovery performance via vessel refilling. We also evaluated leaf functional and xylem structural traits. The xylem recovery index (XRI), defined as the ratio of xylem hydraulic conductivity in plants rewatered after soil drought to that in plants under moist conditions, varied among species. The xylem water potential causing 50% loss of hydraulic conductivity (Ψ50) varied among the species studied, whereas only a slight difference was detected with respect to midday xylem water potential (Ψmin), indicating smaller hydraulic safety margins (Ψmin - Ψ50) for species more vulnerable to cavitation. Cavitation resistance (|Ψ50|) was negatively correlated with XRI across species, with cavitation-vulnerable species showing a higher performance in xylem recovery. Wood density was positively correlated with cavitation resistance and was negatively correlated with XRI. These novel results reveal that coordination exists between cavitation resistance and xylem recovery performance, in association with wood functional traits such as denser wood for cavitation-resistant xylem and less-dense but water-storable wood for refillable xylem. These findings provide insights into long-term maintenance of water transport in tree species growing under variable environmental conditions.
Assessing plant hydraulic architecture with ultrasonic acoustic emission techniques
NASA Astrophysics Data System (ADS)
Meinzer, F. C.; Johnson, D.; McCulloh, K.; Woodruff, D.
2012-12-01
Water is transported through the xylem of plants under tension (negative pressure). If the tension within a xylem conduit exceeds a critical value, cavitation can occur, which if followed by embolism leads to blockage of water transport through the conduit. Plant species and different organs within the plant such as roots, stems and leaves vary widely in the xylem tension thresholds at which cavitation events begin to occur. Massive cavitation and embolism can lead to catastrophic hydraulic failure and plant death from dehydration. Ultrasonic acoustic emission (UAE) transducers provide a non-invasive means of detecting cavitation events in plants and recording the accumulation of these events through time. When used in combination with other techniques, recording of UAEs can be a powerful tool for characterizing and understanding plant hydraulic architecture; the collection of properties that determine the efficiency and vulnerability of water transport from roots to leaves. The hydraulic architecture of leaves is particularly complex because water must traverse the dead cells of the xylem plus an extra-xylary pathway consisting of living cells and intercellular spaces before it arrives at the internal evaporating surfaces. We used UAE, imaging and other techniques to determine the extent to which dehydration-induced declines in leaf hydraulic conductance were associated with xylem cavitation and embolism versus changes in the conductance of the extra-xylary pathway. In most of the evergreen and deciduous tree species studied there was a close correspondence between the trajectories of cumulative UAEs and loss of whole-leaf hydraulic conductance during dehydration. The mean amplitude of UAEs was positively correlated with mean conduit diameter indicating that in addition to detecting cavitation events, analysis of UAE features can provide information about relative changes in xylem hydraulic conductivity because conductivity is a function of conduit radius to the fourth power. We were unable to detect UAEs from conduits smaller than about 4 micrometers in diameter. The occurrence of embolism in leaf xylem was confirmed independently with cryo-SEM and light microscopy imaging. As expected, there was considerable variation among species in the leaf water potential thresholds at which xylem cavitation and loss of hydraulic conductance began to occur. Contrary to suggestions from some published studies, we saw no evidence of reversible partial xylem conduit collapse instead of cavitation during dehydration. Results from our field studies indicate that dehydration-induced embolism in leaf xylem is completely reversible over a 24-hour cycle, but the mechanism for refilling xylem conduits is unknown. We discuss constraints on the application of UAE techniques for studying plant hydraulics.
Renard, Justine; Tjoelker, Mark G.; Salih, Anya
2015-01-01
The transfer of water from phloem into xylem is thought to mitigate increasing hydraulic tension in the vascular system of trees during the diel cycle of transpiration. Although a putative plant function, to date there is no direct evidence of such water transfer or the contributing pathways. Here, we trace the radial flow of water from the phloem into the xylem and investigate its diel variation. Introducing a fluorescent dye (0.1% [w/w] fluorescein) into the phloem water of the tree species Eucalyptus saligna allowed localization of the dye in phloem and xylem tissues using confocal laser scanning microscopy. Our results show that the majority of water transferred between the two tissues is facilitated via the symplast of horizontal ray parenchyma cells. The method also permitted assessment of the radial transfer of water during the diel cycle, where changes in water potential gradients between phloem and xylem determine the extent and direction of radial transfer. When injected during the morning, when xylem water potential rapidly declined, fluorescein was translocated, on average, farther into mature xylem (447 ± 188 µm) compared with nighttime, when xylem water potential was close to zero (155 ± 42 µm). These findings provide empirical evidence to support theoretical predictions of the role of phloem-xylem water transfer in the hydraulic functioning of plants. This method enables investigation of the role of phloem tissue as a dynamic capacitor for water storage and transfer and its contribution toward the maintenance of the functional integrity of xylem in trees. PMID:25588734
Loewenstein, Nancy J.; Pallardy, Stephen G.
1998-07-01
Patterns of water relations, xylem sap abscisic acid concentration ([ABA]) and stomatal aperture were characterized and compared in drought-sensitive black walnut (Juglans nigra L.), less drought-sensitive sugar maple (Acer saccharum Marsh.) and drought-tolerant white oak (Quercus alba L.) trees co-occurring in a second-growth forest in Missouri, USA. There were strong correlations among reduction in predawn leaf water potential, increased xylem sap [ABA] and stomatal closure in all species. Stomatal conductance was more closely correlated with xylem sap ABA concentration than with ABA flux or xylem sap pH and cation concentrations. In isohydric black walnut, increased concentrations of ABA in the xylem sap appeared to be primarily of root origin, causing stomatal closure in response to soil drying. In anisohydric sugar maple and white oak, however, there were reductions in midday leaf water potential associated with stomatal closure, making it uncertain whether drought-induced xylem sap ABA was of leaf or root origin. The role of root-originated xylem sap ABA in these species as a signal to the shoot of the water status of the roots is, therefore, less certain.
NASA Astrophysics Data System (ADS)
Ding, J.; Johnson, E. A.; Martin, Y. E.
2017-12-01
Leaf is the basic production unit of plants. Water is the most critical resource of plants. Its availability controls primary productivity of plants by affecting leaf carbon budget. To avoid the damage of cavitation from lowering vein water potential t caused by evapotranspiration, the leaf must increase the stomatal resistance to reduce evapotranspiration rate. This comes at the cost of reduced carbon fixing rate as increasing stoma resistance meanwhile slows carbon intake rate. Studies suggest that stoma will operate at an optimal resistance to maximize the carbon gain with respect to water. Different plant species have different leaf shapes, a genetically determined trait. Further, on the same plant leaf size can vary many times in size that is related to soil moisture, an indicator of water availability. According to metabolic scaling theory, increasing leaf size will increase total xylem resistance of vein, which may also constrain leaf carbon budget. We present a Constrained Maximization Model of leaf (leaf CMM) that incorporates metabolic theory into the coupling of evapotranspiration and carbon fixation to examine how leaf size, stoma resistance and maximum net leaf primary productivity change with petiole xylem water potential. The model connects vein network structure to leaf shape and use the difference between petiole xylem water potential and the critical minor vein cavitation forming water potential as the budget. The CMM shows that both maximum net leaf primary production and optimal leaf size increase with petiole xylem water potential while optimal stoma resistance decreases. Narrow leaf has overall lower optimal leaf size and maximum net leaf carbon gain and higher optimal stoma resistance than those of broad leaf. This is because with small width to length ratio, total xylem resistance increases faster with leaf size. Total xylem resistance of narrow leaf increases faster with leaf size causing higher average and marginal cost of xylem water potential with respect to net leaf carbon gain. With same leaf area, total xylem resistance of narrow leaf is higher than broad leaf. Given same stoma resistance and petiole water potential, narrow leaf will lose more xylem water potential than broad leaf. Consequently, narrow leaf has smaller size and higher stoma resistance at optimum.
Limitations in the hydraulic pathway: Effects of xylem embolisms on sap velocity and flow
USDA-ARS?s Scientific Manuscript database
Sap flow in plants takes place in the xylem, a hydraulic system that is usually under negative pressure and in which gas and liquid phases are separated by nanoporous, fibrous pit membranes. It has long been known that this system is at risk of drawing gas nanobubbles through these membranes into th...
Santiago, Louis S; De Guzman, Mark E; Baraloto, Christopher; Vogenberg, Jacob E; Brodie, Max; Hérault, Bruno; Fortunel, Claire; Bonal, Damien
2018-05-01
Predicting responses of tropical forests to climate change-type drought is challenging because of high species diversity. Detailed characterization of tropical tree hydraulic physiology is necessary to evaluate community drought vulnerability and improve model parameterization. Here, we measured xylem hydraulic conductivity (hydraulic efficiency), xylem vulnerability curves (hydraulic safety), sapwood pressure-volume curves (drought avoidance) and wood density on emergent branches of 14 common species of Eastern Amazonian canopy trees in Paracou, French Guiana across species with the densest and lightest wood in the plot. Our objectives were to evaluate relationships among hydraulic traits to identify strategies and test the ability of easy-to-measure traits as proxies for hard-to-measure hydraulic traits. Xylem efficiency was related to capacitance, sapwood water content and turgor loss point, and other drought avoidance traits, but not to xylem safety (P 50 ). Wood density was correlated (r = -0.57 to -0.97) with sapwood pressure-volume traits, forming an axis of hydraulic strategy variation. In contrast to drier sites where hydraulic safety plays a greater role, tropical trees in this humid tropical site varied along an axis with low wood density, high xylem efficiency and high capacitance at one end of the spectrum, and high wood density and low turgor loss point at the other. © 2018 The Authors. New Phytologist © 2018 New Phytologist Trust.
Vascular functioning and the water balance of ripening kiwifruit (Actinidia chinensis) berries
Clearwater, Michael J.; Luo, Zhiwei; Ong, Sam Eng Chye; Blattmann, Peter; Thorp, T. Grant
2012-01-01
Indirect evidence suggests that water supply to fleshy fruits during the final stages of development occurs through the phloem, with the xylem providing little water, or acting as a pathway for water loss back to the plant. This inference was tested by examining the water balance and vascular functioning of ripening kiwifruit berries (Actinidia chinensis var. chinensis ‘Hort16A’) exhibiting a pre-harvest ‘shrivel’ disorder in California, and normal development in New Zealand. Dye labelling and mass balance experiments indicated that the xylem and phloem were both functional and contributed approximately equally to the fruit water supply during this stage of development. The modelled fruit water balance was dominated by transpiration, with net water loss under high vapour pressure deficit (Da) conditions in California, but a net gain under cooler New Zealand conditions. Direct measurement of pedicel sap flow under controlled conditions confirmed inward flows in both the phloem and xylem under conditions of both low and high Da. Phloem flows were required for growth, with gradual recovery after a step increase in Da. Xylem flows alone were unable to support growth, but did supply transpiration and were responsive to Da-induced pressure fluctuations. The results suggest that the shrivel disorder was a consequence of a high fruit transpiration rate, and that the perception of complete loss or reversal of inward xylem flows in ripening fruits should be re-examined. PMID:22155631
Bouche, Pauline S; Delzon, Sylvain; Choat, Brendan; Badel, Eric; Brodribb, Timothy J; Burlett, Regis; Cochard, Hervé; Charra-Vaskou, Katline; Lavigne, Bruno; Li, Shan; Mayr, Stefan; Morris, Hugh; Torres-Ruiz, José M; Zufferey, Vivian; Jansen, Steven
2016-04-01
Plants can be highly segmented organisms with an independently redundant design of organs. In the context of plant hydraulics, leaves may be less embolism resistant than stems, allowing hydraulic failure to be restricted to distal organs that can be readily replaced. We quantified drought-induced embolism in needles and stems of Pinus pinaster using high-resolution computed tomography (HRCT). HRCT observations of needles were compared with the rehydration kinetics method to estimate the contribution of extra-xylary pathways to declining hydraulic conductance. High-resolution computed tomography images indicated that the pressure inducing 50% of embolized tracheids was similar between needle and stem xylem (P50 needle xylem = -3.62 MPa, P50 stem xylem = -3.88 MPa). Tracheids in both organs showed no difference in torus overlap of bordered pits. However, estimations of the pressure inducing 50% loss of hydraulic conductance at the whole needle level by the rehydration kinetics method were significantly higher (P50 needle = -1.71 MPa) than P50 needle xylem derived from HRCT. The vulnerability segmentation hypothesis appears to be valid only when considering hydraulic failure at the entire needle level, including extra-xylary pathways. Our findings suggest that native embolism in needles is limited and highlight the importance of imaging techniques for vulnerability curves. © 2015 John Wiley & Sons Ltd.
DOE Office of Scientific and Technical Information (OSTI.GOV)
Powell, Thomas; Moorcroft, Paul
Raw data for xylem vulnerability curves measured on upper canopy branches of mature trees from the Caxiuana and Tapajos National Forests, Para, Brazil. Tapajos samples were harvested from km67 transects, which is nearby the decommissioned throughfall-exclusion, drought-experiment plots. Caxiuana samples were harvested from trees growing in the throughfall-exclusion, drought-experiment plots. Data were collected in 2011 and 2012. Dataset includes: date of measurement, site ID, plot ID, tree ID (species, tree tag #), xylem pressure, percent loss of conductivity. Air injection method was used. Data reference: Powell et al. (2017) Differences in xylem cavitation resistance and leaf hydraulic traits explain differencesmore » in drought tolerance among mature Amazon rainforest trees. Global Change Biology.« less
NASA Technical Reports Server (NTRS)
Cosgrove, D. J.
1987-01-01
This study was carried out to develop improved methods for measuring in-vivo stress relaxation of growing tissues and to compare relaxation in the stems of four different species. When water uptake by growing tissue is prevented, in-vivo stress relaxation occurs because continued wall loosening reduces wall stress and cell turgor pressure. With this procedure one may measure the yield threshold for growth (Y), the turgor pressure in excess of the yield threshold (P-Y), and the physiological wall extensibility (phi). Three relaxation techniques proved useful: "turgor-relaxation", "balance-pressure" and "pressure-block". In the turgor-relaxation method, water is withheld from growing tissue and the reduction in turgor is measured directly with the pressure probe. This technique gives absolute values for P and Y, but requires tissue excision. In the balance-pressure technique, the excised growing region is sealed in a pressure chamber, and the subsequent reduction in water potential is measured as the applied pressure needed to return xylem sap to the cut surface. This method is simple, but only measures (P-Y), not the individual values of P and Y. In the pressure-block technique, the growing tissue is sealed into a pressure chamber, growth is monitored continuously, and just sufficient pressure is applied to the chamber to block growth. The method gives high-resolution kinetics of relaxation and does not require tissue excision, but only measures (P-Y). The three methods gave similar results when applied to the growing stems of pea (Pisum sativum L.), cucumber (Cucumis sativus L.), soybean (Glycine max (L.) Merr.) and zucchini (Curcubita pepo L.) seedlings. Values for (P-Y) averaged between 1.4 and 2.7 bar, depending on species. Yield thresholds averaged between 1.3 and 3.0 bar. Compared with the other methods, relaxation by pressure-block was faster and exhibited dynamic changes in wall-yielding properties. The two pressure-chamber methods were also used to measure the internal water-potential gradient (between the xylem and the epidermis) which drives water uptake for growth. For the four species it was small, between 0.3 and 0.6 bar, and so did not limit growth substantially.
Mayr, Stefan; Gruber, Andreas; Bauer, Helmut
2003-07-01
Freezing and thawing lead to xylem embolism when gas bubbles caused by ice formation expand during the thaw process. However, previous experimental studies indicated that conifers are resistant to freezing-induced embolism, unless xylem pressure becomes very negative during the freezing. In this study, we show that conifers experienced freezing-induced embolism when exposed to repeated freeze-thaw cycles and simultaneously to drought. Simulating conditions at the alpine timberline (128 days with freeze-thaw events and thawing rates of up to 9.5 K h(-1) in the xylem of exposed twigs during winter), young trees of Norway spruce [Picea abies (L.) Karst.] and stone pine (Pinus cembra L.) were exposed to 50 and 100 freeze-thaw cycles. This treatment caused a significant increase in embolism rates in drought-stressed samples. Upon 100 freeze-thaw cycles, vulnerability thresholds (50% loss of conductivity) were shifted 1.8 MPa (Norway spruce) and 0.8 MPa (stone pine) towards less negative water potentials. The results demonstrate that freeze-thaw cycles are a possible reason for winter-embolism in conifers observed in several field studies. Freezing-induced embolism may contribute to the altitudinal limits of conifers.
Boyer, J S; Cavalieri, A J; Schulze, E D
1985-04-01
A new guillotine thermocouple psychrometer was used to make continuous measurements of water potential before and after the excision of elongating and mature regions of darkgrown soybean (Glycine max L. Merr.) stems. Transpiration could not occur, but growth took place during the measurement if the tissue was intact. Tests showed that the instrument measured the average water potential of the sampled tissue and responded rapidly to changes in water potential. By measuring tissue osmotic potential (Ψ s ), turgor pressure (Ψ p ) could be calculated. In the intact plant, Ψ s and Ψ p were essentially constant for the entire 22 h measurement, but Ψ s was lower and Ψ p higher in the elongating region than in the mature region. This caused the water potential in the elongating region to be lower than in the mature region. The mature tissue equilibrated with the water potential of the xylem. Therefore, the difference in water potential between mature and elongating tissue represented a difference between the xylem and the elongating region, reflecting a water potential gradient from the xylem to the epidermis that was involved in supplying water for elongation. When mature tissue was excised with the guillotine, Ψ s and Ψ p did not change. However, when elongating tissue was excised, water was absorbed from the xylem, whose water potential decreased. This collapsed the gradient and prevented further water uptake. Tissue Ψ p then decreased rapidly (5 min) by about 0.1 MPa in the elongating tissue. The Ψ p decreased because the cell walls relaxed as extension, caused by Ψ p , continued briefly without water uptake. The Ψ p decreased until the minimum for wall extension (Y) was reached, whereupon elongation ceased. This was followed by a slow further decrease in Y but no additional elongation. In elongating tissue excised with mature tissue attached, there was almost no effect on water potential or Ψ p for several hours. Nevertheless, growth was reduced immediately and continued at a decreasing rate. In this case, the mature tissue supplied water to the elongating tissue and the cell walls did not relax. Based on these measurements, a theory is presented for simultaneously evaluating the effects of water supply and water demand associated with growth. Because wall relaxation measured with the psychrometer provided a new method for determining Y and wall extensibility, all the factors required by the theory could be evaluated for the first time in a single sample. The analysis showed that water uptake and wall extension co-limited elongation in soybean stems under our conditions. This co-limitation explains why elongation responded immediately to a decrease in the water potential of the xylem and why excision with attached mature tissue caused an immediate decrease in growth rate without an immediate change in Ψ p.
How to become a tree without wood--biomechanical analysis of the stem of Carica papaya L.
Kempe, A; Lautenschläger, T; Lange, A; Neinhuis, C
2014-01-01
Carica papaya L. does not contain wood, according to the botanical definition of wood as lignified secondary xylem. Despite its parenchymatous secondary xylem, these plants are able to grow up to 10-m high. This is surprising, as wooden structural elements are the ubiquitous strategy for supporting height growth in plants. Proposed possible alternative principles to explain the compensation for lack of wood in C. papaya are turgor pressure of the parenchyma, lignified phloem fibres in the bark, or a combination of the two. Interestingly, lignified tissue comprises only 5-8% of the entire stem mass. Furthermore, the phloem fibres do not form a compact tube enclosing the xylem, but instead form a mesh tubular structure. To investigate the mechanism of papaya's unusually high mechanical strength, a set of mechanical measurements were undertaken on whole stems and tissue sections of secondary phloem and xylem. The structural Young's modulus of mature stems reached 2.5 GPa. Since this is low compared to woody plants, the flexural rigidity of papaya stem construction may mainly be based on a higher second moment of inertia. Additionally, stem turgor pressure was determined indirectly by immersing specimens in sucrose solutions of different osmolalities, followed by mechanical tests; turgor pressure was between 0.82 and 1.25 MPa, indicating that turgor is essential for flexural rigidity of the entire stem. © 2013 German Botanical Society and The Royal Botanical Society of the Netherlands.
Yin, Xiao-Han; Sterck, Frank; Hao, Guang-You
2018-04-23
Some temperate tree species mitigate the negative impacts of frost-induced xylem cavitation by restoring impaired hydraulic function via positive pressures, and may therefore be more resistant to frost fatigue (the phenomenon that post-freezing xylem becomes more susceptible to hydraulic dysfunction) than nonpressure-generating species. We test this hypothesis and investigate underlying anatomical/physiological mechanisms. Using a common garden experiment, we studied key hydraulic traits and detailed xylem anatomical characteristics of 18 sympatric tree species. These species belong to three functional groups, that is, one generating both root and stem pressures (RSP), one generating only root pressure (RP), and one unable to generate such pressures (NP). The three functional groups diverged substantially in hydraulic efficiency, resistance to drought-induced cavitation, and frost fatigue resistance. Most notably, RSP and RP were more resistant to frost fatigue than NP, but this was at the cost of reduced hydraulic conductivity for RSP and reduced resistance to drought-induced cavitation for RP. Our results show that, in environments with strong frost stress: these groups diverge in hydraulic functioning following multiple trade-offs between hydraulic efficiency, resistance to drought and resistance to frost fatigue; and how differences in anatomical characteristics drive such divergence across species. © 2018 The Authors. New Phytologist © 2018 New Phytologist Trust.
Scaling of xylem and phloem transport capacity and resource usage with tree size
Hölttä, Teemu; Kurppa, Miika; Nikinmaa, Eero
2013-01-01
Xylem and phloem need to maintain steady transport rates of water and carbohydrates to match the exchange rates of these compounds at the leaves. A major proportion of the carbon and nitrogen assimilated by a tree is allocated to the construction and maintenance of the xylem and phloem long distance transport tissues. This proportion can be expected to increase with increasing tree size due to the growing transport distances between the assimilating tissues, i.e., leaves and fine roots, at the expense of their growth. We formulated whole tree level scaling relations to estimate how xylem and phloem volume, nitrogen content and hydraulic conductance scale with tree size, and how these properties are distributed along a tree height. Xylem and phloem thicknesses and nitrogen contents were measured within varying positions in four tree species from Southern Finland. Phloem volume, nitrogen amount and hydraulic conductance were found to be concentrated toward the branch and stem apices, in contrast to the xylem where these properties were more concentrated toward the tree base. All of the species under study demonstrated very similar trends. Total nitrogen amount allocated to xylem and phloem was predicted to be comparable to the nitrogen amount allocated to the leaves in small and medium size trees, and to increase significantly above the nitrogen content of the leaves in larger trees. Total volume, hydraulic conductance and nitrogen content of the xylem were predicted to increase faster than that of the phloem with increasing tree height in small trees (<~10 m in height). In larger trees, xylem sapwood turnover to heartwood, if present, would maintain phloem conductance at the same level with xylem conductance with further increases in tree height. Further simulations with a previously published xylem-phloem transport model demonstrated that the Münch pressure flow hypothesis could explain phloem transport with increasing tree height even for the tallest trees. PMID:24367373
Loewenstein, Nancy J.; Pallardy, Stephen G.
1998-07-01
Patterns of water relations, xylem sap abscisic acid (ABA) concentration ([ABA]) and stomatal aperture were compared in drought-sensitive black walnut (Juglans nigra L.) and black willow (Salix nigra Marsh.), less drought-sensitive sugar maple (Acer saccharum Marsh.) and drought-tolerant white oak (Quercus alba L.). Strong correlations among reduction in predawn water potential, increase in xylem sap [ABA] and stomatal closure were observed in all species. Stomatal response was more highly correlated with xylem [ABA] than with ABA flux. Xylem sap pH and ion concentrations appeared not to play a major role in the stomatal response of these species. Stomata were more sensitive to relative changes in [ABA] in drought-sensitive black walnut and black willow than in sugar maple and white oak. In the early stages of drought, increased [ABA] in the xylem sap of black walnut and black willow was probably of root origin and provided a signal to the shoot of the water status of the roots. In sugar maple and white oak, leaf water potential declined with the onset of stomatal closure, so that stomatal closure also may have occurred in response to the change in leaf water potential.
A closed-form solution for steady-state coupled phloem/xylem flow using the Lambert-W function.
Hall, A J; Minchin, P E H
2013-12-01
A closed-form solution for steady-state coupled phloem/xylem flow is presented. This incorporates the basic Münch flow model of phloem transport, the cohesion model of xylem flow, and local variation in the xylem water potential and lateral water flow along the transport pathway. Use of the Lambert-W function allows this solution to be obtained under much more general and realistic conditions than has previously been possible. Variation in phloem resistance (i.e. viscosity) with solute concentration, and deviations from the Van't Hoff expression for osmotic potential are included. It is shown that the model predictions match those of the equilibrium solution of a numerical time-dependent model based upon the same mechanistic assumptions. The effect of xylem flow upon phloem flow can readily be calculated, which has not been possible in any previous analytical model. It is also shown how this new analytical solution can handle multiple sources and sinks within a complex architecture, and can describe competition between sinks. The model provides new insights into Münch flow by explicitly including interactions with xylem flow and water potential in the closed-form solution, and is expected to be useful as a component part of larger numerical models of entire plants. © 2013 John Wiley & Sons Ltd.
Powell, Thomas L; Wheeler, James K; de Oliveira, Alex A R; da Costa, Antonio Carlos Lola; Saleska, Scott R; Meir, Patrick; Moorcroft, Paul R
2017-10-01
Considerable uncertainty surrounds the impacts of anthropogenic climate change on the composition and structure of Amazon forests. Building upon results from two large-scale ecosystem drought experiments in the eastern Brazilian Amazon that observed increases in mortality rates among some tree species but not others, in this study we investigate the physiological traits underpinning these differential demographic responses. Xylem pressure at 50% conductivity (xylem-P 50 ), leaf turgor loss point (TLP), cellular osmotic potential (π o ), and cellular bulk modulus of elasticity (ε), all traits mechanistically linked to drought tolerance, were measured on upper canopy branches and leaves of mature trees from selected species growing at the two drought experiment sites. Each species was placed a priori into one of four plant functional type (PFT) categories: drought-tolerant versus drought-intolerant based on observed mortality rates, and subdivided into early- versus late-successional based on wood density. We tested the hypotheses that the measured traits would be significantly different between the four PFTs and that they would be spatially conserved across the two experimental sites. Xylem-P 50 , TLP, and π o , but not ε, occurred at significantly higher water potentials for the drought-intolerant PFT compared to the drought-tolerant PFT; however, there were no significant differences between the early- and late-successional PFTs. These results suggest that these three traits are important for determining drought tolerance, and are largely independent of wood density-a trait commonly associated with successional status. Differences in these physiological traits that occurred between the drought-tolerant and drought-intolerant PFTs were conserved between the two research sites, even though they had different soil types and dry-season lengths. This more detailed understanding of how xylem and leaf hydraulic traits vary between co-occuring drought-tolerant and drought-intolerant tropical tree species promises to facilitate a much-needed improvement in the representation of plant hydraulics within terrestrial ecosystem and biosphere models, which will enhance our ability to make robust predictions of how future changes in climate will affect tropical forests. © 2017 John Wiley & Sons Ltd.
Hydraulic Limits on Maximum Plant Transpiration
NASA Astrophysics Data System (ADS)
Manzoni, S.; Vico, G.; Katul, G. G.; Palmroth, S.; Jackson, R. B.; Porporato, A. M.
2011-12-01
Photosynthesis occurs at the expense of water losses through transpiration. As a consequence of this basic carbon-water interaction at the leaf level, plant growth and ecosystem carbon exchanges are tightly coupled to transpiration. In this contribution, the hydraulic constraints that limit transpiration rates under well-watered conditions are examined across plant functional types and climates. The potential water flow through plants is proportional to both xylem hydraulic conductivity (which depends on plant carbon economy) and the difference in water potential between the soil and the atmosphere (the driving force that pulls water from the soil). Differently from previous works, we study how this potential flux changes with the amplitude of the driving force (i.e., we focus on xylem properties and not on stomatal regulation). Xylem hydraulic conductivity decreases as the driving force increases due to cavitation of the tissues. As a result of this negative feedback, more negative leaf (and xylem) water potentials would provide a stronger driving force for water transport, while at the same time limiting xylem hydraulic conductivity due to cavitation. Here, the leaf water potential value that allows an optimum balance between driving force and xylem conductivity is quantified, thus defining the maximum transpiration rate that can be sustained by the soil-to-leaf hydraulic system. To apply the proposed framework at the global scale, a novel database of xylem conductivity and cavitation vulnerability across plant types and biomes is developed. Conductivity and water potential at 50% cavitation are shown to be complementary (in particular between angiosperms and conifers), suggesting a tradeoff between transport efficiency and hydraulic safety. Plants from warmer and drier biomes tend to achieve larger maximum transpiration than plants growing in environments with lower atmospheric water demand. The predicted maximum transpiration and the corresponding leaf water potential compare well with measured peak transpiration and minimum water potentials across plant types and biomes, suggesting that plant water transport system and stomatal regulation co-evolved to meet peak atmospheric demands, thus sustaining carbon uptake while avoiding tissue damage even in such harsh conditions.
Carvalho, Mónica R; Losada, Juan M; Niklas, Karl J
2018-06-01
The survival of all vascular plants depends on phloem and xylem, which comprise a hydraulically coupled tissue system that transports photosynthates, water, and a variety of other molecules and ions. Although xylem hydraulics has been extensively studied, until recently, comparatively little is known quantitatively about the phloem hydraulic network and how it is functionally coupled to the xylem network, particularly in photosynthetic leaves. Here, we summarize recent advances in quantifying phloem hydraulics in fully expanded mature leaves with different vascular architectures and show that (1) the size of phloem conducting cells across phylogenetically different taxa scales isometrically with respect to xylem conducting cell size, (2) cell transport areas and lengths increase along phloem transport pathways in a manner that can be used to model Münch's pressure-flow hypothesis, and (3) report observations that invalidate da Vinci's and Murray's hydraulic models as plausible constructs for understanding photosynthate transport in the leaf lamina. Copyright © 2017 Elsevier Ltd. All rights reserved.
Gas flow in plant microfluidic networks controlled by capillary valves
NASA Astrophysics Data System (ADS)
Capron, M.; Tordjeman, Ph.; Charru, F.; Badel, E.; Cochard, H.
2014-03-01
The xylem vessels of trees constitute a model natural microfluidic system. In this work, we have studied the mechanism of air flow in the Populus xylem. The vessel microstructure was characterized by optical microscopy, transmission electronic microscopy (TEM), and atomic force microscopy (AFM) at different length scales. The xylem vessels have length ≈15 cm and diameter ≈20μm. Flow from one vessel to the next occurs through ˜102 pits, which are grouped together at the ends of the vessels. The pits contain a thin, porous pit membrane with a thickness of 310 nm. We have measured the Young's moduli of the vessel wall and of the pits (both water-saturated and after drying) by specific nanoindentation and nanoflexion experiments with AFM. We found that both the dried and water-saturated pit membranes have Young's modulus around 0.4 MPa, in agreement with values obtained by micromolding of pits deformed by an applied pressure difference. Air injection experiments reveal that air flows through the xylem vessels when the differential pressure across a sample is larger than a critical value ΔPc=1.8 MPa. In order to model the air flow rate for ΔP ⩾ΔPc, we assumed the pit membrane to be a porous medium that is strained by the applied pressure difference. Water menisci in the pit pores play the role of capillary valves, which open at ΔP =ΔPc. From the point of view of the plant physiology, this work presents a basic understanding of the physics of bordered pits.
Hao, Guang-You; Wheeler, James K; Holbrook, N Michele; Goldstein, Guillermo
2013-05-01
Trunks of large trees play an important role in whole-plant water balance but technical difficulties have limited most hydraulic research to small stems, leaves, and roots. To investigate the dynamics of water-related processes in tree trunks, such as winter embolism refilling, xylem hydraulic vulnerability, and water storage, volumetric water content (VWC) in the main stem was monitored continuously using frequency domain moisture sensors in adult Betula papyrifera trees from early spring through the beginning of winter. An air injection technique was developed to estimate hydraulic vulnerability of the trunk xylem. Trunk VWC increased in early spring and again in autumn, concurrently with root pressure during both seasons. Diurnal fluctuations and a gradual decrease in trunk VWC through the growing season were observed, which, in combination with VWC increase after significant rainfall events and depletion during periods of high water demand, indicate the importance of stem water storage in both short- and long-term water balance. Comparisons between the trunk air injection results and conventional branch hydraulic vulnerability curves showed no evidence of 'vulnerability segmentation' between the main stem and small branches in B. papyrifera. Measurements of VWC following air injection, together with evidence from air injection and xylem dye perfusion, indicate that embolized vessels can be refilled by active root pressure but not in the absence of root pressure. The precise, continuous, and non-destructive measurement of wood water content using frequency domain sensors provides an ideal way to probe many hydraulic processes in large tree trunks that are otherwise difficult to investigate.
Hao, Guang-You; Wheeler, James K.; Holbrook, N. Michele; Goldstein, Guillermo
2013-01-01
Trunks of large trees play an important role in whole-plant water balance but technical difficulties have limited most hydraulic research to small stems, leaves, and roots. To investigate the dynamics of water-related processes in tree trunks, such as winter embolism refilling, xylem hydraulic vulnerability, and water storage, volumetric water content (VWC) in the main stem was monitored continuously using frequency domain moisture sensors in adult Betula papyrifera trees from early spring through the beginning of winter. An air injection technique was developed to estimate hydraulic vulnerability of the trunk xylem. Trunk VWC increased in early spring and again in autumn, concurrently with root pressure during both seasons. Diurnal fluctuations and a gradual decrease in trunk VWC through the growing season were observed, which, in combination with VWC increase after significant rainfall events and depletion during periods of high water demand, indicate the importance of stem water storage in both short- and long-term water balance. Comparisons between the trunk air injection results and conventional branch hydraulic vulnerability curves showed no evidence of ‘vulnerability segmentation’ between the main stem and small branches in B. papyrifera. Measurements of VWC following air injection, together with evidence from air injection and xylem dye perfusion, indicate that embolized vessels can be refilled by active root pressure but not in the absence of root pressure. The precise, continuous, and non-destructive measurement of wood water content using frequency domain sensors provides an ideal way to probe many hydraulic processes in large tree trunks that are otherwise difficult to investigate. PMID:23585669
Manzoni, Stefano; Vico, Giulia; Katul, Gabriel; Palmroth, Sari; Jackson, Robert B; Porporato, Amilcare
2013-04-01
Soil and plant hydraulics constrain ecosystem productivity by setting physical limits to water transport and hence carbon uptake by leaves. While more negative xylem water potentials provide a larger driving force for water transport, they also cause cavitation that limits hydraulic conductivity. An optimum balance between driving force and cavitation occurs at intermediate water potentials, thus defining the maximum transpiration rate the xylem can sustain (denoted as E(max)). The presence of this maximum raises the question as to whether plants regulate transpiration through stomata to function near E(max). To address this question, we calculated E(max) across plant functional types and climates using a hydraulic model and a global database of plant hydraulic traits. The predicted E(max) compared well with measured peak transpiration across plant sizes and growth conditions (R = 0.86, P < 0.001) and was relatively conserved among plant types (for a given plant size), while increasing across climates following the atmospheric evaporative demand. The fact that E(max) was roughly conserved across plant types and scales with the product of xylem saturated conductivity and water potential at 50% cavitation was used here to explain the safety-efficiency trade-off in plant xylem. Stomatal conductance allows maximum transpiration rates despite partial cavitation in the xylem thereby suggesting coordination between stomatal regulation and xylem hydraulic characteristics. © 2013 The Authors. New Phytologist © 2013 New Phytologist Trust.
NASA Astrophysics Data System (ADS)
Maneta, M. P.; Simeone, C.; Dobrowski, S.; Holden, Z.; Sapes, G.; Sala, A.; Begueria, S.
2017-12-01
In semiarid regions, drought-induced seedling mortality is considered to be caused by failure in the tree hydraulic column. Understanding the mechanisms that cause hydraulic failure and death in seedlings is important, among other things, to diagnose where some tree species may fail to regenerate, triggering demographic imbalances in the forest that could result in climate-driven shifts of tree species. Ponderosa pine is a common lower tree line species in the western US. Seedlings of ponderosa pine are often subject to low soil water potentials, which require lower water potentials in the xylem and leaves to maintain the negative pressure gradient that drives water upward. The resilience of the hydraulic column to hydraulic tension is species dependent, but from greenhouse experiments, we have identified general tension thresholds beyond which loss of xylem conductivity becomes critical, and mortality in Ponderosa pine seedlings start to occur. We describe this hydraulic behavior of plants using a mechanistic soil-vegetation-atmosphere transfer model. Before we use this models to understand water-stress induced seedling mortality at the landscape scale, we perform a modeling analysis of the dynamics of soil moisture, transpiration, leaf water potential and loss of plant water conductivity using detailed data from our green house experiments. The analysis is done using a spatially distributed model that simulates water fluxes, energy exchanges and water potentials in the soil-vegetation-atmosphere continuum. Plant hydraulic and physiological parameters of this model were calibrated using Monte Carlo methods against information on soil moisture, soil hydraulic potential, transpiration, leaf water potential and percent loss of conductivity in the xylem. This analysis permits us to construct a full portrait of the parameter space for Ponderosa pine seedling and generate posterior predictive distributions of tree response to understand the sensitivity of transpiration, hydraulic tension in the plant, and percent loss of conductivity to environmental stresses.
Jacobsen, Anna L; Pratt, R Brandon
2012-06-01
Vulnerability to cavitation curves are used to estimate xylem cavitation resistance and can be constructed using multiple techniques. It was recently suggested that a technique that relies on centrifugal force to generate negative xylem pressures may be susceptible to an open vessel artifact in long-vesselled species. Here, we used custom centrifuge rotors to measure different sample lengths of 1-yr-old stems of grapevine to examine the influence of open vessels on vulnerability curves, thus testing the hypothesized open vessel artifact. These curves were compared with a dehydration-based vulnerability curve. Although samples differed significantly in the number of open vessels, there was no difference in the vulnerability to cavitation measured on 0.14- and 0.271-m-long samples of Vitis vinifera. Dehydration and centrifuge-based curves showed a similar pattern of declining xylem-specific hydraulic conductivity (K(s)) with declining water potential. The percentage loss in hydraulic conductivity (PLC) differed between dehydration and centrifuge curves and it was determined that grapevine is susceptible to errors in estimating maximum K(s) during dehydration because of the development of vessel blockages. Our results from a long-vesselled liana do not support the open vessel artifact hypothesis. © 2012 The Authors. New Phytologist © 2012 New Phytologist Trust.
Spatial and temporal patterns of xylem sap pH derived from stems and twigs of Populus deltoides L.
DOE Office of Scientific and Technical Information (OSTI.GOV)
Aubrey, Doug P.; Boyles, Justin G.; Krysinsky, Laura S.
2011-02-12
Xylem sap pH (pHX) is critical in determining the quantity of inorganic carbon dissolved in xylem solution from gaseous [CO2] measurements. Studies of internal carbon transport have generally assumed that pHX derived from stems and twigs is similar and that pHX remains constant through time; however, no empirical studies have investigated these assumptions. If any of these assumptions are violated, potentially large errors can be introduced into calculations of dissolved CO 2 in xylem and resulting estimates of internal carbon transport.Wetested the validity of assumptions related to pHX in Populus deltoides L. with a series of non-manipulative experiments. The pHXmore » derived from stems and twigs was generally similar and remained relatively constant through a diel period. The only exception was that pHX derived from lower stem sections at night was higher than that derived from twigs. The pHX derived from stems was similar on clear days when solar radiation and vapor pressure deficit (VPD) were similar, but higher on an overcast day when solar radiation and VPD were lower. Similarly, cloudy conditions immediately before an afternoon thunderstorm increased pHX derived from twigs. The pHX derived from twigs remained similar when measured on sunny afternoons between July and October. Our results suggest that common assumptions of pHX used in studies of internal carbon transport appear valid for P. deltoides and further suggest pHX is influenced by environmental factors, such as solar radiation and VPD that affect transpiration rates.« less
Identity and Behavior of Xylem-Residing Bacteria in Rough Lemon Roots of Florida Citrus Trees †
Gardner, John M.; Feldman, Albert W.; Zablotowicz, Robert M.
1982-01-01
An aseptic vacuum extraction technique was used to obtain xylem fluid from the roots of rough lemon (Citrus jambhiri Lush.) rootstock of Florida citrus trees. Bacteria were consistently isolated from vascular fluid of both healthy and young tree decline-affected trees. Thirteen genera of bacteria were found, the most frequently occurring genera being Pseudomonas (40%), Enterobacter (18%), Bacillus, Corynebacterium, and other gram-positive bacteria (16%), and Serratia (6%). Xylem bacterial counts fluctuated seasonally. Bacterial populations ranged from 0.1 to 22 per mm3 of root tissue (about 102 to 2 × 104 bacteria per g of xylem) when bacterial counts were made on vascular fluid, but these numbers were 10- to 1,000-fold greater when aseptically homogenized xylem tissue was examined similarly. Some of the resident bacteria (4%) are potentially phytopathogenic. It is proposed that xylem bacteria have an important role in the physiology of citrus. PMID:16346030
Siebers, Tyche; Catarino, Bruno; Agusti, Javier
2017-03-01
We have identified new potential regulators of xylem cell-type determination and cellular proliferation in cassava and studied their expression in roots. Results are highly relevant for cassava biotechnology. Cassava's root system is composed of two types of root that coexist in every individual: the fibrous and the storage roots. Whether a root becomes fibrous or storage depends on the xylem cell types that it develops: fibrous roots develop xylem fibres and vessels while storage roots develop parenchyma xylem, the starch-storing tissue. A crucial question in cassava root development is how the specific xylem cell types differentiate and proliferate in the fibrous and storage roots. Using phylogenetic, protein sequence and synteny analyses we identified (1) MeVND6, MeVND7.1, MeVND7.2, MeNST3.1 and MeNST3.2 as the potential cassava orthologues of the Arabidopsis regulators of xylem cell type determination AtVND6, AtVND7 and AtNST3; and (2) MeWOX4.1 and MeWOX4.2 as the potential cassava orthologues of the Arabidopsis cambium regulator AtWOX4. Fibrous and storage roots were anatomically characterised and tested for the expression of the identified genes. Results revealed that (1) MeVND7.1 and MeVND7.2 are expressed in the fibrous but not in the storage roots; (2) MeVND6 shows low expression in both root types; (3) MeNST3.1 is not expressed in the fibrous or storage roots, while MeNST3.2 is highly expressed in both root-types and (4) MeWOX4.1 and, to a higher level, MeWOX4.2 are expressed in both the fibrous and storage roots. Results open new avenues for research in cassava root development and for food security-oriented biotechnology programmes.
NASA Technical Reports Server (NTRS)
Zimmermann, R.; McDonald, K.; Way, J.; Oren, R.
1994-01-01
Tree canopy microclimate, xylem water flux and xylem dielectric constant have been monitored in situ since June 1993 in two adjacent natural forest stands in central Alaska. The deciduous stand represents a mature balsam poplar site on the Tanana River floodplain, while the coniferous stand consists of mature white spruce with some black spruce mixed in. During solstice in June and later in summer, diurnal changes of xylem water potential were measured to investigate the occurrence and magnitude of tree transpiration and dielectric constant changes in stems.
Biles, C L; Abeles, F B
1991-06-01
Xylem sap from apple (Malus domestica Borkh), peach (Prunus persica Batsch), and pear (Pyrus communis L.) twigs was collected by means of pressure extrusion. This sap contained a number of acidic peroxidases and other proteins. Two other sources of xylem sap used in this study were stem exudates and guttation fluid. Similar peroxidases were also found in stem exudates and guttation fluids of strawberry (Fragaria x ananassa Duch.), tomato (Lycopersicum esculentum L.), and cucumber (Cucumis sativus L.). Isoelectric focusing activity gels showed that two peroxidases (isoelectric point [pl] 9 and pl 4.6) were present in initial stem exudates collected in the first 30 minutes after excision. Subsequent samples of stem exudate collected contained only the pl 4.6 isozyme. The pl 4.6 peroxidase isozyme was also found in root tissue and guttation fluid. These observations suggest that roots produce and secrete the pl 4.6 peroxidase into xylem sap. Cucumber seedlings were treated with 100 microliters per liter ethylene for 16 hours and the exudate from decapitated hypocotyl stumps was collected over a 3 hour period. Ethylene increased the peroxidase activity of stem exudates and inhibited the amount of exudate released. These observations suggest that xylem sap peroxidase may play a role in plugging damaged vascular tissue.
Lintunen, A.; Hölttä, T.; Kulmala, M.
2013-01-01
Water in the xylem, the water transport system of plants, is vulnerable to freezing and cavitation, i.e. to phase change from liquid to ice or gaseous phase. The former is a threat in cold and the latter in dry environmental conditions. Here we show that a small xylem conduit diameter, which has previously been shown to be associated with lower cavitation pressure thus making a plant more drought resistant, is also associated with a decrease in the temperature required for ice nucleation in the xylem. Thus the susceptibility of freezing and cavitation are linked together in the xylem of plants. We explain this linkage by the regulation of the sizes of the nuclei catalysing freezing and drought cavitation. Our results offer better understanding of the similarities of adaption of plants to cold and drought stress, and offer new insights into the ability of plants to adapt to the changing environment. PMID:23778457
Species climate range influences hydraulic and stomatal traits in Eucalyptus species.
Bourne, Aimee E; Creek, Danielle; Peters, Jennifer M R; Ellsworth, David S; Choat, Brendan
2017-07-01
Plant hydraulic traits influence the capacity of species to grow and survive in water-limited environments, but their comparative study at a common site has been limited. The primary aim of this study was to determine whether selective pressures on species originating in drought-prone environments constrain hydraulic traits among related species grown under common conditions. Leaf tissue water relations, xylem anatomy, stomatal behaviour and vulnerability to drought-induced embolism were measured on six Eucalyptus species growing in a common garden to determine whether these traits were related to current species climate range and to understand linkages between the traits. Hydraulically weighted xylem vessel diameter, leaf turgor loss point, the water potential at stomatal closure and vulnerability to drought-induced embolism were significantly ( P < 0·05) correlated with climate parameters from the species range. There was a co-ordination between stem and leaf parameters with the water potential at turgor loss, 12 % loss of conductivity and the point of stomatal closure significantly correlated. The correlation of hydraulic, stomatal and anatomical traits with climate variables from the species' original ranges suggests that these traits are genetically constrained. The conservative nature of xylem traits in Eucalyptus trees has important implications for the limits of species responses to changing environmental conditions and thus for species survival and distribution into the future, and yields new information for physiological models. © The Author 2017. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please email: journals.permissions@oup.com
Dié, Agathe; Kitin, Peter; Kouamé, François N'Guessan; Van den Bulcke, Jan; Van Acker, Joris; Beeckman, Hans
2012-01-01
Background and Aims Teak forms xylem rings that potentially carry records of carbon sequestration and climate in the tropics. These records are only useful when the structural variations of tree rings and their periodicity of formation are known. Methods The seasonality of ring formation in mature teak trees was examined via correlative analysis of cambial activity, xylem and phloem formation, and climate throughout 1·5 years. Xylem and phloem differentiation were visualized by light microscopy and scanning electron microscopy. Key Results A 3 month dry season resulted in semi-deciduousness, cambial dormancy and formation of annual xylem growth rings (AXGRs). Intra-annual xylem and phloem growth was characterized by variable intensity. Morphometric features of cambium such as cambium thickness and differentiating xylem layers were positively correlated. Cambium thickness was strongly correlated with monthly rainfall (R2 = 0·7535). In all sampled trees, xylem growth zones (XGZs) were formed within the AXGRs during the seasonal development of new foliage. When trees achieved full leaf, the xylem in the new XGZs appeared completely differentiated and functional for water transport. Two phloem growth rings were formed in one growing season. Conclusions The seasonal formation pattern and microstructure of teak xylem suggest that AXGRs and XGZs can be used as proxies for analyses of the tree history and climate at annual and intra-annual resolution. PMID:22805529
Gleason, Sean M; Westoby, Mark; Jansen, Steven; Choat, Brendan; Hacke, Uwe G; Pratt, Robert B; Bhaskar, Radika; Brodribb, Tim J; Bucci, Sandra J; Cao, Kun-Fang; Cochard, Hervé; Delzon, Sylvain; Domec, Jean-Christophe; Fan, Ze-Xin; Feild, Taylor S; Jacobsen, Anna L; Johnson, Daniel M; Lens, Frederic; Maherali, Hafiz; Martínez-Vilalta, Jordi; Mayr, Stefan; McCulloh, Katherine A; Mencuccini, Maurizio; Mitchell, Patrick J; Morris, Hugh; Nardini, Andrea; Pittermann, Jarmila; Plavcová, Lenka; Schreiber, Stefan G; Sperry, John S; Wright, Ian J; Zanne, Amy E
2016-01-01
The evolution of lignified xylem allowed for the efficient transport of water under tension, but also exposed the vascular network to the risk of gas emboli and the spread of gas between xylem conduits, thus impeding sap transport to the leaves. A well-known hypothesis proposes that the safety of xylem (its ability to resist embolism formation and spread) should trade off against xylem efficiency (its capacity to transport water). We tested this safety-efficiency hypothesis in branch xylem across 335 angiosperm and 89 gymnosperm species. Safety was considered at three levels: the xylem water potentials where 12%, 50% and 88% of maximal conductivity are lost. Although correlations between safety and efficiency were weak (r(2) < 0.086), no species had high efficiency and high safety, supporting the idea for a safety-efficiency tradeoff. However, many species had low efficiency and low safety. Species with low efficiency and low safety were weakly associated (r(2) < 0.02 in most cases) with higher wood density, lower leaf- to sapwood-area and shorter stature. There appears to be no persuasive explanation for the considerable number of species with both low efficiency and low safety. These species represent a real challenge for understanding the evolution of xylem. No claim to US government works. New Phytologist © 2015 New Phytologist Trust.
Domec, J C; Scholz, F G; Bucci, S J; Meinzer, F C; Goldstein, G; Villalobos-Vega, R
2006-01-01
Vulnerability to water-stress-induced embolism and variation in the degree of native embolism were measured in lateral roots of four co-occurring neotropical savanna tree species. Root embolism varied diurnally and seasonally. Late in the dry season, loss of root xylem conductivity reached 80% in the afternoon when root water potential (psi root) was about -2.6 MPa, and recovered to 25-40% loss of conductivity in the morning when psi root was about -1.0 MPa. Daily variation in psi root decreased, and root xylem vulnerability and capacitance increased with rooting depth. However, all species experienced seasonal minimum psi root close to complete hydraulic failure independent of their rooting depth or resistance to embolism. Predawn psi root was lower than psi soil when psi soil was relatively high (> -0.7 MPa) but became less negative than psi soil, later in the dry season, consistent with a transition from a disequilibrium between plant and soil psi induced by nocturnal transpiration to one induced by hydraulic redistribution of water from deeper soil layers. Shallow longitudinal root incisions external to the xylem prevented reversal of embolism overnight, suggesting that root mechanical integrity was necessary for recovery, consistent with the hypothesis that if embolism is a function of tension, refilling may be a function of internal pressure imbalances. All species shared a common relationship in which maximum daily stomatal conductance declined linearly with increasing afternoon loss of root conductivity over the course of the dry season. Daily embolism and refilling in roots is a common occurrence and thus may be an inherent component of a hydraulic signaling mechanism enabling stomata to maintain the integrity of the hydraulic pipeline in long-lived structures such as stems.
Dodd, Ian C; Egea, Gregorio; Davies, William J
2008-01-01
When soil moisture is heterogeneous, sap flow from, and ABA status of, different parts of the root system impact on leaf xylem ABA concentration ([X-ABA]leaf). The robustness of a model for predicting [X-ABA]leaf was assessed. 'Two root-one shoot' grafted sunflower (Helianthus annuus L.) plants received either deficit irrigation (DI, each root system received the same irrigation volumes) or partial rootzone drying (PRD, only one root system was watered and the other dried the soil). Irrespective of whether relative sap flow was assessed using sap flow sensors in vivo or by pressurization of de-topped roots, each root system contributed similarly to total sap flow during DI, while sap flow from roots in drying soil declined linearly with soil water potential (Psisoil) during PRD. Although Psisoil of the irrigated pot determined the threshold Psisoil at which sap flow from roots in drying soil decreased, the slope of this decrease was independent of the wet pot Psisoil. Irrespective of whether sap was collected from the wet or dry root system of PRD plants, or a DI plant, root xylem ABA concentration increased as Psisoil declined. The model, which weighted ABA contributions of each root system according to the sap flow from each, almost perfectly explained [X-ABA] immediately above the graft union. That the model overestimated measured [X-ABA]leaf may result from changes in [X-ABA] along the transport pathway or an artefact of collecting xylem sap from detached leaves. The implications of declining sap flow through partially dry roots during PRD for the control of stomatal behaviour and irrigation scheduling are discussed.
Pragmatic hydraulic theory predicts stomatal responses to climatic water deficits.
Sperry, John S; Wang, Yujie; Wolfe, Brett T; Mackay, D Scott; Anderegg, William R L; McDowell, Nate G; Pockman, William T
2016-11-01
Ecosystem models have difficulty predicting plant drought responses, partially from uncertainty in the stomatal response to water deficits in soil and atmosphere. We evaluate a 'supply-demand' theory for water-limited stomatal behavior that avoids the typical scaffold of empirical response functions. The premise is that canopy water demand is regulated in proportion to threat to supply posed by xylem cavitation and soil drying. The theory was implemented in a trait-based soil-plant-atmosphere model. The model predicted canopy transpiration (E), canopy diffusive conductance (G), and canopy xylem pressure (P canopy ) from soil water potential (P soil ) and vapor pressure deficit (D). Modeled responses to D and P soil were consistent with empirical response functions, but controlling parameters were hydraulic traits rather than coefficients. Maximum hydraulic and diffusive conductances and vulnerability to loss in hydraulic conductance dictated stomatal sensitivity and hence the iso- to anisohydric spectrum of regulation. The model matched wide fluctuations in G and P canopy across nine data sets from seasonally dry tropical forest and piñon-juniper woodland with < 26% mean error. Promising initial performance suggests the theory could be useful in improving ecosystem models. Better understanding of the variation in hydraulic properties along the root-stem-leaf continuum will simplify parameterization. © 2016 The Authors. New Phytologist © 2016 New Phytologist Trust.
Leaf Pressure Volume Data in Caxiuana and Tapajos National Forest, Para, Brazil (2011)
DOE Office of Scientific and Technical Information (OSTI.GOV)
Powell, Thomas; Moorcroft, Paul
Pressure volume curve measurements on leaves of canopy trees from the from the Caxiuana and Tapajos National Forests, Para, Brazil. Tapajos samples were harvested from the km 67 forested area, which is adjacent to the decommissioned throughfall exclusion drought experimental plot. Caxiuana samples were harvested from trees growing in the throughfall exclusion plots. Data were collected in 2011. Dataset includes: date of measurement, site ID, plot ID, tree ID (species, tree tag #), leaf area, fresh weight, relative weight, leaf water potential, and leaf water loss. P-V curve parameters (turgor loss point, osmotic potential, and bulk modulus of elasticity) canmore » be found in Powell et al. (2017) Differences in xylem cavitation resistance and leaf hydraulic traits explain differences in drought tolerance among mature Amazon rainforest trees. Global Change Biology.« less
Medeiros, Juliana S.; Ward, Joy K.
2013-01-01
Summary Changes in atmospheric carbon dioxide concentration ([CO2]) affect plant carbon/water trade-offs, with implications for drought tolerance. Leaf-level studies often indicate that drought tolerance may increase with rising [CO2], but integrated leaf and xylem responses are not well understood in this respect. In addition, the influence of low [CO2] of the last glacial period on drought tolerance and xylem properties is not well understood.We investigated the interactive effects of a broad range of [CO2] and plant water potentials on leaf function, xylem structure and function and the integration of leaf and xylem function in Phaseolus vulgaris.Elevated [CO2] decreased vessel implosion strength, reduced conduit specific hydraulic conductance, and compromised leaf specific xylem hydraulic conductance under moderate drought. By contrast, at glacial [CO2], transpiration was maintained under moderate drought via greater conduit specific and leaf specific hydraulic conductance in association with increased vessel implosion strength.Our study involving the integration of leaf and xylem responses suggests that increasing [CO2] does not improve drought tolerance. We show that under glacial conditions changes in leaf and xylem properties could increase drought tolerance, while under future conditions greater productivity may only occur when higher water use can be accommodated. PMID:23668237
McCully
1999-03-01
Embolism and refilling of vessels was monitored directly by cryomicroscopy of field-grown corn (Zea mays L.) roots. To test the reliability of an earlier study showing embolism refilling in roots at negative leaf water potentials, embolisms were counted, and root water potentials (Psiroot) and osmotic potentials of exuded xylem sap from the same roots were measured by isopiestic psychrometry. All vessels were full at dawn (Psiroot -0.1 MPa). Embolisms were first seen in late metaxylem vessels at 8 AM. Embolized late metaxylem vessels peaked at 50% at 10 AM (Psiroot -0.1 MPa), fell to 44% by 12 PM (Psiroot -0.23 MPa), then dropped steadily to zero by early evening (Psiroot -0.28 MPa). Transpiration was highest (8.5 μg cm-2 s-1) between 12 and 2 PM when the percentage of vessels embolized was falling. Embolized vessels were refilled by liquid moving through their lateral walls. Xylem sap was very low in solutes. The mechanism of vessel refilling, when Psiroot is negative, requires further investigation. Daily embolism and refilling in roots of well-watered plants is a normal occurrence and may be a component of an important hydraulic signaling mechanism between roots and shoots.
Daniel M. Johnson; Katherine A. McCulloh; David R. Woodruff; Frederick C. Meinzer
2012-01-01
Angiosperm and coniferous tree species utilize a continuum of hydraulic strategies. Hydraulic safety margins (defined as differences between naturally occurring xylem pressures and pressures that would cause hydraulic dysfunction, or differences between pressures resulting in loss of hydraulic function in adjacent organs (e.g., stems vs. leaves) tend to be much greater...
Modeling the dynamics of pressure propagation and diameter variation in tree sapwood.
Perämäki, Martti; Vesala, Timo; Nikinmaa, Eero
2005-09-01
A non-steady-state model of water tension propagation in tree stems was developed. The model is based on the cohesion theory and the assumption that fluctuating water tension driven by transpiration together with the elasticity of wood cause variations in the diameter of a tree stem. The change in xylem diameter can be linked to water tension in accordance with Hooke's law. The model was tested against field measurements of the diurnal change in xylem diameter at different heights in a 180-year-old Scots pine tree at Hyytiälä, southern Finland. Model predictions agreed well with measurements. The effect of tree dimensions on pressure propagation was examined with the model. The model outcomes were also consistent with results of several field measurements presented in the literature.
James P. Dunn; Peter L. Jr. Lorio
1993-01-01
We modified soil water supply to two groups of juvenile loblolly pines, Pinus taeda L., by sheltering or irrigating root systems in early summer or in later summer and measured oleoresin flow (primary defense), net photosynthesis, xylem water potential, and cambial growth throughout the growing season. When consistent significant differences in...
Tocochromanols in wood: a potential new tool for dendrometabolomics.
Fleta-Soriano, Eva; Fernández-Marín, Beatriz; Olano, José Miguel; Míguez, Fátima; Molinero, Jon; Camarero, Jesús Julio; García-Plazaola, José Ignacio
2014-12-01
Tocochromanols are the most abundant lipid-soluble antioxidants in plants. Among them, α-tocopherol (α-Toc) shows a particularly high sensitivity to environmental stressors and its content is used as a stress biomarker even in non-photosynthetic tissues. Nevertheless, the presence of tocochromanols has not been described yet in the xylem of woody plants, even when their functions regarding cell membrane protection and the transport of photoassimilates may be crucial in this tissue and despite its potential utility in dendrometabolomics. Considering all these, we aimed to determine the presence and distribution of tocochromanols in the xylem of woody plants, to examine their responsiveness to high temperature and to evaluate their potential as environmental bioindicators. The analysis of 29 phyllogenetically diverse species showed that α-Toc is the most abundant and frequent tocochromanol in the xylem and is ubiquitously present in all the studied species, with a concentration ranging from 0.5 to 39.3 μg g(-1) of dry weight. α-Tocopherol appeared to be mainly located in the parenchyma rays and was found in both the sapwood and the heartwood, suggesting that it is present even in dead parenchyma cells. The levels of α-Toc in the xylem did not change in response to locally induced xylem heating, but responded positively to the 3-year moving average of annual precipitation. The present findings suggest that α-Toc may be linked to changes in climatic stress. This should enhance further research on the environmental controls of α-Toc variation in the xylem as a first step towards a deeper understanding of dendrometabolomics. © The Author 2014. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.
Boyle, Richard K A; McAinsh, Martin; Dodd, Ian C
2016-01-01
Soil water deficits applied at different rates and for different durations can decrease both stomatal conductance (gs ) and leaf water potential (Ψleaf ). Understanding the physiological mechanisms regulating these responses is important in sustainable irrigation scheduling. Glasshouse-grown, containerized Pelargonium × hortorum BullsEye plants were irrigated either daily at various fractions of plant evapotranspiration (100, 75 and 50% ET) for 20 days or irrigation was withheld for 4 days. Xylem sap was collected and gs and Ψleaf were measured on days 15 and 20, and on days 16-19 for the respective treatments. Xylem sap pH and NO3 (-) and Ca(2+) concentrations did not differ between irrigation treatments. Xylem abscisic acid (ABA) concentrations ([ABA]xyl ) increased within 24 h of irrigation being withheld whilst gs and Ψleaf decreased. Supplying irrigation at a fraction of daily ET produced a similar relationship between [ABA]xyl and gs , but did not change Ψleaf . Treatment differences occurred independently of whether Ψleaf was measured in whole leaves with a pressure chamber, or in the lamina with a thermocouple psychrometer. Plants that were irrigated daily showed lower [ABA]xyl than plants from which irrigation was withheld, even at comparable soil moisture content. This implies that regular re-watering attenuates ABA signaling due to maintenance of soil moisture in the upper soil levels. Crucially, detached leaves supplied with synthetic ABA showed a similar relationship between [ABA]xyl and gs as intact plants, suggesting that stomatal closure of P. hortorum in response to soil water deficit is primarily an ABA-induced response, independent of changes in Ψleaf . © 2015 Scandinavian Plant Physiology Society.
Lopisso, Daniel Teshome; Knüfer, Jessica; Koopmann, Birger; von Tiedemann, Andreas
2017-09-01
As Verticillium stem striping of oilseed rape (OSR), a vascular disease caused by Verticillium longisporum, is extending into new geographic regions and no control with fungicides exists, the demand for understanding mechanisms of quantitative resistance increases. Because V. longisporum is strictly limited to the xylem and resistance is expressed in the systemic stage post root invasion, we investigated a potential antifungal role of soluble constituents and nutritional conditions in xylem sap as determinants of cultivar resistance of OSR to V. longisporum. Assessment of biometric and molecular genetic parameters applied to describe V. longisporum resistance (net area under disease progress curve, stunting, stem thickness, plant biomass, and V. longisporum DNA content) showed consistent susceptibility of cultivar 'Falcon' in contrast to two resistant genotypes, 'SEM' and 'Aviso'. Spectrophotometric analysis revealed a consistently stronger in vitro growth of V. longisporum in xylem sap extracted from OSR compared with the water control. Further comparisons of fungal growth in xylem sap of different cultivars revealed the absence of constitutive or V. longisporum induced antifungal activity in the xylem sap of resistant versus susceptible genotypes. The similar growth of V. longisporum in xylem sap, irrespective of cultivar, infection with V. longisporum and xylem sap filtration, was correlated with about equal amounts of total soluble proteins in xylem sap from these treatments. Interestingly, compared with younger plants, xylem sap from older plants induced significantly stronger fungal growth. Growth enhancement of V. longisporum in xylem sap of aging plants was reflected by increased contents of carbohydrates, which was consistent in mock or V. longisporum-infected plants and independent from cultivar resistance. The improved nutritional conditions in the xylem of more mature plants may explain the late appearance of disease symptoms, which are observed only in late maturity stages of plants in the field. While falsifying the presence of antifungal activity in xylem sap of resistant cultivars, this study strengthens previous findings that indicated a significant role of physical cell wall bound resistance factors involved in quantitative, cultivar-related resistance of B. napus to V. longisporum.
Sugars from woody tissue photosynthesis reduce xylem vulnerability to cavitation.
De Baerdemaeker, Niels J F; Salomón, Roberto Luis; De Roo, Linus; Steppe, Kathy
2017-11-01
Reassimilation of internal CO 2 via woody tissue photosynthesis has a substantial effect on tree carbon income and wood production. However, little is known about its role in xylem vulnerability to cavitation and its implications in drought-driven tree mortality. Young trees of Populus nigra were subjected to light exclusion at the branch and stem levels. After 40 d, measurements of xylem water potential, diameter variation and acoustic emission (AE) were performed in detached branches to obtain acoustic vulnerability curves to cavitation following bench-top dehydration. Acoustic vulnerability curves and derived AE 50 values (i.e. water potential at which 50% of cavitation-related acoustic emissions occur) differed significantly between light-excluded and control branches (AE 50,light-excluded = -1.00 ± 0.13 MPa; AE 50,control = -1.45 ± 0.09 MPa; P = 0.007) denoting higher vulnerability to cavitation in light-excluded trees. Woody tissue photosynthesis represents an alternative and immediate source of nonstructural carbohydrates (NSC) that confers lower xylem vulnerability to cavitation via sugar-mediated mechanisms. Embolism repair and xylem structural changes could not explain this observation as the amount of cumulative AE and basic wood density did not differ between treatments. We suggest that woody tissue assimilates might play a role in the synthesis of xylem surfactants for nanobubble stabilization under tension. © 2017 The Authors. New Phytologist © 2017 New Phytologist Trust.
Torres-Ruiz, José M; Sperry, John S; Fernández, José E
2012-10-01
Xylem hydraulic conductivity (K) is typically defined as K = F/(P/L), where F is the flow rate through a xylem segment associated with an applied pressure gradient (P/L) along the segment. This definition assumes a linear flow-pressure relationship with a flow intercept (F(0)) of zero. While linearity is typically the case, there is often a non-zero F(0) that persists in the absence of leaks or evaporation and is caused by passive uptake of water by the sample. In this study, we determined the consequences of failing to account for non-zero F(0) for both K measurements and the use of K to estimate the vulnerability to xylem cavitation. We generated vulnerability curves for olive root samples (Olea europaea) by the centrifuge technique, measuring a maximally accurate reference K(ref) as the slope of a four-point F vs P/L relationship. The K(ref) was compared with three more rapid ways of estimating K. When F(0) was assumed to be zero, K was significantly under-estimated (average of -81.4 ± 4.7%), especially when K(ref) was low. Vulnerability curves derived from these under-estimated K values overestimated the vulnerability to cavitation. When non-zero F(0) was taken into account, whether it was measured or estimated, more accurate K values (relative to K(ref)) were obtained, and vulnerability curves indicated greater resistance to cavitation. We recommend accounting for non-zero F(0) for obtaining accurate estimates of K and cavitation resistance in hydraulic studies. Copyright © Physiologia Plantarum 2012.
Scaling of angiosperm xylem structure with safety and efficiency.
Hacke, Uwe G; Sperry, John S; Wheeler, James K; Castro, Laura
2006-06-01
We tested the hypothesis that greater cavitation resistance correlates with less total inter-vessel pit area per vessel (the pit area hypothesis) and evaluated a trade-off between cavitation safety and transport efficiency. Fourteen species of diverse growth form (vine, ring- and diffuse-porous tree, shrub) and family affinity were added to published data predominately from the Rosaceae (29 species total). Two types of vulnerability-to-cavitation curves were found. Ring-porous trees and vines showed an abrupt drop in hydraulic conductivity with increasing negative pressure, whereas hydraulic conductivity in diffuse-porous species generally decreased gradually. The ring-porous type curve was not an artifact of the centrifuge method because it was obtained also with the air-injection technique. A safety versus efficiency trade-off was evident when curves were compared across species: for a given pressure, there was a limited range of optimal vulnerability curves. The pit area hypothesis was supported by a strong relationship (r2 = 0.77) between increasing cavitation resistance and diminishing pit membrane area per vessel (A(P)). Small A(P) was associated with small vessel surface area and hence narrow vessel diameter (D) and short vessel length (L)--consistent with an increase in vessel flow resistance with cavitation resistance. This trade-off was amplified at the tissue level by an increase in xylem/vessel area ratio with cavitation resistance. Ring-porous species were more efficient than diffuse-porous species on a vessel basis but not on a xylem basis owing to higher xylem/vessel area ratios in ring-porous anatomy. Across four orders of magnitude, lumen and end-wall resistivities maintained a relatively tight proportionality with a near-optimal mean of 56% of the total vessel resistivity residing in the end-wall. This was consistent with an underlying scaling of L to D(3/2) across species. Pit flow resistance did not increase with cavitation safety, suggesting that cavitation pressure was not related to mean pit membrane porosity.
Maximum sustainable xylem sap tensions in Rhododendron and other species.
Crombie, D S; Milburn, J A; Hipkins, M F
1985-01-01
The acoustic technique was used in conjunction with the pressure chamber to determine the tensions causing cavitation of xylem sap in leaves of five woody angiosperms (Acer pseudoplatanus L., Alnus glutinosa L. Gaertn., Eucalyptus globulus Labill., Fraxinus excelsior L. and Rhododendron ponticum L.) and three species of herbs (Lycopersicum esculentum Mill., Plantago major L. and Ricinus communis L.). The results showed leaves of most species to suffer considerably from cavitation at sap tensions of 1.6-3 MPa. Two of the herbs, Lycopersicum and Ricinus, cavitated extensively at sap tensions below 1 MPa. Additional evidence is presented that clicks, detected by acoustic amplification, are caused by cavitation of sap in the xylem conduits. A rapid method is suggested for the determination of sap tensions in cavitating leaves and which is suitable for surveys of the critical sap tension in a large number of species.
Sack, Lawren; Scoffoni, Christine
2012-12-31
Water is a key resource, and the plant water transport system sets limits on maximum growth and drought tolerance. When plants open their stomata to achieve a high stomatal conductance (gs) to capture CO2 for photosynthesis, water is lost by transpiration(1,2). Water evaporating from the airspaces is replaced from cell walls, in turn drawing water from the xylem of leaf veins, in turn drawing from xylem in the stems and roots. As water is pulled through the system, it experiences hydraulic resistance, creating tension throughout the system and a low leaf water potential (Ψ(leaf)). The leaf itself is a critical bottleneck in the whole plant system, accounting for on average 30% of the plant hydraulic resistance(3). Leaf hydraulic conductance (K(leaf) = 1/ leaf hydraulic resistance) is the ratio of the water flow rate to the water potential gradient across the leaf, and summarizes the behavior of a complex system: water moves through the petiole and through several orders of veins, exits into the bundle sheath and passes through or around mesophyll cells before evaporating into the airspace and being transpired from the stomata. K(leaf) is of strong interest as an important physiological trait to compare species, quantifying the effectiveness of the leaf structure and physiology for water transport, and a key variable to investigate for its relationship to variation in structure (e.g., in leaf venation architecture) and its impacts on photosynthetic gas exchange. Further, K(leaf) responds strongly to the internal and external leaf environment(3). K(leaf) can increase dramatically with irradiance apparently due to changes in the expression and activation of aquaporins, the proteins involved in water transport through membranes(4), and K(leaf) declines strongly during drought, due to cavitation and/or collapse of xylem conduits, and/or loss of permeability in the extra-xylem tissues due to mesophyll and bundle sheath cell shrinkage or aquaporin deactivation(5-10). Because K(leaf) can constrain gs and photosynthetic rate across species in well watered conditions and during drought, and thus limit whole-plant performance they may possibly determine species distributions especially as droughts increase in frequency and severity(11-14). We present a simple method for simultaneous determination of K(leaf) and gs on excised leaves. A transpiring leaf is connected by its petiole to tubing running to a water source on a balance. The loss of water from the balance is recorded to calculate the flow rate through the leaf. When steady state transpiration (E, mmol • m(-2) • s(-1)) is reached, gs is determined by dividing by vapor pressure deficit, and K(leaf) by dividing by the water potential driving force determined using a pressure chamber (K(leaf)= E /- Δψ(leaf), MPa)(15). This method can be used to assess K(leaf) responses to different irradiances and the vulnerability of K(leaf) to dehydration(14,16,17).
Gonzalez-Rodriguez, David; Cournède, Paul-Henry; de Langre, Emmanuel
2016-06-07
Water stress is a major cause of tree mortality. In response to drought, leaves wilt due to an increase in petiole flexibility. We present an analytical model coupling petiole mechanics, thermal balance, and xylem hydraulics to investigate the role of petiole flexibility in protecting a tree from water stress. Our model suggests that turgidity-dependent petiole flexibility can significantly attenuate the minimal xylem pressure and thus reduce the risk of cavitation. Moreover, we show that petiole flexibility increases water use efficiency by trees under water stress. Copyright © 2016 Elsevier Ltd. All rights reserved.
Coordination of leaf and stem water transport properties in tropical forest trees
Frederick C. Meinzer; David R. Woodruff; Jean-Christophe Domec; Guillermo Goldstein; Paula I. Campanello; Genoveva M. Gatti; Randol Villalobos-Vega
2008-01-01
Stomatal regulation of transpiration constrains leaf water potential (ψ l) within species-specific ranges that presumably avoid excessive tension and embolism in the stem xylem upstream. However, the hydraulic resistance of leaves can be highly variable over short time scales, uncoupling tension in the xylem of leaves from that in the...
Love, Christopher J.; Zhang, Shuguang; Mershin, Andreas
2008-01-01
It has long been known that there is a sustained electrical potential (voltage) difference between the xylem of many plants and their surrounding soil, but the mechanism behind this voltage has remained controversial. After eliminating any extraneous capacitive or inductive couplings and ground-mediated electric current flows, we have measured sustained differences of 50–200 mV between the xylem region of a Faraday-caged, intact, potted Ficus benjamina tree and its soil, as well as between its cut branches and soils and ionic solutions standardized to various pH values. Using identical platinum electrodes, no correlation between the voltage and time of day, illumination, sap flow, electrode elevation, or ionic composition of soil was found, suggesting no direct connection to simple dissimilar-metal redox reactions or transpirational activity. Instead, a clear relationship between the voltage polarity and magnitude and the pH difference between xylem and soil was observed. We attribute these sustained voltages to a biological concentration cell likely set up by the homeostatic mechanisms of the tree. Potential applications of this finding are briefly explored. PMID:18698415
Neghliz, Hayet; Cochard, Hervé; Brunel, Nicole; Martre, Pierre
2016-01-01
Seed dehydration is the normal terminal event in the development of orthodox seeds and is physiologically related to the cessation of grain dry mass accumulation and crop grain yield. For a better understanding of grain dehydration, we evaluated the hypothesis that hydraulic conductance of the ear decreases during the latter stages of development and that this decrease results from disruption or occlusion of xylem conduits. Whole ear, rachis, and stem nodes hydraulic conductance and percentage loss of xylem conductivity were measured from flowering to harvest-ripeness on bread wheat (Triticum aestivum L.) cv. Récital grown under controlled environments. Flag leaf transpiration, stomatal conductance, chlorophyll content and grain and ear water potentials were also measured during grain development. We show that grain dehydration was not related with whole plant physiology and leaf senescence, but closely correlated with the hydraulic properties of the xylem conduits irrigating the grains. Indeed, there was a substantial decrease in rachis hydraulic conductance at the onset of the grain dehydration phase. This hydraulic impairment was not caused by the presence of air embolism in xylem conduits of the stem internodes or rachis but by the occlusion of the xylem lumens by polysaccharides (pectins and callose). Our results demonstrate that xylem hydraulics plays a key role during grain maturation. PMID:27446150
Reversible Leaf Xylem Collapse: A Potential “Circuit Breaker” against Cavitation1[OPEN
Zhang, Yong-Jiang; Rockwell, Fulton E.; Graham, Adam C.; Alexander, Teressa; Holbrook, N. Michele
2016-01-01
We report a novel form of xylem dysfunction in angiosperms: reversible collapse of the xylem conduits of the smallest vein orders that demarcate and intrusively irrigate the areoles of red oak (Quercus rubra) leaves. Cryo-scanning electron microscopy revealed gradual increases in collapse from approximately −2 MPa down to −3 MPa, saturating thereafter (to −4 MPa). Over this range, cavitation remained negligible in these veins. Imaging of rehydration experiments showed spatially variable recovery from collapse within 20 s and complete recovery after 2 min. More broadly, the patterns of deformation induced by desiccation in both mesophyll and xylem suggest that cell wall collapse is unlikely to depend solely on individual wall properties, as mechanical constraints imposed by neighbors appear to be important. From the perspective of equilibrium leaf water potentials, petioles, whose vessels extend into the major veins, showed a vulnerability to cavitation that overlapped in the water potential domain with both minor vein collapse and buckling (turgor loss) of the living cells. However, models of transpiration transients showed that minor vein collapse and mesophyll capacitance could effectively buffer major veins from cavitation over time scales relevant to the rectification of stomatal wrong-way responses. We suggest that, for angiosperms, whose subsidiary cells give up large volumes to allow large stomatal apertures at the cost of potentially large wrong-way responses, vein collapse could make an important contribution to these plants’ ability to transpire near the brink of cavitation-inducing water potentials. PMID:27733514
Vitor, Simone C; do Amarante, Luciano; Sodek, Ladaslav
2018-05-16
A substantial increase in malate in the xylem sap of soybean subjected to mineral N starvation originates mainly from aspartate, a prominent amino acid of the phloem. A substantial increase in xylem malate was found when non-nodulated soybean plants were transferred to a N-free medium. Nodulated plants growing in the absence of mineral N and, therefore, dependent on symbiotic N 2 fixation also contained elevated concentrations of malate in the xylem sap. When either nitrate or ammonium was supplied, malate concentrations in the xylem sap were low, both for nodulated and non-nodulated plants. Evidence was obtained that the elevated malate concentration of the xylem was derived from amino acids supplied by the phloem. Aspartate was a prominent component of the phloem sap amino acids and, therefore, a potential source of malate. Supplying the roots of intact plants with 13 C-aspartate revealed that malate of the xylem sap was readily labelled under N starvation. A hypothetical scheme is proposed whereby aspartate supplied by the phloem is metabolised in the roots and the products of this metabolism cycled back to the shoot. Under N starvation, aspartate metabolism is diverted from asparagine synthesis to supply N for the synthesis of other amino acids via transaminase activity. The by-product of aspartate transaminase activity, oxaloacetate, is transformed to malate and its export accounts for much of the elevated concentration of malate found in the xylem sap. This mechanism represents a new additional role for malate during mineral N starvation of soybean, beyond that of charge balance.
Zwieniecki, Maciej A; Melcher, Peter J; Ahrens, Eric T
2013-01-01
We report results of an analysis of embolism formation and subsequent refilling observed in stems of Acer rubrum L. using magnetic resonance imaging (MRI). MRI is one of the very few techniques that can provide direct non-destructive observations of the water content within opaque biological materials at a micrometer resolution. Thus, it has been used to determine temporal dynamics and water distributions within xylem tissue. In this study, we found good agreement between MRI measures of pixel brightness to assess xylem liquid water content and the percent loss in hydraulic conductivity (PLC) in response to water stress (P50 values of 2.51 and 2.70 for MRI and PLC, respectively). These data provide strong support that pixel brightness is well correlated to PLC and can be used as a proxy of PLC even when single vessels cannot be resolved on the image. Pressure induced embolism in moderately stressed plants resulted in initial drop of pixel brightness. This drop was followed by brightness gain over 100 min following pressure application suggesting that plants can restore water content in stem after induced embolism. This recovery was limited only to current-year wood ring; older wood did not show signs of recovery within the length of experiment (16 h). In vivo MRI observations of the xylem of moderately stressed (~-0.5 MPa) A. rubrum stems revealed evidence of a spontaneous embolism formation followed by rapid refilling (~30 min). Spontaneous (not induced) embolism formation was observed only once, despite over 60 h of continuous MRI observations made on several plants. Thus this observation provide evidence for the presence of naturally occurring embolism-refilling cycle in A. rubrum, but it is impossible to infer any conclusions in relation to its frequency in nature.
18O Spatial Patterns of Vein Xylem Water, Leaf Water, and Dry Matter in Cotton Leaves
Gan, Kim Suan; Wong, Suan Chin; Yong, Jean Wan Hong; Farquhar, Graham Douglas
2002-01-01
Three leaf water models (two-pool model, Péclet effect, and string-of-lakes) were assessed for their robustness in predicting leaf water enrichment and its spatial heterogeneity. This was achieved by studying the 18O spatial patterns of vein xylem water, leaf water, and dry matter in cotton (Gossypium hirsutum) leaves grown at different humidities using new experimental approaches. Vein xylem water was collected from intact transpiring cotton leaves by pressurizing the roots in a pressure chamber, whereas the isotopic content of leaf water was determined without extracting it from fresh leaves with the aid of a purpose-designed leaf punch. Our results indicate that veins have a significant degree of lateral exchange with highly enriched leaf water. Vein xylem water is thus slightly, but progressively enriched in the direction of water flow. Leaf water enrichment is dependent on the relative distances from major veins, with water from the marginal and intercostal regions more enriched and that next to veins and near the leaf base more depleted than the Craig-Gordon modeled enrichment of water at the sites of evaporation. The spatial pattern of leaf water enrichment varies with humidity, as expected from the string-of-lakes model. This pattern is also reflected in leaf dry matter. All three models are realistic, but none could fully account for all of the facets of leaf water enrichment. Our findings acknowledge the presence of capacitance in the ground tissues of vein ribs and highlight the essential need to incorporate Péclet effects into the string-of-lakes model when applying it to leaves. PMID:12376664
Zwieniecki, Maciej A.; Melcher, Peter J.; Ahrens, Eric T.
2013-01-01
We report results of an analysis of embolism formation and subsequent refilling observed in stems of Acer rubrum L. using magnetic resonance imaging (MRI). MRI is one of the very few techniques that can provide direct non-destructive observations of the water content within opaque biological materials at a micrometer resolution. Thus, it has been used to determine temporal dynamics and water distributions within xylem tissue. In this study, we found good agreement between MRI measures of pixel brightness to assess xylem liquid water content and the percent loss in hydraulic conductivity (PLC) in response to water stress (P50 values of 2.51 and 2.70 for MRI and PLC, respectively). These data provide strong support that pixel brightness is well correlated to PLC and can be used as a proxy of PLC even when single vessels cannot be resolved on the image. Pressure induced embolism in moderately stressed plants resulted in initial drop of pixel brightness. This drop was followed by brightness gain over 100 min following pressure application suggesting that plants can restore water content in stem after induced embolism. This recovery was limited only to current-year wood ring; older wood did not show signs of recovery within the length of experiment (16 h). In vivo MRI observations of the xylem of moderately stressed (~-0.5 MPa) A. rubrum stems revealed evidence of a spontaneous embolism formation followed by rapid refilling (~30 min). Spontaneous (not induced) embolism formation was observed only once, despite over 60 h of continuous MRI observations made on several plants. Thus this observation provide evidence for the presence of naturally occurring embolism-refilling cycle in A. rubrum, but it is impossible to infer any conclusions in relation to its frequency in nature. PMID:23885258
Li, Shan; Feifel, Marion; Karimi, Zohreh; Schuldt, Bernhard; Choat, Brendan; Jansen, Steven
2016-02-01
Establishing physiological thresholds to drought-induced mortality in a range of plant species is crucial in understanding how plants respond to severe drought. Here, five common European tree species were selected (Acer campestre L., Acer pseudoplatanus L., Carpinus betulus L., Corylus avellana L. and Fraxinus excelsior L.) to study their hydraulic thresholds to mortality. Photosynthetic parameters during desiccation and the recovery of leaf gas exchange after rewatering were measured. Stem vulnerability curves and leaf pressure-volume curves were investigated to understand the hydraulic coordination of stem and leaf tissue traits. Stem and root samples from well-watered and severely drought-stressed plants of two species were observed using transmission electron microscopy to visualize mortality of cambial cells. The lethal water potential (ψlethal) correlated with stem P99 (i.e., the xylem water potential at 99% loss of hydraulic conductivity, PLC). However, several plants that were stressed beyond the water potential at 100% PLC showed complete recovery during the next spring, which suggests that the ψlethal values were underestimated. Moreover, we observed a 1 : 1 relationship between the xylem water potential at the onset of embolism and stomatal closure, confirming hydraulic coordination between leaf and stem tissues. Finally, ultrastructural changes in the cytoplasm of cambium tissue and mortality of cambial cells are proposed to provide an alternative approach to investigate the point of no return associated with plant death. © The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.
Franco, A C; de Soyza, A G; Virginia, R A; Reynolds, J F; Whitford, W G
1994-03-01
Larrea tridentata is a xerophytic evergreen shrub, dominant in the arid regions of the southwestern United States. We examined relationships between gasexchange characteristics, plant and soil water relations, and growth responses of large versus small shrubs of L. tridentata over the course of a summer growing season in the Chihuahuan Desert of southern New Mexico, USA. The soil wetting front did not reach 0.6 m, and soils at depths of 0.6 and 0.9 m remained dry throughout the summer, suggesting that L. tridentata extracts water largely from soil near the surface. Surface soil layers (<0.3 m) were drier under large plants, but predawn xylem water potentials were similar for both plant sizes suggesting some access to deeper soil moisture reserves by large plants. Stem elongation rates were about 40% less in large, reproductively active shrubs than in small, reproductively inactive shrubs. Maximal net photosynthetic rates (P max ) occurred in early summer (21.3 μ mol m -2 s -1 ), when pre-dawn xylem water potential (XWP) reached ca. -1 MPa. Although both shrub sizes exhibited similar responses to environmental factors, small shrubs recovered faster from short-term drought, when pre-dawn XWP reached about -4.5 MPa and P max decreased to only ca. 20% of unstressed levels. Gas exchange measurements yielded a strong relationship between stomatal conductance and photosynthesis, and the relationship between leaf-to-air vapor pressure deficit and stomatal conductance was found to be influenced by pre-dawn XWP. Our results indicate that stomatal responses to water stress and vapor pressure deficit are important in determining rates of carbon gain and water loss in L. tridentata.
Domec, Jean-Christophe; Pruyn, Michele L
2008-10-01
Effects of trunk girdling on seasonal patterns of xylem water status, water transport and woody tissue metabolic properties were investigated in ponderosa pine (Pinus ponderosa Dougl. ex P. Laws.) trees. At the onset of summer, there was a sharp decrease in stomatal conductance (g(s)) in girdled trees followed by a full recovery after the first major rainfall in September. Eliminating the root as a carbohydrate sink by girdling induced a rapid reversible reduction in g(s). Respiratory potential (a laboratory measure of tissue-level respiration) increased above the girdle (branches and upper trunk) and decreased below the girdle (lower trunk and roots) relative to control trees during the growing season, but the effect was reversed after the first major rainfall. The increase in branch respiratory potential induced by girdling suggests that the decrease in g(s) was caused by the accumulation of carbohydrates above the girdle, which is consistent with an observed increase in leaf mass per area in the girdled trees. Trunk girdling did not affect native xylem embolism or xylem conductivity. Both treated and control trunks experienced loss of xylem conductivity ranging from 10% in spring to 30% in summer. Girdling reduced xylem growth and sapwood to leaf area ratio, which in turn reduced branch leaf specific conductivity (LSC). The girdling-induced reductions in g(s) and transpiration were associated with a decrease in leaf hydraulic conductance. Two years after girdling, when root-to-shoot phloem continuity had been restored, girdled trees had a reduced density of new wood, which increased xylem conductivity and whole-tree LSC, but also vulnerability to embolism.
Joshi, Ankur; Knipfer, Thorsten; Steudle, Ernst
2009-11-01
In standard techniques (root pressure probe or high-pressure flowmeter), the hydraulic conductivity of roots is calculated from transients of root pressure using responses following step changes in volume or pressure, which may be affected by a storage of water in the stele. Storage effects were examined using both experimental data of root pressure relaxations and clamps and a physical capacity model. Young roots of corn and barley were treated as a three-compartment system, comprising a serial arrangement of xylem/probe, stele and outside medium/cortex. The hydraulic conductivities of the endodermis and of xylem vessels were derived from experimental data. The lower limit of the storage capacity of stelar tissue was caused by the compressibility of water. This was subsequently increased to account for realistic storage capacities of the stele. When root water storage was varied over up to five orders of magnitude, the results of simulations showed that storage effects could not explain the experimental data, suggesting a major contribution of effects other than water storage. It is concluded that initial water flows may be used to measure root hydraulic conductivity provided that the volumes of water used are much larger than the volumes stored.
Speirs, Jim; Binney, Allan; Collins, Marisa; Edwards, Everard; Loveys, Brian
2013-04-01
The influence of different levels of irrigation and of variation in atmospheric vapour pressure deficit (VPD) on the synthesis, metabolism, and transport of abscisic acid (ABA) and the effects on stomatal conductance were examined in field-grown Cabernet Sauvignon grapevines. Xylem sap, leaf tissue, and root tissue were collected at regular intervals during two seasons in conjunction with measurements of leaf water potential (Ψleaf) and stomatal conductance (gs). The different irrigation levels significantly altered the Ψleaf and gs of the vines across both seasons. ABA abundance in the xylem sap was correlated with gs. The expression of genes associated with ABA synthesis, NCED1 and NCED2, was higher in the roots than in the leaves throughout and highest in the roots in mid January, a time when soil moisture declined and VPD was at its highest. Their expression in roots was also inversely related to the levels of irrigation and correlated with ABA abundance in the roots, xylem sap, and leaves. Three genes encoding ABA 8'-hydroxylases were isolated and their identities confirmed by expression in yeast cells. The expression of one of these, Hyd1, was elevated in leaves when VPD was below 2.0-2.5 kPa and minimal at higher VPD levels. The results provide evidence that ABA plays an important role in linking stomatal response to soil moisture status and that changes in ABA catabolism at or near its site of action allows optimization of gas exchange to current environmental conditions.
Primary events regulating stem growth at low water potentials.
Nonami, H; Boyer, J S
1990-08-01
Cell enlargement is inhibited by inadequate water. As a first step toward understanding the mechanism, all the physical parameters affecting enlargement were monitored to identify those that changed first, particularly in coincidence with the inhibition. The osmotic potential, turgor, yield threshold turgor, growth-induced water potential, wall extensibility, and conductance to water were measured in the elongating region, and the water potential was measured in the xylem of stems of dark-grown soybean (Glycine max [L.] Merr.) seedlings. A stepdown in water potential was achieved around the roots by transplanting the seedlings to vermiculite of low water content, and each of the parameters was measured simultaneously in the same plants while intact or within a few minutes of being intact using a newly developed guillotine psychrometer. The gradient of decreasing water potential from the xylem to the enlarging cells (growth-induced water potential) was the first of the parameters to decrease to a growth-limiting level. The kinetics were the same as for the inhibition of growth. The decreased gradient was caused mostly by a decreased water potential of the xylem. This was followed after 5 to 10 hours by a similar decrease in cell wall extensibility and tissue conductance for water. Later, the growth-induced water potential recovered as a result of osmotic adjustment and a rise in the water potential of the xylem. Still later, moderate growth resumed at a rate apparently determined by the low wall extensibility and tissue conductance for water. The turgor did not change significantly during the experiment. These results indicate that the primary event during the growth inhibition was the change in the growth-induced water potential. Because the growth limitation subsequently shifted to the low wall extensibility and tissue conductance for water, the initial change in potential may have set in motion subsequent metabolic changes that altered the characteristics of the wall and cell membranes.
Primary Events Regulating Stem Growth at Low Water Potentials 1
Nonami, Hiroshi; Boyer, John S.
1990-01-01
Cell enlargement is inhibited by inadequate water. As a first step toward understanding the mechanism, all the physical parameters affecting enlargement were monitored to identify those that changed first, particularly in coincidence with the inhibition. The osmotic potential, turgor, yield threshold turgor, growth-induced water potential, wall extensibility, and conductance to water were measured in the elongating region, and the water potential was measured in the xylem of stems of dark-grown soybean (Glycine max [L.] Merr.) seedlings. A stepdown in water potential was achieved around the roots by transplanting the seedlings to vermiculite of low water content, and each of the parameters was measured simultaneously in the same plants while intact or within a few minutes of being intact using a newly developed guillotine psychrometer. The gradient of decreasing water potential from the xylem to the enlarging cells (growth-induced water potential) was the first of the parameters to decrease to a growth-limiting level. The kinetics were the same as for the inhibition of growth. The decreased gradient was caused mostly by a decreased water potential of the xylem. This was followed after 5 to 10 hours by a similar decrease in cell wall extensibility and tissue conductance for water. Later, the growth-induced water potential recovered as a result of osmotic adjustment and a rise in the water potential of the xylem. Still later, moderate growth resumed at a rate apparently determined by the low wall extensibility and tissue conductance for water. The turgor did not change significantly during the experiment. These results indicate that the primary event during the growth inhibition was the change in the growth-induced water potential. Because the growth limitation subsequently shifted to the low wall extensibility and tissue conductance for water, the initial change in potential may have set in motion subsequent metabolic changes that altered the characteristics of the wall and cell membranes. PMID:16667663
USDA-ARS?s Scientific Manuscript database
In phytophagous hemipterans, salivary sheath-feeding species are described as xylem or phloem sap-ingesting specialists. Because these two food sources have different characteristics, two different feeding tactics are associated with this supposed specialization. Study of feeding behavior is crucial...
Water relations of Calycanthus flowers: Hydraulic conductance, capacitance, and embolism resistance.
Roddy, Adam B; Simonin, Kevin A; McCulloh, Katherine A; Brodersen, Craig R; Dawson, Todd E
2018-03-30
For most angiosperms, producing and maintaining flowers is critical to sexual reproduction, yet little is known about the physiological processes involved in maintaining flowers throughout anthesis. Among extant species, flowers of the genus Calycanthus have the highest hydraulic conductance and vein densities of species measured to date, yet they can wilt by late morning under hot conditions. Here, we combine diurnal measurements of gas exchange and water potential, pressure-volume relations, functional responses of gas exchange, and characterization of embolism formation using high resolution X-ray computed microtomography to determine drought responses of Calycanthus flowers. Transpiration from flowers frequently exceeded transpiration from leaves, and flowers were unable to limit transpiration under conditions of high vapour pressure deficit. As a result, they rely heavily on hydraulic capacitance to prevent water potential declines. Despite having high water potentials at turgor loss, flowers were very resistant to embolism formation, with no embolism apparent until tepal water potentials had declined to -2 MPa. Although Calycanthus flowers remain connected to the stem xylem and have high hydraulic capacitance, their inability to curtail transpiration leads to turgor loss. These results suggest that extreme climate events may cause flower failure, potentially preventing successful reproduction. © 2018 John Wiley & Sons Ltd.
A novel approach for diagnosing isohydric and anisohydric plant water use during drought
NASA Astrophysics Data System (ADS)
Novick, K. A.; Roman, D. T.; Brzostek, E. R.; Dragoni, D.; Phillips, R.
2014-12-01
Recent years have seen the emergence of a new framework for describing plant water use, whereby species-specific water use strategies during periods of hydrologic stress are classified as falling on a spectrum of isohydric to anisohydric behavior. Trees that regulate water potential to within a relatively narrow range, and thereby reduce the risk of damaging xylem cavitation, are categorized as isohydric. In contrast, anisohydric trees allow their leaf water potential to decrease during drought, which may improve gas exchange rates, but at the cost of a greater risk of cavitation in the xylem. To date, most of the approaches to diagnose and characterize isohydric as compared to anisohydric behavior rely on observations of stem or leaf water potential measurements, which are difficult to collect at a high temporal and spatial frequency and rely on destructive techniques. Here, we use cohesion-tension theory to develop a novel approach for diagnosing isohydric/anisohydric behavior in observations of leaf- or canopy-scale stomatal conductance, which are data that may be collected in situ and with relative ease. The approach is particularly focused on exploring how the relationship between stomatal conductance and vapor pressure deficit changes during dry-down periods. The theoretical predictions suggest that the sensitivity of stomatal conductance to vapor pressure deficit may decrease over the course of the drought event for more anisohydric trees, and increase in the case of more isohydric trees. Species-specific, leaf-level observations of the relevant variables collected during the course of a severe drought event affecting the Morgan-Monroe State Forest in 2012 are shown to confirm the theoretical predictions. Finally, the diagnostic approach is evaluated in the context of other emerging approaches for describing stomatal behavior, including the growing recognition of the role of hydraulic capacitance during drought, and recent advances in stomatal optimization theory. Ultimately, placing species along the isohydric-anisohydric contiuum may advance our understanding of the magnitude of drought-related declines in productivity and other physiological processes in forest ecosystems.
Active water transport in unicellular algae: where, why, and how.
Raven, John A; Doblin, Martina A
2014-12-01
The occurrence of active water transport (net transport against a free energy gradient) in photosynthetic organisms has been debated for several decades. Here, active water transport is considered in terms of its roles, where it is found, and the mechanisms by which it could occur. First there is a brief consideration of the possibility of active water transport into plant xylem in the generation of root pressure and the refilling of embolized xylem elements, and from an unsaturated atmosphere into terrestrial organisms living in habitats with limited availability of liquid water. There is then a more detailed consideration of volume and osmotic regulation in wall-less freshwater unicells, and the possibility of generation of buoyancy in marine phytoplankton such as large-celled diatoms. Calculations show that active water transport is a plausible mechanism to assist cells in upwards vertical movements, requires less energy than synthesis of low-density organic solutes, and potentially on a par with excluding certain ions from the vacuole. © The Author 2014. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.
Comparative genomics reveals conservative evolution of the xylem transcriptome in vascular plants.
Li, Xinguo; Wu, Harry X; Southerton, Simon G
2010-06-21
Wood is a valuable natural resource and a major carbon sink. Wood formation is an important developmental process in vascular plants which played a crucial role in plant evolution. Although genes involved in xylem formation have been investigated, the molecular mechanisms of xylem evolution are not well understood. We use comparative genomics to examine evolution of the xylem transcriptome to gain insights into xylem evolution. The xylem transcriptome is highly conserved in conifers, but considerably divergent in angiosperms. The functional domains of genes in the xylem transcriptome are moderately to highly conserved in vascular plants, suggesting the existence of a common ancestral xylem transcriptome. Compared to the total transcriptome derived from a range of tissues, the xylem transcriptome is relatively conserved in vascular plants. Of the xylem transcriptome, cell wall genes, ancestral xylem genes, known proteins and transcription factors are relatively more conserved in vascular plants. A total of 527 putative xylem orthologs were identified, which are unevenly distributed across the Arabidopsis chromosomes with eight hot spots observed. Phylogenetic analysis revealed that evolution of the xylem transcriptome has paralleled plant evolution. We also identified 274 conifer-specific xylem unigenes, all of which are of unknown function. These xylem orthologs and conifer-specific unigenes are likely to have played a crucial role in xylem evolution. Conifers have highly conserved xylem transcriptomes, while angiosperm xylem transcriptomes are relatively diversified. Vascular plants share a common ancestral xylem transcriptome. The xylem transcriptomes of vascular plants are more conserved than the total transcriptomes. Evolution of the xylem transcriptome has largely followed the trend of plant evolution.
Comparative genomics reveals conservative evolution of the xylem transcriptome in vascular plants
2010-01-01
Background Wood is a valuable natural resource and a major carbon sink. Wood formation is an important developmental process in vascular plants which played a crucial role in plant evolution. Although genes involved in xylem formation have been investigated, the molecular mechanisms of xylem evolution are not well understood. We use comparative genomics to examine evolution of the xylem transcriptome to gain insights into xylem evolution. Results The xylem transcriptome is highly conserved in conifers, but considerably divergent in angiosperms. The functional domains of genes in the xylem transcriptome are moderately to highly conserved in vascular plants, suggesting the existence of a common ancestral xylem transcriptome. Compared to the total transcriptome derived from a range of tissues, the xylem transcriptome is relatively conserved in vascular plants. Of the xylem transcriptome, cell wall genes, ancestral xylem genes, known proteins and transcription factors are relatively more conserved in vascular plants. A total of 527 putative xylem orthologs were identified, which are unevenly distributed across the Arabidopsis chromosomes with eight hot spots observed. Phylogenetic analysis revealed that evolution of the xylem transcriptome has paralleled plant evolution. We also identified 274 conifer-specific xylem unigenes, all of which are of unknown function. These xylem orthologs and conifer-specific unigenes are likely to have played a crucial role in xylem evolution. Conclusions Conifers have highly conserved xylem transcriptomes, while angiosperm xylem transcriptomes are relatively diversified. Vascular plants share a common ancestral xylem transcriptome. The xylem transcriptomes of vascular plants are more conserved than the total transcriptomes. Evolution of the xylem transcriptome has largely followed the trend of plant evolution. PMID:20565927
Rita, Angelo; Cherubini, Paolo; Leonardi, Stefano; Todaro, Luigi; Borghetti, Marco
2015-08-01
The present study assessed the effects of climatic conditions on radial growth and functional anatomical traits, including ring width, vessel size, vessel frequency and derived variables, i.e., potential hydraulic conductivity and xylem vulnerability to cavitation in Ilex aquifolium L. trees using long-term tree-ring time series obtained at two climatically contrasting sites, one mesic site in Switzerland (CH) and one drought-prone site in Italy (ITA). Relationships were explored by examining different xylem traits, and point pattern analysis was applied to investigate vessel clustering. We also used generalized additive models and bootstrap correlation functions to describe temperature and precipitation effects. Results indicated modified radial growth and xylem anatomy in trees over the last century; in particular, vessel frequency increased markedly at both sites in recent years, and all xylem traits examined, with the exception of xylem cavitation vulnerability, were higher at the CH mesic compared with the ITA drought site. A significant vessel clustering was observed at the ITA site, which could contribute to an enhanced tolerance to drought-induced embolism. Flat and negative relationships between vessel size and ring width were observed, suggesting carbon was not allocated to radial growth under conditions which favored stem water conduction. Finally, in most cases results indicated that climatic conditions influenced functional anatomical traits more substantially than tree radial growth, suggesting a crucial role of functional xylem anatomy in plant acclimation to future climatic conditions. © The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.
J-C. Domec; F.G. Scholz; S.J. Bucci; F.C. Meinzer; G. Goldstein; R. Villalobos-Vega
2006-01-01
Vulnerability to water-stress-induced embolism and variation in the degree of native embolism were measured in lateral roots of four co-occuring neotropical savanna tree species. Root embolism varied diurnally and seasonally. Late in the dry season, loss of root xylem conductivity reached 80% in the afternoon when root water potential (ψroot...
Ratnayake, Kamani; Joyce, Daryl C; Webb, Richard I
2013-08-01
Maintaining a high rate of water uptake is crucial for maximum longevity of cut stems. Physiological gel/tylosis formation decreases water transport efficiency in the xylem. The primary mechanism of action for post-harvest Cu(2+) treatments in improving cut flower and foliage longevity has been elusive. The effect of Cu(2+) on wound-induced xylem vessel occlusion was investigated for Acacia holosericea A. Cunn. ex G. Don. Experiments were conducted using a Cu(2+) pulse (5 h, 2.2 mM) and a Cu(2+) vase solution (0.5 mM) vs a deionized water (DIW) control. Development of xylem blockage in the stem-end region 10 mm proximal to the wounded stem surface was examined over 21 days by light and transmission electron microscopy. Xylem vessels of stems stood into DIW were occluded with gels secreted into vessel lumens via pits from surrounding axial parenchyma cells. Gel secretion was initiated within 1-2 days post-wounding and gels were detected in the xylem from day 3. In contrast, Cu(2+) treatments disrupted the surrounding parenchyma cells, thereby inhibiting gel secretion and maintaining the vessel lumens devoid of occlusions. The Cu(2+) treatments significantly improved water uptake by the cut stems as compared to the control. © 2013 Scandinavian Plant Physiology Society.
Sevanto, Sanna Annika; Ryan, Max; Turin Dickman, L.; ...
2018-03-22
Plants close their stomata during drought to avoid excessive water loss, but species differ in respect to the drought severity at which stomata close. The stomatal closure point is related to xylem anatomy and vulnerability to embolism, but it also has implications for phloem transport, and possibly phloem anatomy to allow sugar transport at low water potentials. Desiccation tolerant plants that close their stomata at severe drought should have smaller xylem conduits and/or fewer and smaller inter-conduit pits to reduce vulnerability to embolism, but more phloem tissue and larger phloem conduits compared to plants that avoid desiccation. These anatomical differencesmore » could be expected to increase in response to long-term reduction in precipitation. To test these hypotheses we used tridimensional synchroton X-ray microtomograph and light microscope imaging of combined xylem and phloem tissues of two coniferous species: one-seed juniper (Juniperus monosperma) and piñon pine (Pinus edulis) subjected to precipitation manipulation treatments. These species show different xylem vulnerability to embolism, contrasting desiccation tolerance, and stomatal closure points. Our results support the hypothesis that desiccation tolerant plants require higher phloem transport capacity than desiccation avoiding plants, but this can be gained through various anatomical adaptations in addition to changing conduit or tissue size.« less
DOE Office of Scientific and Technical Information (OSTI.GOV)
Sevanto, Sanna Annika; Ryan, Max; Turin Dickman, L.
Plants close their stomata during drought to avoid excessive water loss, but species differ in respect to the drought severity at which stomata close. The stomatal closure point is related to xylem anatomy and vulnerability to embolism, but it also has implications for phloem transport, and possibly phloem anatomy to allow sugar transport at low water potentials. Desiccation tolerant plants that close their stomata at severe drought should have smaller xylem conduits and/or fewer and smaller inter-conduit pits to reduce vulnerability to embolism, but more phloem tissue and larger phloem conduits compared to plants that avoid desiccation. These anatomical differencesmore » could be expected to increase in response to long-term reduction in precipitation. To test these hypotheses we used tridimensional synchroton X-ray microtomograph and light microscope imaging of combined xylem and phloem tissues of two coniferous species: one-seed juniper (Juniperus monosperma) and piñon pine (Pinus edulis) subjected to precipitation manipulation treatments. These species show different xylem vulnerability to embolism, contrasting desiccation tolerance, and stomatal closure points. Our results support the hypothesis that desiccation tolerant plants require higher phloem transport capacity than desiccation avoiding plants, but this can be gained through various anatomical adaptations in addition to changing conduit or tissue size.« less
Dynamic relationship between the VOC emissions from a Scots pine stem and the tree water relations
NASA Astrophysics Data System (ADS)
Vanhatalo, Anni; Chan, Tommy; Aalto, Juho; Kolari, Pasi; Rissanen, Kaisa; Hakola, Hannele; Hölttä, Teemu; Bäck, Jaana
2013-04-01
The stems of coniferous trees contain huge storages of oleoresin. The composition of oleoresin depends on e.g. tree species, age, provenance, health status, and environmental conditions. Oleoresin is under pressure in the extensive network of resin ducts in wood and needles. It flows out from a mechanically damaged site to protect the tree by sealing the wounded site. Once in contact with air, volatile parts of oleoresin evaporate, and the residual compounds harden to make a solid protective seal over damaged tissues. The hardening time of the resin depends on evaporation rate of the volatiles which in turn depends on temperature. The storage is also toxic to herbivores and attracts predators that restrict the herbivore damage. Despite abundant knowledge on emissions of volatile isoprenoids from foliage, very little is known about their emissions from woody plant parts. We set up an experiment to measure emissions of isoprene and monoterpenes as well as two oxygenated VOCs, methanol and acetone, from a Scots pine (Pinus sylvestris) stem and branches. The measurements were started in early April and continued until mid-June, 2012. Simultaneously, we measured the dynamics of whole stem and xylem diameter changes, stem sap flow rate and foliage transpiration rate. These measurements were used to estimate A) pressure changes inside the living stem tissue and the water conducting xylem, B) the refilling of stem water stores after winter dehydration (the ratio of sap flow at the stem base to water loss by foliage), and C) the increase in tree water transport capacity (the ratio of maximum daily sap flow rate to the diurnal variation in xylem pressure) during spring due to winter embolism refilling and/or the temperature dependent root water uptake capacity. The results show that already very early in spring, significant VOC emissions from pine stem can be detected, and that they exhibit a diurnal cycle similar to that of ambient temperature. During the highest emission period a sudden decrease in stem diameter was observed, which we hypothesize could either indicate a decrease in the pressure of living cells in connection with stem VOC emissions, or result mechanically from exudation of oleoresin from the stem. We also found that the stem water stores and xylem water transport capacity increased during periods of VOC emissions, which indicates xylem embolism refilling during times of VOC emissions. A qualitative difference was found between VOC emissions from pine stem and thick branches, the stem emissions containing more sesquiterpenes. Most of the tree biomass is in the lower part of the stem, and as stem tissues are lacking green photosynthesizing tissue the emissions are supposed to be related to damage. Our results show that emissions from tree stems are connected to the tree water relations and that they are important during the period when the foliage still is rather inactive.
Cavitation Events in Thuja occidentalis L.? 1
Tyree, Melvin T.; Dixon, Michael A.
1983-01-01
Ultrasonic acoustic emissions (AE) in the frequency range of 0.1 to 1 megahertz appear to originate in the sapwood of Thuja occidentalis L. The AE are vibrations of an impulsive nature. The vibrations can be transduced to a voltage waveform and amplified. The vibrations of each AE event begin at a large amplitude which decays over 20 to 100 microseconds. Strong circumstantial evidence indicates that the ultrasonic AE result from cavitation events because: (a) they occur only when the xylem pressure potential Ψxp is more negative than a threshold level of about —1 megapascal; (b) the rate of AE events increases as Ψxp decreases and when the net rate of water loss increases; (c) the AE can be stopped by raising Ψxp above —1 megapascal. Ultrasonic AE have been measured in whole terminal shoots allowed to dry in the laboratory, in isolated pieces of sapwood as they dried in the laboratory, and in whole terminal shoots in a pressure bomb when Ψxp was decreased by lowering the gas pressure in the pressure bomb. PMID:16663126
Hydraulic Function in Australian Tree Species during Drought-Induced Mortality
NASA Astrophysics Data System (ADS)
Tissue, D.; Maier, C.; Creek, D.; Choat, B.
2016-12-01
Drought induced tree mortality and decline are key issues facing forest ecology and management. Here, we primarily investigated the hydraulic limitations underpinning drought-induced mortality in three Australian tree species. Using field-based large rainout shelters, three angiosperm species (Casuarina cunninghamiana, Eucalyptus sideroxylon, Eucalyptus tereticornis) were subjected to two successive drought and recovery cycles, prior to a subsequent long and extreme drought to mortality; total duration of experiment was 2.5 years. Leaf gas exchange, leaf and stem hydraulics, and carbon reserves were monitored during the experiment. Trees died as a result of failure in the hydraulic transport system, primarily related to water stress induced embolism. Stomatal closure occurred prior to the induction of significant embolism in the stem xylem of all species. Nonetheless, trees suffered a rapid decline in xylem water potential and increase in embolism during the severe drought treatment. Trees died at water potentials causing greater than 90% loss of hydraulic conductivity in the stem, providing support for the theory that lethal water potential is correlated with complete loss of hydraulic function in the stem xylem of angiosperms.
Root resistance to cavitation is accurately measured using a centrifuge technique.
Pratt, R B; MacKinnon, E D; Venturas, M D; Crous, C J; Jacobsen, A L
2015-02-01
Plants transport water under negative pressure and this makes their xylem vulnerable to cavitation. Among plant organs, root xylem is often highly vulnerable to cavitation due to water stress. The use of centrifuge methods to study organs, such as roots, that have long vessels are hypothesized to produce erroneous estimates of cavitation resistance due to the presence of open vessels through measured samples. The assumption that roots have long vessels may be premature since data for root vessel length are sparse; moreover, recent studies have not supported the existence of a long-vessel artifact for stems when a standard centrifuge technique was used. We examined resistance to cavitation estimated using a standard centrifuge technique and compared these values with native embolism measurements for roots of seven woody species grown in a common garden. For one species we also measured vulnerability using single-vessel air injection. We found excellent agreement between root native embolism and the levels of embolism measured using a centrifuge technique, and with air-seeding estimates from single-vessel injection. Estimates of cavitation resistance measured from centrifuge curves were biologically meaningful and were correlated with field minimum water potentials, vessel diameter (VD), maximum xylem-specific conductivity (Ksmax) and vessel length. Roots did not have unusually long vessels compared with stems; moreover, root vessel length was not correlated to VD or to the vessel length of stems. These results suggest that root cavitation resistance can be accurately and efficiently measured using a standard centrifuge method and that roots are highly vulnerable to cavitation. The role of root cavitation resistance in determining drought tolerance of woody species deserves further study, particularly in the context of climate change. © The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.
Speirs, Jim; Binney, Allan; Collins, Marisa; Edwards, Everard; Loveys, Brian
2013-01-01
The influence of different levels of irrigation and of variation in atmospheric vapour pressure deficit (VPD) on the synthesis, metabolism, and transport of abscisic acid (ABA) and the effects on stomatal conductance were examined in field-grown Cabernet Sauvignon grapevines. Xylem sap, leaf tissue, and root tissue were collected at regular intervals during two seasons in conjunction with measurements of leaf water potential (Ψleaf) and stomatal conductance (gs). The different irrigation levels significantly altered the Ψleaf and gs of the vines across both seasons. ABA abundance in the xylem sap was correlated with gs. The expression of genes associated with ABA synthesis, NCED1 and NCED2, was higher in the roots than in the leaves throughout and highest in the roots in mid January, a time when soil moisture declined and VPD was at its highest. Their expression in roots was also inversely related to the levels of irrigation and correlated with ABA abundance in the roots, xylem sap, and leaves. Three genes encoding ABA 8’-hydroxylases were isolated and their identities confirmed by expression in yeast cells. The expression of one of these, Hyd1, was elevated in leaves when VPD was below 2.0–2.5 kPa and minimal at higher VPD levels. The results provide evidence that ABA plays an important role in linking stomatal response to soil moisture status and that changes in ABA catabolism at or near its site of action allows optimization of gas exchange to current environmental conditions. PMID:23630325
Optimal stomatal behavior with competition for water and risk of hydraulic impairment.
Wolf, Adam; Anderegg, William R L; Pacala, Stephen W
2016-11-15
For over 40 y the dominant theory of stomatal behavior has been that plants should open stomates until the carbon gained by an infinitesimal additional opening balances the additional water lost times a water price that is constant at least over short periods. This theory has persisted because of its remarkable success in explaining strongly supported simple empirical models of stomatal conductance, even though we have also known for over 40 y that the theory is not consistent with competition among plants for water. We develop an alternative theory in which plants maximize carbon gain without pricing water loss and also add two features to both this and the classical theory, which are strongly supported by empirical evidence: (i) water flow through xylem that is progressively impaired as xylem water potential drops and (ii) fitness or carbon costs associated with low water potentials caused by a variety of mechanisms, including xylem damage repair. We show that our alternative carbon-maximization optimization is consistent with plant competition because it yields an evolutionary stable strategy (ESS)-species with the ESS stomatal behavior that will outcompete all others. We further show that, like the classical theory, the alternative theory also explains the functional forms of empirical stomatal models. We derive ways to test between the alternative optimization criteria by introducing a metric-the marginal xylem tension efficiency, which quantifies the amount of photosynthesis a plant will forego from opening stomatal an infinitesimal amount more to avoid a drop in water potential.
NASA Astrophysics Data System (ADS)
Deng, Zijuan; Guan, Huade; Hutson, John; Forster, Michael A.; Wang, Yunquan; Simmons, Craig T.
2017-06-01
A novel simple soil-plant-atmospheric continuum model that emphasizes the vegetation's role in controlling water transfer (v-SPAC) has been developed in this study. The v-SPAC model aims to incorporate both plant and soil hydrological measurements into plant water transfer modeling. The model is different from previous SPAC models in which v-SPAC uses (1) a dynamic plant resistance system in the form of a vulnerability curve that can be easily obtained from sap flow and stem xylem water potential time series and (2) a plant capacitance parameter to buffer the effects of transpiration on root water uptake. The unique representation of root resistance and capacitance allows the model to embrace SPAC hydraulic pathway from bulk soil, to soil-root interface, to root xylem, and finally to stem xylem where the xylem water potential is measured. The v-SPAC model was tested on a native tree species in Australia, Eucalyptus crenulata saplings, with controlled drought treatment. To further validate the robustness of the v-SPAC model, it was compared against a soil-focused SPAC model, LEACHM. The v-SPAC model simulation results closely matched the observed sap flow and stem water potential time series, as well as the soil moisture variation of the experiment. The v-SPAC model was found to be more accurate in predicting measured data than the LEACHM model, underscoring the importance of incorporating root resistance into SPAC models and the benefit of integrating plant measurements to constrain SPAC modeling.
Else, Mark A; Taylor, June M; Atkinson, Christopher J
2006-01-01
In flooded soils, the rapid effects of decreasing oxygen availability on root metabolic activity are likely to generate many potential chemical signals that may impact on stomatal apertures. Detached leaf transpiration tests showed that filtered xylem sap, collected at realistic flow rates from plants flooded for 2 h and 4 h, contained one or more factors that reduced stomatal apertures. The closure could not be attributed to increased root output of the glucose ester of abscisic acid (ABA-GE), since concentrations and deliveries of ABA conjugates were unaffected by soil flooding. Although xylem sap collected from the shoot base of detopped flooded plants became more alkaline within 2 h of flooding, this rapid pH change of 0.5 units did not alter partitioning of root-sourced ABA sufficiently to prompt a transient increase in xylem ABA delivery. More shoot-sourced ABA was detected in the xylem when excised petiole sections were perfused with pH 7 buffer, compared with pH 6 buffer. Sap collected from the fifth oldest leaf of "intact" well-drained plants and plants flooded for 3 h was more alkaline, by approximately 0.4 pH units, than sap collected from the shoot base. Accordingly, xylem [ABA] was increased 2-fold in sap collected from the fifth oldest petiole compared with the shoot base of flooded plants. However, water loss from transpiring, detached leaves was not reduced when the pH of the feeding solution containing 3-h-flooded [ABA] was increased from 6.7 to 7.1 Thus, the extent of the pH-mediated, shoot-sourced ABA redistribution was not sufficient to raise xylem [ABA] to physiologically active levels. Using a detached epidermis bioassay, significant non-ABA anti-transpirant activity was also detected in xylem sap collected at intervals during the first 24 h of soil flooding.
Tao, Qi; Jupa, Radek; Luo, Jipeng; Lux, Alexander; Kováč, Ján; Wen, Yue; Zhou, Yimei; Jan, Japenga; Liang, Yongchao
2017-01-01
Abstract Although the significance of apoplasmic barriers in roots with regards to the uptake of toxic elements is generally known, the contribution of apoplasmic bypasses (ABs) to cadmium (Cd) hyperaccumulation is little understood. Here, we employed a combination of stable isotopic tracer techniques, an ABs tracer, hydraulic measurements, suberin lamellae staining, metabolic inhibitors, and antitranspirants to investigate and quantify the impact of the ABs on translocation of Cd to the xylem in roots of a hyperaccumulating (H) ecotype and a non-hyperaccumulating (NH) ecotype of Sedum alfredii. In the H ecotype, the Cd content in the xylem sap was proportional to hydrostatic pressure, which was attributed to pressure-driven flow via the ABs. The contribution of the ABs to Cd transportation to the xylem was dependent on the Cd concentration applied to the H ecotype (up to 37% at the highest concentration used). Cd-treated H ecotype roots showed significantly higher hydraulic conductance compared with the NH ecotype (76 vs 52 × 10–8 m s–1MPa–1), which is in accordance with less extensive suberization due to reduced expression of suberin-related genes. The main entry sites of apoplasmically transported Cd were localized in the root apexes and lateral roots of the H ecotype, where suberin lamellae were not well developed. These findings highlight the significance of the apoplasmic bypass in Cd hyperaccumulation in hyperaccumulating ecotypes of S. alfredii. PMID:28204505
New insights into the mechanisms of water-stress-induced cavitation in conifers.
Cochard, Hervé; Hölttä, Teemu; Herbette, Stéphane; Delzon, Sylvain; Mencuccini, Maurizio
2009-10-01
Cavitation resistance is a key parameter to understand tree drought tolerance but little is known about the mechanisms of air entry into xylem conduits. For conifers three mechanisms have been proposed: (1) a rupture of pit margo microfibrils, (2) a displacement of the pit torus from its normal sealing position over the pit aperture, and (3) a rupture of an air-water menisci in a pore of the pit margo. In this article, we report experimental results on three coniferous species suggesting additional mechanisms. First, when xylem segments were injected with a fluid at a pressure sufficient to aspirate pit tori and well above the pressure for cavitation induction we failed to detect the increase in sample conductance that should have been caused by torus displacement from blocking the pit aperture or by membrane rupture. Second, by injecting xylem samples with different surfactant solutions, we found a linear relation between sample vulnerability to cavitation and fluid surface tension. This suggests that cavitation in conifers could also be provoked by the capillary failure of an air-water meniscus in coherence with the prediction of Young-Laplace's equation. Within the bordered pit membrane, the exact position of this capillary seeding is unknown. The possible Achilles' heel could be the seal between tori and pit walls or holes in the torus. The mechanism of water-stress-induced cavitation in conifers could then be relatively similar to the one currently proposed for angiosperms.
Uptake of water via branches helps timberline conifers refill embolized xylem in late winter.
Mayr, Stefan; Schmid, Peter; Laur, Joan; Rosner, Sabine; Charra-Vaskou, Katline; Dämon, Birgit; Hacke, Uwe G
2014-04-01
Xylem embolism is a limiting factor for woody species worldwide. Conifers at the alpine timberline are exposed to drought and freeze-thaw stress during winter, which induce potentially lethal embolism. Previous studies indicated that timberline trees survive by xylem refilling. In this study on Picea abies, refilling was monitored during winter and spring seasons and analyzed in the laboratory and in situ experiments, based on hydraulic, anatomical, and histochemical methods. Refilling started in late winter, when the soil was frozen and soil water not available for the trees. Xylem embolism caused up to 86.2% ± 3.1% loss of conductivity and was correlated with the ratio of closed pits. Refilling of xylem as well as recovery in shoot conductance started in February and corresponded with starch accumulation in secondary phloem and in the mesophyll of needles, where we also observed increasing aquaporin densities in the phloem and endodermis. This indicates that active, cellular processes play a role for refilling even under winter conditions. As demonstrated by our experiments, water for refilling was thereby taken up via the branches, likely by foliar water uptake. Our results suggest that refilling is based on water shifts to embolized tracheids via intact xylem, phloem, and parenchyma, whereby aquaporins reduce resistances along the symplastic pathway and aspirated pits facilitate isolation of refilling tracheids. Refilling must be taken into account as a key process in plant hydraulics and in estimating future effects of climate change on forests and alpine tree ecosystems.
Borel, C; Audran, C; Frey, A; Marion-Poll, A; Tardieu, F; Simonneau, T
2001-03-01
A series of transgenic lines of Nicotiana plumbaginifolia with modified expression of zeaxanthin epoxidase gene (ZEP) provided contrasting ABA accumulation in roots and xylem sap. For mild water stress, concentration of ABA in the xylem sap ([ABA](xylem)) was clearly lower in plants underexpressing ZEP mRNA (complemented mutants and antisense transgenic lines) than in wild-type. In well-watered conditions, all lines presented similar [ABA](xylem) and similar ABA accumulation rates in detached roots. Plants could, therefore, be grown under normal light intensities and evaporative demand. Both ZEP mRNA abundance and ABA accumulation rate in roots increased with water deficit in all transgenic lines, except in complemented aba2-s1 mutants in which the ZEP gene was controlled by a constitutive promoter which does not respond to water deficit. These lines presented no change in root ABA content either with time or dehydration. The increase in ZEP mRNA abundance in roots with decreasing RWC was more pronounced in detached roots than in whole plants, suggesting a difference in mechanism. In all transgenic lines, a linear relationship was observed between predawn leaf water potential and [ABA](xylem), which could be reproduced in several experiments in the greenhouse and in the growth chamber. It is therefore possible to represent the effect of the transformation by a single parameter, thereby allowing the use of a quantitative approach to assist understanding of the behaviour of transgenic lines.
Tomasella, Martina; Beikircher, Barbara; Häberle, Karl-Heinz; Hesse, Benjamin; Kallenbach, Christian; Matyssek, Rainer; Mayr, Stefan
2018-02-01
Decreasing water availability due to climate change poses the question of whether and to what extent tree species are able to hydraulically acclimate and how hydraulic traits of stems and leaves are coordinated under drought. In a through-fall exclusion experiment, hydraulic acclimation was analyzed in a mixed forest stand of Fagus sylvatica L. and Picea abies (L.) Karst. In drought-stressed (TE, through-fall exclusion over 2 years) and control (CO) trees, hydraulic vulnerability was studied in branches as well as in leaves (F. sylvatica) and end-twigs (P. abies, entirely formed during the drought period) sampled at the same height in sun-exposed portions of the tree crown. In addition, relevant xylem anatomical traits and leaf pressure-volume relations were analyzed. The TE trees reached pre-dawn water potentials down to -1.6 MPa. In both species, water potentials at 50% loss of xylem hydraulic conductivity were ~0.4 MPa more negative in TE than in CO branches. Foliage hydraulic vulnerability (expressed as water potential at 50% loss of leaf/end-twig hydraulic conductance) and water potential at turgor loss point were also, respectively, 0.4 and 0.5 MPa lower in TE trees. Minor differences were observed in conduit mean hydraulic diameter and cell wall reinforcement. Our findings indicate significant and fast hydraulic acclimation under relatively mild drought in both tree species. Acclimation was well coordinated between branches and foliage, which might be essential for survival and productivity of mature trees under future drought periods. © The Author 2017. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.
Differences in delta13C and diameter growth among remnant Scots pine populations in Scotland.
Brendel, Oliver; Handley, Linda; Griffiths, Howard
2002-10-01
Published data suggest that differences in wood cellulose carbon isotope composition (delta13C) and xylem ring width among natural populations of Scots pine in Scotland (Pinus sylvestris L.) are attributable to the persistence of palaeotypes of various post-glacial migratory origins. We assessed differences in wood cellulose delta13C and ring width among Scottish Scots pine populations grown in a clone bank and in natural stands at various locations in northern and central Scotland. Ring width and wood cellulose delta13C varied significantly among natural stands. Potential water deficit was positively correlated with wood cellulose delta13C and xylem ring width in the natural stands. Neither wood cellulose delta13C nor xylem ring width of clone bank trees correlated with any climate variables at the sites from which the trees originated, indicating little adaptation to climate for these traits. Xylem ring width showed a site x population interaction for the growth sites (i.e., natural stands versus clone bank), but wood cellulose delta13C did not. These results suggest that climate variation in Scotland has not resulted in significant genetic variation in wood cellulose delta13C or xylem ring width in post-glacial populations.
Wang, Peng; Lee, Yunho; Igo, Michele M; Roper, M Caroline
2017-09-01
Bacterial plant pathogens often encounter reactive oxygen species (ROS) during host invasion. In foliar bacterial pathogens, multiple regulatory proteins are involved in the sensing of oxidative stress and the activation of the expression of antioxidant genes. However, it is unclear whether xylem-limited bacteria, such as Xylella fastidiosa, experience oxidative stress during the colonization of plants. Examination of the X. fastidiosa genome uncovered only one homologue of oxidative stress regulatory proteins, OxyR. Here, a knockout mutation in the X. fastidiosa oxyR gene was constructed; the resulting strain was significantly more sensitive to hydrogen peroxide (H 2 O 2 ) relative to the wild-type. In addition, during early stages of grapevine infection, the survival rate was 1000-fold lower for the oxyR mutant than for the wild-type. This supports the hypothesis that grapevine xylem represents an oxidative environment and that X. fastidiosa must overcome this challenge to achieve maximal xylem colonization. Finally, the oxyR mutant exhibited reduced surface attachment and cell-cell aggregation and was defective in biofilm maturation, suggesting that ROS could be a potential environmental cue stimulating biofilm development during the early stages of host colonization. © 2016 BSPP AND JOHN WILEY & SONS LTD.
Mirfenderesgi, Golnazalsadat; Bohrer, Gil; Matheny, Ashley M.; ...
2016-06-21
The finite difference ecosystem-scale tree crown hydrodynamics model version 2 (FETCH2) is a tree-scale hydrodynamic model of transpiration. The FETCH2 model employs a finite difference numerical methodology and a simplified single-beam conduit system to explicitly resolve xylem water potentials throughout the vertical extent of a tree. Empirical equations relate water potential within the stem to stomatal conductance of the leaves at each height throughout the crown. While highly simplified, this approach brings additional realism to the simulation of transpiration by linking stomatal responses to stem water potential rather than directly to soil moisture, as is currently the case in themore » majority of land surface models. FETCH2 accounts for plant hydraulic traits, such as the degree of anisohydric/isohydric response of stomata, maximal xylem conductivity, vertical distribution of leaf area, and maximal and minimal xylem water content. We used FETCH2 along with sap flow and eddy covariance data sets collected from a mixed plot of two genera (oak/pine) in Silas Little Experimental Forest, NJ, USA, to conduct an analysis of the intergeneric variation of hydraulic strategies and their effects on diurnal and seasonal transpiration dynamics. We define these strategies through the parameters that describe the genus level transpiration and xylem conductivity responses to changes in stem water potential. Here, our evaluation revealed that FETCH2 considerably improved the simulation of ecosystem transpiration and latent heat flux in comparison to more conventional models. In conclusion, a virtual experiment showed that the model was able to capture the effect of hydraulic strategies such as isohydric/anisohydric behavior on stomatal conductance under different soil-water availability conditions.« less
DOE Office of Scientific and Technical Information (OSTI.GOV)
Mirfenderesgi, Golnazalsadat; Bohrer, Gil; Matheny, Ashley M.
The finite difference ecosystem-scale tree crown hydrodynamics model version 2 (FETCH2) is a tree-scale hydrodynamic model of transpiration. The FETCH2 model employs a finite difference numerical methodology and a simplified single-beam conduit system to explicitly resolve xylem water potentials throughout the vertical extent of a tree. Empirical equations relate water potential within the stem to stomatal conductance of the leaves at each height throughout the crown. While highly simplified, this approach brings additional realism to the simulation of transpiration by linking stomatal responses to stem water potential rather than directly to soil moisture, as is currently the case in themore » majority of land surface models. FETCH2 accounts for plant hydraulic traits, such as the degree of anisohydric/isohydric response of stomata, maximal xylem conductivity, vertical distribution of leaf area, and maximal and minimal xylem water content. We used FETCH2 along with sap flow and eddy covariance data sets collected from a mixed plot of two genera (oak/pine) in Silas Little Experimental Forest, NJ, USA, to conduct an analysis of the intergeneric variation of hydraulic strategies and their effects on diurnal and seasonal transpiration dynamics. We define these strategies through the parameters that describe the genus level transpiration and xylem conductivity responses to changes in stem water potential. Here, our evaluation revealed that FETCH2 considerably improved the simulation of ecosystem transpiration and latent heat flux in comparison to more conventional models. In conclusion, a virtual experiment showed that the model was able to capture the effect of hydraulic strategies such as isohydric/anisohydric behavior on stomatal conductance under different soil-water availability conditions.« less
NASA Astrophysics Data System (ADS)
Mirfenderesgi, Golnazalsadat; Bohrer, Gil; Matheny, Ashley M.; Fatichi, Simone; de Moraes Frasson, Renato Prata; Schäfer, Karina V. R.
2016-07-01
The finite difference ecosystem-scale tree crown hydrodynamics model version 2 (FETCH2) is a tree-scale hydrodynamic model of transpiration. The FETCH2 model employs a finite difference numerical methodology and a simplified single-beam conduit system to explicitly resolve xylem water potentials throughout the vertical extent of a tree. Empirical equations relate water potential within the stem to stomatal conductance of the leaves at each height throughout the crown. While highly simplified, this approach brings additional realism to the simulation of transpiration by linking stomatal responses to stem water potential rather than directly to soil moisture, as is currently the case in the majority of land surface models. FETCH2 accounts for plant hydraulic traits, such as the degree of anisohydric/isohydric response of stomata, maximal xylem conductivity, vertical distribution of leaf area, and maximal and minimal xylem water content. We used FETCH2 along with sap flow and eddy covariance data sets collected from a mixed plot of two genera (oak/pine) in Silas Little Experimental Forest, NJ, USA, to conduct an analysis of the intergeneric variation of hydraulic strategies and their effects on diurnal and seasonal transpiration dynamics. We define these strategies through the parameters that describe the genus level transpiration and xylem conductivity responses to changes in stem water potential. Our evaluation revealed that FETCH2 considerably improved the simulation of ecosystem transpiration and latent heat flux in comparison to more conventional models. A virtual experiment showed that the model was able to capture the effect of hydraulic strategies such as isohydric/anisohydric behavior on stomatal conductance under different soil-water availability conditions.
Tombesi, Sergio; Johnson, R. Scott; Day, Kevin R.; DeJong, Theodore M.
2010-01-01
Background and Aims Previous studies indicate that the size-controlling capacity of peach rootstocks is associated with reductions of scion water potential during mid-day that are caused by the reduced hydraulic conductance of the rootstock. Thus, shoot growth appears to be reduced by decreases in stem water potential. The aim of this study was to investigate the mechanism of reduced hydraulic conductance in size-controlling peach rootstocks. Methods Anatomical measurements (diameter and frequency) of xylem vessels were determined in shoots, trunks and roots of three contrasting peach rootstocks grown as trees, each with different size-controlling characteristics: ‘Nemaguard’ (vigorous), ‘P30-135’ (intermediate vigour) and ‘K146-43’ (substantially dwarfing). Based on anatomical measurements, the theoretical axial xylem conductance of each tissue type and rootstock genotype was calculated via the Poiseuille–Hagen law. Key Results Larger vessel dimensions were found in the vigorous rootstock (‘Nemaguard’) than in the most dwarfing one (‘K146-43’) whereas vessels of ‘P30-135’ had intermediate dimensions. The density of vessels per xylem area in ‘Nemaguard’ was also less than in ‘P30-135’and ‘K146-43’. These characteristics resulted in different estimated hydraulic conductance among rootstocks: ‘Nemaguard’ had higher theoretical values followed by ‘P30-135’ and ‘K146-43’. Conclusions These data indicate that phenotypic differences in xylem anatomical characteristics of rootstock genotypes appear to influence hydraulic conductance capacity directly, and therefore may be the main determinant of dwarfing in these peach rootstocks. PMID:19939979
Hydraulic conductivity and embolism in the mangrove tree Laguncularia racemosa.
Ewers, Frank W; Lopez-Portillo, Jórge; Angeles, Guillermo; Fisher, Jack B
2004-09-01
We measured xylem pressure potentials, soil osmotic potentials, hydraulic conductivity and percent loss of conductivity (PLC) due to embolism, and made microscopic observations of perfused dye in the white mangrove tree, Laguncularia racemosa (L.) Gaertn. f., (1) to determine its vulnerability to air embolism compared with published results for the highly salt-tolerant red mangrove tree, Rhizophora mangle L., and (2) to identify possible relationships between air embolism, permanent blockage of vessels and stem diameter. Laguncularia racemosa was more vulnerable to embolism than reported for R. mangle, with 50 PLC at -3.4 MPa. Narrow stems (5-mm diameter) had higher PLC than larger stems (8.4- or 14-mm diameter) of the same plants. Basic fuchsin dye indicated that up to 89% of the vessels, especially in the narrow stems, had permanent blockage that could not be reversed by high pressure perfusion. Air embolism could lead to permanent vessel blockage and eventual stem mortality. Such vulnerability to embolism may restrict the growth of L. racemosa and limit its distribution to less salty areas of mangrove communities.
A synthetic leaf: the biomimetic potential of graphene oxide
NASA Astrophysics Data System (ADS)
Lamb, Marilla; Koch, George W.; Morgan, Eric R.; Shafer, Michael W.
2015-03-01
Emerging materials such as graphene oxide (GO) have micro and nano features that are functionally similar to those in plant cell walls involved in water transport. Therefore, it may now be possible to design and build biomimetic trees to lift water via mechanisms similar to those employed by trees, allowing for potential applications such as passive water pumping, filtering, and evaporative cooling. The tallest trees can raise large volumes of water to over 100 meters using only the vapor pressure gradient between their leaves and the atmosphere. This phenomenon occurs in all terrestrial plants when capillary forces generated in the microscopic pores in the cell walls of leaves are collectively applied to large diameter xylem conduits. The design of a synthetic tree that mimics these mechanisms will allow water to be moved to heights greater than is currently possible by any engineered system that does not require the use of a positive pressure pump. We are testing the suitability of membranous GO as the leaf of a synthetic tree and present an analysis in support of this design. In addition, we include results from a preliminary design using ceramics.
Jiang, Guo-Feng; Goodale, Uromi Manage; Liu, Yan-Yan; Hao, Guang-You; Cao, Kun-Fang
2017-03-01
Mangroves in hypersaline coastal habitats are under constant high xylem tension and face great risk of hydraulic dysfunction. To investigate the relationships between functional traits and salt management, we measured 20 hydraulic and photosynthetic traits in four salt-adapted (SA) and two non-SA (NSA) mangrove tree species in south China. The SA species included two salt secretors (SSs), Avicennia marina (Forsskål) Vierhapper and Aegiceras corniculatum (L.) Blanco and two salt excluders (SEs), Bruguiera gymnorrhiza (L.) Savigny and Kandelia obovata (L.) Sheue et al. The two NSA species were Hibiscus tiliaceus (L.) and Pongamia pinnata (L.) Merr. Extremely high xylem cavitation resistance, indicated by water potential at 50% loss of xylem conductivity (Ψ50; -7.85 MPa), was found in SEs. Lower cavitation resistance was observed in SSs, and may result from incomplete salt removal that reduces the magnitude of xylem tension required to maintain water uptake from the soil. Surprisingly, the NSA species, P. pinnata, had very low Ψ50 (-5.44 MPa). Compared with NSAs, SAs had lower photosynthesis, vessel density, hydraulic conductivity and vessel diameter, but higher sapwood density. Eight traits were strongly associated with species' salt management strategies, with predawn water potential (ΨPD) and mean vessel diameter accounting for 95% flow (D95) having the most significant association; D95 separated SAs from NSAs and SEs had the lowest ΨPD. There was significant coupling between hydraulic traits and carbon assimilation traits. Instead of hydraulic safety being compromised by xylem efficiency, mangrove species with higher safety had higher efficiency and greater sapwood density (ρSapwood), but there was no relationship between ρSapwood and efficiency. Principal component analysis differentiated the species of the three salt management strategies by loading D, D95 and vessel density on the first axis and loading ΨPD, Ψ50 and water potential at 12% loss of xylem conductivity (Ψ12), ρSapwood and quantum yield on the second axis. Our results provide the first comparative characterization of hydraulic and photosynthetic traits among mangroves with different salt management strategies. © The Author 2017. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.
Monitoring of Freezing Dynamics in Trees: A Simple Phase Shift Causes Complexity1[OPEN
Charra-Vaskou, Katline
2017-01-01
During winter, trees have to cope with harsh conditions, including extreme freeze-thaw stress. This study focused on ice nucleation and propagation, related water shifts and xylem cavitation, as well as cell damage and was based on in situ monitoring of xylem (thermocouples) and surface temperatures (infrared imaging), ultrasonic emissions, and dendrometer analysis. Field experiments during late winter on Picea abies growing at the alpine timberline revealed three distinct freezing patterns: (1) from the top of the tree toward the base, (2) from thin branches toward the main stem’s top and base, and (3) from the base toward the top. Infrared imaging showed freezing within branches from their base toward distal parts. Such complex freezing causes dynamic and heterogenous patterns in water potential and probably in cavitation. This study highlights the interaction between environmental conditions upon freezing and thawing and demonstrates the enormous complexity of freezing processes in trees. Diameter shrinkage, which indicated water fluxes within the stem, and acoustic emission analysis, which indicated cavitation events near the ice front upon freezing, were both related to minimum temperature and, upon thawing, related to vapor pressure deficit and soil temperature. These complex patterns, emphasizing the common mechanisms between frost and drought stress, shed new light on winter tree physiology. PMID:28242655
Mayr, Stefan; Schmid, Peter; Laur, Joan; Rosner, Sabine; Charra-Vaskou, Katline; Dämon, Birgit; Hacke, Uwe G.
2014-01-01
Xylem embolism is a limiting factor for woody species worldwide. Conifers at the alpine timberline are exposed to drought and freeze-thaw stress during winter, which induce potentially lethal embolism. Previous studies indicated that timberline trees survive by xylem refilling. In this study on Picea abies, refilling was monitored during winter and spring seasons and analyzed in the laboratory and in situ experiments, based on hydraulic, anatomical, and histochemical methods. Refilling started in late winter, when the soil was frozen and soil water not available for the trees. Xylem embolism caused up to 86.2% ± 3.1% loss of conductivity and was correlated with the ratio of closed pits. Refilling of xylem as well as recovery in shoot conductance started in February and corresponded with starch accumulation in secondary phloem and in the mesophyll of needles, where we also observed increasing aquaporin densities in the phloem and endodermis. This indicates that active, cellular processes play a role for refilling even under winter conditions. As demonstrated by our experiments, water for refilling was thereby taken up via the branches, likely by foliar water uptake. Our results suggest that refilling is based on water shifts to embolized tracheids via intact xylem, phloem, and parenchyma, whereby aquaporins reduce resistances along the symplastic pathway and aspirated pits facilitate isolation of refilling tracheids. Refilling must be taken into account as a key process in plant hydraulics and in estimating future effects of climate change on forests and alpine tree ecosystems. PMID:24521876
Mayr, Stefan; Bertel, Clara; Dämon, Birgit; Beikircher, Barbara
2014-01-01
The xylem hydraulic efficiency and safety is usually measured on mechanically unstressed samples, although trees may be exposed to combined hydraulic and mechanical stress in the field. We analysed changes in hydraulic conductivity and vulnerability to drought-induced embolism during static bending of Picea abies and Pinus sylvestris branches as well as the effect of dynamic bending on the vulnerability. We hypothesized this mechanical stress to substantially impair xylem hydraulics. Intense static bending caused an only small decrease in hydraulic conductance (−19.5 ± 2.4% in P. abies) but no shift in vulnerability thresholds. Dynamic bending caused a 0.4 and 0.8 MPa decrease of the water potential at 50 and 88% loss of conductivity in P. sylvestris, but did not affect vulnerability thresholds in P. abies. With respect to applied extreme bending radii, effects on plant hydraulics were surprisingly small and are thus probably of minor eco-physiological importance. More importantly, results indicate that available xylem hydraulic analyses (of conifers) sufficiently reflect plant hydraulics under field conditions. PMID:24697679
Martorell, Sebastià; Diaz-Espejo, Antonio; Medrano, Hipólito; Ball, Marilyn C; Choat, Brendan
2014-03-01
In woody plants, photosynthetic capacity is closely linked to rates at which the plant hydraulic system can supply water to the leaf surface. Drought-induced embolism can cause sharp declines in xylem hydraulic conductivity that coincide with stomatal closure and reduced photosynthesis. Recovery of photosynthetic capacity after drought is dependent on restored xylem function, although few data exist to elucidate this coordination. We examined the dynamics of leaf gas exchange and xylem function in Eucalyptus pauciflora seedlings exposed to a cycle of severe water stress and recovery after re-watering. Stomatal closure and leaf turgor loss occurred at water potentials that delayed the extensive spread of embolism through the stem xylem. Stem hydraulic conductance recovered to control levels within 6 h after re-watering despite a severe drought treatment, suggesting an active mechanism embolism repair. However, stomatal conductance did not recover after 10 d of re-watering, effecting tighter control of transpiration post drought. The dynamics of recovery suggest that a combination of hydraulic and non-hydraulic factors influenced stomatal behaviour post drought. © 2013 John Wiley & Sons Ltd.
USDA-ARS?s Scientific Manuscript database
Drought induces xylem embolism formation, but grapevines can refill blocked conduits to restore transport capacity. It is unknown whether vulnerability to embolism formation and ability to repair differ among grapevine species. We analyzed in vivo embolism formation and repair using x-ray microtomog...
Dynamic variation in sapwood specific conductivity in six woody species.
J.C. Domec; F.C. Meinzer; B.L. Gartner; J. Housset
2007-01-01
Relationships between pressure gradients and flow rates in the xylem are incompletely understood because steady-state conductivity coefficients are inadequate for predicting and interpreting flow under the non-steady-state conditions more prevalent in intact trees. The goal of this study was to determine the magnitude of deviation of trunk sapwood specific conductivity...
Domec, J-C; Warren, J M; Meinzer, F C; Brooks, J R; Coulombe, R
2004-09-01
Hydraulic redistribution (HR), the passive movement of water via roots from moist to drier portions of the soil, occurs in many ecosystems, influencing both plant and ecosystem-water use. We examined the effects of HR on root hydraulic functioning during drought in young and old-growth Douglas-fir [ Pseudotsuga menziesii (Mirb.) Franco] and ponderosa pine ( Pinus ponderosa Dougl. Ex Laws) trees growing in four sites. During the 2002 growing season, in situ xylem embolism, water deficit and xylem vulnerability to embolism were measured on medium roots (2-4-mm diameter) collected at 20-30 cm depth. Soil water content and water potentials were monitored concurrently to determine the extent of HR. Additionally, the water potential and stomatal conductance ( g(s)) of upper canopy leaves were measured throughout the growing season. In the site with young Douglas-fir trees, root embolism increased from 20 to 55 percent loss of conductivity (PLC) as the dry season progressed. In young ponderosa pine, root embolism increased from 45 to 75 PLC. In contrast, roots of old-growth Douglas-fir and ponderosa pine trees never experienced more than 30 and 40 PLC, respectively. HR kept soil water potential at 20-30 cm depth above -0.5 MPa in the old-growth Douglas-fir site and -1.8 MPa in the old-growth ponderosa pine site, which significantly reduced loss of shallow root function. In the young ponderosa pine stand, where little HR occurred, the water potential in the upper soil layers fell to about -2.8 MPa, which severely impaired root functioning and limited recovery when the fall rains returned. In both species, daily maximum g(s) decreased linearly with increasing root PLC, suggesting that root xylem embolism acted in concert with stomata to limit water loss, thereby maintaining minimum leaf water potential above critical values. HR appears to be an important mechanism for maintaining shallow root function during drought and preventing total stomatal closure.
Secchi, Francesca; Zwieniecki, Maciej A
2016-11-01
Severe water stress constrains, or even stops, water transport in the xylem due to embolism formation. Previously, the xylem of poplar trees was shown to respond to embolism formation by accumulating carbohydrates in the xylem apoplast and dropping xylem sap pH. We hypothesize that these two processes may be functionally linked as lower pH activates acidic invertases degrading sucrose and inducing accumulation of monosaccharides in xylem apoplast. Using a novel in vivo method to measure xylem apoplast pH, we show that pH drops from ~6.2 to ~5.6 in stems of severely stressed plants and rises following recovery of stem water status. We also show that in a lower pH environment, sugars are continuously accumulating in the xylem apoplast. Apoplastic carbohydrate accumulation was reduced significantly in the presence of a proton pump blocker (orthovanadate). These observations suggest that a balance in sugar concentrations exists between the xylem apoplast and symplast that can be controlled by xylem pH and sugar concentration. We conclude that lower pH is related to loss of xylem transport function, eventually resulting in accumulation of sugars that primes stems for recovery from embolism when water stress is relieved. © 2016 John Wiley & Sons Ltd.
Lengthening of the duration of xylogenesis engenders disproportionate increases in xylem production.
Rossi, Sergio; Girard, Marie-Josée; Morin, Hubert
2014-07-01
In cold climates, the expected global warming will lead to earlier cambial resumptions in spring, with a resultant lengthening of the growing season but unknown consequences on forest productivity. The phenological traits of cambium activity and xylem formation were analyzed at a short time scale along a thermal gradient represented by an alti-latitudinal range from the 48th to 53rd parallels and covering the whole closed black-spruce [Picea mariana (Mill.) BSP] forest in Quebec, Canada. A hypothesis was tested that warmer temperatures influence cambium phenology, allowing longer duration and higher intensity of growth, and resulting in proportionally increased xylem production. From April to October 2012, cell division in cambium and post-cambial differentiation of xylem were observed on anatomical sections obtained from microcores collected weekly from the stem of fifty trees. The southern and warmer site was characterized by the highest radial growth, which corresponded to both the highest rates and longest durations of cell production. The differences in terms of xylem phenology and growth were marginal between the other sites. Xylem growth was positively correlated with rate and duration of cell production, with the latter explaining most variability in growth. Within the range analyzed, the relationship between temperature and most phenological phases of xylogenesis was linear. On the contrary, temperature was related with cell production according to an exponential pattern. Periods of xylogenesis of 14 days longer (+13.1%) corresponded to a massive increase in cell production (33 cells, +109%). This disproportionate change occurred at a May-September average temperature of ca. 14 °C and a snow-free period of 210-235 days. At the lower boundary of the distribution of black spruce, small environmental changes allowing marginal lengthening of the period of cell division could potentially lead to disproportionate increases in xylem cell production, with substantial consequences for the productivity of this boreal species. © 2013 John Wiley & Sons Ltd.
Husby, Chad E.; Delatorre, José; Oreste, Vittorio; Oberbauer, Steven F.; Palow, Danielle T.; Novara, Lázaro; Grau, Alfredo
2011-01-01
Background and aims The basic set of adaptations necessary for salinity tolerance in vascular plants remains unknown. Although much has been published on salinity stress, almost all studies deal with spermatophytes. Studies of salinity tolerance in pteridophytes are relatively rare but hold promise for revealing the fundamental adaptations that all salt-tolerant vascular plants may share. The most basal pteridophytes to exhibit salinity tolerance are members of the genus Equisetum, including the giant horsetail, Equisetum giganteum, the only pteridophyte to occur in salinity-affected regions of the Atacama Desert valleys of northern Chile. Here it can constitute a significant vegetation component, forming dense stands of shoots >4 m high. Methodology Physiological parameters (stomatal conductances; efficiency of photosystem II; sap osmotic potential) were measured in E. giganteum populations in northern Chile across a range of groundwater salinities at 11 sites. In addition, Na, K, electrical conductivity and total plant water potential were measured in the plants and groundwater from each site. Principal results Equisetum giganteum exhibits similar stomatal conductances and photochemical efficiencies of photosystem II across a wide range of groundwater salinities. It lowers cell sap osmotic potential with increasing salinity and produces positive root pressure, as evidenced by guttation, at the full range of salinities experienced in the Atacama Desert. Equisetum giganteum maintains low Na concentrations in its xylem fluid and cell sap when soil water Na is high. It also maintains high K/Na ratios in xylem fluid and cell sap when soil water has low K/Na ratios. Conclusions Equisetum giganteum is well adapted to salinity stress. Efficient K uptake and Na exclusion are important adaptations and closely similar to those of the facultative halophyte fern Acrostichum aureum. PMID:22476492
Ion-mediated changes of xylem hydraulic resistance in planta: fact or fiction?
van Ieperen, Wim
2007-04-01
Although xylem provides an efficient transport pathway for water in plants, the hydraulic conductivity of xylem (K(h)) can still influence plant water status. For decades, the K(h) of functional xylem has been assumed to be constant in the short term because xylem consists of a network of dead interconnected capillary elements (conduits). Recent research has shown that K(h) can change in response to the cation content of the xylem fluid. Volume changes of pectin gel in nanometer-sized pores at inter-conduit connections are hypothesized to be the cause, and implications for xylem transport in planta are suggested. However, it seems too early to be conclusive about this phenomenon because the phenomenon has not been measured in planta with xylem fluids that realistically mimic natural xylem sap and the applied methods used to measure ion-mediated changes in K(h) have drawbacks.
NASA Astrophysics Data System (ADS)
Love, D. M.; Venturas, M.; Sperry, J.; Wang, Y.; Anderegg, W.
2017-12-01
Modeling approaches for tree stomatal control often rely on empirical fitting to provide accurate estimates of whole tree transpiration (E) and assimilation (A), which are limited in their predictive power by the data envelope used to calibrate model parameters. Optimization based models hold promise as a means to predict stomatal behavior under novel climate conditions. We designed an experiment to test a hydraulic trait based optimization model, which predicts stomatal conductance from a gain/risk approach. Optimal stomatal conductance is expected to maximize the potential carbon gain by photosynthesis, and minimize the risk to hydraulic transport imposed by cavitation. The modeled risk to the hydraulic network is assessed from cavitation vulnerability curves, a commonly measured physiological trait in woody plant species. Over a growing season garden grown plots of aspen (Populus tremuloides, Michx.) and ponderosa pine (Pinus ponderosa, Douglas) were subjected to three distinct drought treatments (moderate, severe, severe with rehydration) relative to a control plot to test model predictions. Model outputs of predicted E, A, and xylem pressure can be directly compared to both continuous data (whole tree sapflux, soil moisture) and point measurements (leaf level E, A, xylem pressure). The model also predicts levels of whole tree hydraulic impairment expected to increase mortality risk. This threshold is used to estimate survivorship in the drought treatment plots. The model can be run at two scales, either entirely from climate (meteorological inputs, irrigation) or using the physiological measurements as a starting point. These data will be used to study model performance and utility, and aid in developing the model for larger scale applications.
Hydraulic efficiency and safety of vascular and non-vascular components in Pinus pinaster leaves.
Charra-Vaskou, Katline; Badel, Eric; Burlett, Régis; Cochard, Hervé; Delzon, Sylvain; Mayr, Stefan
2012-09-01
Leaves, the distal section of the soil-plant-atmosphere continuum, exhibit the lowest water potentials in a plant. In contrast to angiosperm leaves, knowledge of the hydraulic architecture of conifer needles is scant. We investigated the hydraulic efficiency and safety of Pinus pinaster needles, comparing different techniques. The xylem hydraulic conductivity (k(s)) and embolism vulnerability (P(50)) of both needle and stem were measured using the cavitron technique. The conductance and vulnerability of whole needles were measured via rehydration kinetics, and Cryo-SEM and 3D X-ray microtomographic observations were used as reference tools to validate physical measurements. The needle xylem of P. pinaster had lower hydraulic efficiency (k(s) = 2.0 × 10(-4) m(2) MPa(-1) s(-1)) and safety (P(50) = - 1.5 MPa) than stem xylem (k(s) = 7.7 × 10(-4) m(2) MPa(-1) s(-1); P(50) = - 3.6 to - 3.2 MPa). P(50) of whole needles (both extra-vascular and vascular pathways) was - 0.5 MPa, suggesting that non-vascular tissues were more vulnerable than the xylem. During dehydration to - 3.5 MPa, collapse and embolism in xylem tracheids, and gap formation in surrounding tissues were observed. However, a discrepancy in hydraulic and acoustic results appeared compared with visualizations, arguing for greater caution with these techniques when applied to needles. Our results indicate that the most distal parts of the water transport pathway are limiting for hydraulics of P. pinaster. Needle tissues exhibit a low hydraulic efficiency and low hydraulic safety, but may also act to buffer short-term water deficits, thus preventing xylem embolism.
Trade-offs between xylem hydraulic properties, wood anatomy and yield in Populus.
Hajek, Peter; Leuschner, Christoph; Hertel, Dietrich; Delzon, Sylvain; Schuldt, Bernhard
2014-07-01
Trees face the dilemma that achieving high plant productivity is accompanied by a risk of drought-induced hydraulic failure due to a trade-off in the trees' vascular system between hydraulic efficiency and safety. By investigating the xylem anatomy of branches and coarse roots, and measuring branch axial hydraulic conductivity and vulnerability to cavitation in 4-year-old field-grown aspen plants of five demes (Populus tremula L. and Populus tremuloides Michx.) differing in growth rate, we tested the hypotheses that (i) demes differ in wood anatomical and hydraulic properties, (ii) hydraulic efficiency and safety are related to xylem anatomical traits, and (iii) aboveground productivity and hydraulic efficiency are negatively correlated to cavitation resistance. Significant deme differences existed in seven of the nine investigated branch-related anatomical and hydraulic traits but only in one of the four coarse-root-related anatomical traits; this likely is a consequence of high intra-plant variation in root morphology and the occurrence of a few 'high-conductivity roots'. Growth rate was positively related to branch hydraulic efficiency (xylem-specific conductivity) but not to cavitation resistance; this indicates that no marked trade-off exists between cavitation resistance and growth. Both branch hydraulic safety and hydraulic efficiency significantly depended on vessel size and were related to the genetic distance between the demes, while the xylem pressure causing 88% loss of hydraulic conductivity (P88 value) was more closely related to hydraulic efficiency than the commonly used P50 value. Deme-specific variation in the pit membrane structure may explain why vessel size was not directly linked to growth rate. We conclude that branch hydraulic efficiency is an important growth-influencing trait in aspen, while the assumed trade-off between productivity and hydraulic safety is weak. © The Author 2014. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.
Unraveling the Effects of Plant Hydraulics on Stomatal Closure during Water Stress in Walnut
Cochard, Hervé; Coll, Lluis; Le Roux, Xavier; Améglio, Thierry
2002-01-01
The objectives of the study were to identify the relevant hydraulic parameters associated with stomatal regulation during water stress and to test the hypothesis of a stomatal control of xylem embolism in walnut (Juglans regia × nigra) trees. The hydraulic characteristics of the sap pathway were experimentally altered with different methods to alter plant transpiration (Eplant) and stomatal conductance (gs). Potted trees were exposed to a soil water depletion to alter soil water potential (Ψsoil), soil resistance (Rsoil), and root hydraulic resistances (Rroot). Soil temperature was changed to alter Rroot alone. Embolism was created in the trunk to increase shoot resistance (Rshoot). Stomata closed in response to these stresses with the effect of maintaining the water pressure in the leaf rachis xylem (Prachis) above −1.4 MPa and the leaf water potential (Ψleaf) above −1.6 MPa. The same dependence of Eplant and gs on Prachis or Ψleaf was always observed. This suggested that stomata were not responding to changes in Ψsoil, Rsoil, Rroot, or Rshoot per se but rather to their impact on Prachis and/or Ψleaf. Leaf rachis was the most vulnerable organ, with a threshold Prachis for embolism induction of −1.4 MPa. The minimum Ψleaf values corresponded to leaf turgor loss point. This suggested that stomata are responding to leaf water status as determined by transpiration rate and plant hydraulics and that Prachis might be the physiological parameter regulated by stomatal closure during water stress, which would have the effect of preventing extensive developments of cavitation during water stress. PMID:11788773
USDA-ARS?s Scientific Manuscript database
It is not currently well-understood how much xylem conductance is lost in maize plants during the day, if conductance is recovered during the night, or what soil water conditions are required for recovery to take place. To answer these questions we designed a greenhouse experiment whereby two genet...
de Bernonville, Thomas Dugé; Albenne, Cécile; Arlat, Matthieu; Hoffmann, Laurent; Lauber, Emmanuelle; Jamet, Elisabeth
2014-01-01
Proteomic analysis of xylem sap has recently become a major field of interest to understand several biological questions related to plant development and responses to environmental clues. The xylem sap appears as a dynamic fluid undergoing changes in its proteome upon abiotic and biotic stresses. Unlike cell compartments which are amenable to purification in sufficient amount prior to proteomic analysis, the xylem sap has to be collected in particular conditions to avoid contamination by intracellular proteins and to obtain enough material. A model plant like Arabidopsis thaliana is not suitable for such an analysis because efficient harvesting of xylem sap is difficult. The analysis of the xylem sap proteome also requires specific procedures to concentrate proteins and to focus on proteins predicted to be secreted. Indeed, xylem sap proteins appear to be synthesized and secreted in the root stele or to originate from dying differentiated xylem cells. This chapter describes protocols to collect xylem sap from Brassica species and to prepare total and N-glycoprotein extracts for identification of proteins by mass spectrometry analyses and bioinformatics.
Knipfer, Thorsten; Eustis, Ashley; Brodersen, Craig; Walker, Andrew M; McElrone, Andrew J
2015-08-01
Drought induces xylem embolism formation, but grapevines can refill non-functional vessels to restore transport capacity. It is unknown whether vulnerability to embolism formation and ability to repair differ among grapevine species. We analysed in vivo embolism formation and repair using x-ray computed microtomography in three wild grapevine species from varied native habitats (Vitis riparia, V. arizonica, V. champinii) and related responses to measurements of leaf gas exchange and root pressure. Vulnerability to embolism formation was greatest in V. riparia, intermediate in V. arizonica and lowest in V. champinii. After re-watering, embolism repair was rapid and pronounced in V. riparia and V. arizonica, but limited or negligible in V. champinii even after numerous days. Similarly, root pressure measured after re-watering was positively correlated with drought stress severity for V. riparia and V. arizonica (species exhibiting embolism repair) but not for V. champinii. Drought-induced reductions in transpiration were greatest for V. riparia and least in V. champinii. Recovery of transpiration after re-watering was delayed for all species, but was greatest for V. champinii and most rapid in V. arizonica. These species exhibit varied responses to drought stress that involve maintenance/recovery of xylem transport capacity coordinated with root pressure and gas exchange responses. © 2014 John Wiley & Sons Ltd.
Differential expression of genes of Xylella fastidiosa in xylem fluid of citrus and grapevine.
Shi, Xiangyang; Bi, Jianlong; Morse, Joseph G; Toscano, Nick C; Cooksey, Donald A
2010-03-01
Xylella fastidiosa causes a serious Pierce's disease (PD) in grapevine. Xylella fastidiosa cells from a PD strain were grown in a pure xylem fluid of a susceptible grapevine cultivar vs. xylem fluid from citrus, which is not a host for this strain of X. fastidiosa. When grown in grapevine xylem fluid, cells of the PD strain formed clumps and biofilm formed to a greater extent than in citrus xylem fluid, although the PD strain did grow in xylem fluid of three citrus varieties. The differential expression of selected genes of a PD X. fastidiosa strain cultured in the two xylem fluids was analyzed using a DNA macroarray. Compared with citrus xylem fluid, grapevine xylem fluid stimulated the expression of X. fastidiosa genes involved in virulence regulation, such as gacA, algU, xrvA, and hsq, and also genes involved in the biogenesis of pili and twitching motility, such as fimT, pilI, pilU, and pilY1. Increased gene expression likely contributes to PD expression in grapevine, whereas citrus xylem fluid did not support or possibly suppressed the expression of these virulence genes.
Zaller, Johann G; Cantelmo, Clemens; Santos, Gabriel Dos; Muther, Sandrina; Gruber, Edith; Pallua, Paul; Mandl, Karin; Friedrich, Barbara; Hofstetter, Ingrid; Schmuckenschlager, Bernhard; Faber, Florian
2018-06-03
Herbicides are increasingly applied in vineyards worldwide. However, not much is known on potential side effects on soil organisms or on the nutrition of grapevines (Vitis vinifera). In an experimental vineyard in Austria, we examined the impacts of three within-row herbicide treatments (active ingredients: flazasulfuron, glufosinate, glyphosate) and mechanical weeding on grapevine root mycorrhization; soil microorganisms; earthworms; and nutrient concentration in grapevine roots, leaves, xylem sap and grape juice. The three herbicides reduced grapevine root mycorrhization on average by 53% compared to mechanical weeding. Soil microorganisms (total colony-forming units, CFU) were significantly affected by herbicides with highest CFUs under glufosinate and lowest under glyphosate. Earthworms (surface casting activity, density, biomass, reproduction) or litter decomposition in soil were unaffected by herbicides. Herbicides altered nutrient composition in grapevine roots, leaves, grape juice and xylem sap that was collected 11 months after herbicide application. Xylem sap under herbicide treatments also contained on average 70% more bacteria than under mechanical weeding; however, due to high variability, this was not statistically significant. We conclude that interdisciplinary approaches should receive more attention when assessing ecological effects of herbicides in vineyard ecosystems.
Resistance to Dutch Elm Disease Reduces Presence of Xylem Endophytic Fungi in Elms (Ulmus spp.)
Martín, Juan A.; Witzell, Johanna; Blumenstein, Kathrin; Rozpedowska, Elzbieta; Helander, Marjo; Sieber, Thomas N.; Gil, Luis
2013-01-01
Efforts to introduce pathogen resistance into landscape tree species by breeding may have unintended consequences for fungal diversity. To address this issue, we compared the frequency and diversity of endophytic fungi and defensive phenolic metabolites in elm (Ulmus spp.) trees with genotypes known to differ in resistance to Dutch elm disease. Our results indicate that resistant U. minor and U. pumila genotypes exhibit a lower frequency and diversity of fungal endophytes in the xylem than susceptible U. minor genotypes. However, resistant and susceptible genotypes showed a similar frequency and diversity of endophytes in the leaves and bark. The resistant and susceptible genotypes could be discriminated on the basis of the phenolic profile of the xylem, but not on basis of phenolics in the leaves or bark. As the Dutch elm disease pathogen develops within xylem tissues, the defensive chemistry of resistant elm genotypes thus appears to be one of the factors that may limit colonization by both the pathogen and endophytes. We discuss a potential trade-off between the benefits of breeding resistance into tree species, versus concomitant losses of fungal endophytes and the ecosystem services they provide. PMID:23468900
Mayr, Stefan; Bertel, Clara; Dämon, Birgit; Beikircher, Barbara
2014-09-01
The xylem hydraulic efficiency and safety is usually measured on mechanically unstressed samples, although trees may be exposed to combined hydraulic and mechanical stress in the field. We analysed changes in hydraulic conductivity and vulnerability to drought-induced embolism during static bending of Picea abies and Pinus sylvestris branches as well as the effect of dynamic bending on the vulnerability. We hypothesized this mechanical stress to substantially impair xylem hydraulics. Intense static bending caused an only small decrease in hydraulic conductance (-19.5 ± 2.4% in P. abies) but no shift in vulnerability thresholds. Dynamic bending caused a 0.4 and 0.8 MPa decrease of the water potential at 50 and 88% loss of conductivity in P. sylvestris, but did not affect vulnerability thresholds in P. abies. With respect to applied extreme bending radii, effects on plant hydraulics were surprisingly small and are thus probably of minor eco-physiological importance. More importantly, results indicate that available xylem hydraulic analyses (of conifers) sufficiently reflect plant hydraulics under field conditions. © 2014 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.
Huang, Ping; Wan, Xianchong; Lieffers, Victor J
2016-05-01
This study tested how wind in daytime and nighttime affects hydraulic properties and thigmomorphogenic response of poplar saplings. It shows that wind in daytime interrupted water balance of poplar plants by aggravating cavitation in the stem xylem under high xylem tension in the daytime, reducing water potential in midday and hence reducing gas exchange, including stomatal conductance and CO2 assimilation. The wind blowing in daytime significantly reduced plant growth, including height, diameter, leaf size, leaf area, root and whole biomass, whereas wind blowing in nighttime only caused a reduction in radial and height growth at the early stage compared with the control but decreased height:diameter ratios. In summary, the interaction between wind loading and xylem tension exerted a negative impact on water balance, gas exchanges and growth of poplar plants, and wind in nighttime caused only a small thigmomorphogenic response. © 2015 Scandinavian Plant Physiology Society.
Plant xylem hydraulics: What we understand, current research, and future challenges.
Venturas, Martin D; Sperry, John S; Hacke, Uwe G
2017-06-01
Herein we review the current state-of-the-art of plant hydraulics in the context of plant physiology, ecology, and evolution, focusing on current and future research opportunities. We explain the physics of water transport in plants and the limits of this transport system, highlighting the relationships between xylem structure and function. We describe the great variety of techniques existing for evaluating xylem resistance to cavitation. We address several methodological issues and their connection with current debates on conduit refilling and exponentially shaped vulnerability curves. We analyze the trade-offs existing between water transport safety and efficiency. We also stress how little information is available on molecular biology of cavitation and the potential role of aquaporins in conduit refilling. Finally, we draw attention to how plant hydraulic traits can be used for modeling stomatal responses to environmental variables and climate change, including drought mortality. © 2017 Institute of Botany, Chinese Academy of Sciences.
Townes, Shatara V.; Bartlett, Megan K.; Buckley, Thomas N.; McElrone, Andrew J.; Sack, Lawren
2017-01-01
Leaf hydraulic supply is crucial to maintaining open stomata for CO2 capture and plant growth. During drought-induced dehydration, the leaf hydraulic conductance (Kleaf) declines, which contributes to stomatal closure and, eventually, to leaf death. Previous studies have tended to attribute the decline of Kleaf to embolism in the leaf vein xylem. We visualized at high resolution and quantified experimentally the hydraulic vulnerability of xylem and outside-xylem pathways and modeled their respective influences on plant water transport. Evidence from all approaches indicated that the decline of Kleaf during dehydration arose first and foremost due to the vulnerability of outside-xylem tissues. In vivo x-ray microcomputed tomography of dehydrating leaves of four diverse angiosperm species showed that, at the turgor loss point, only small fractions of leaf vein xylem conduits were embolized, and substantial xylem embolism arose only under severe dehydration. Experiments on an expanded set of eight angiosperm species showed that outside-xylem hydraulic vulnerability explained 75% to 100% of Kleaf decline across the range of dehydration from mild water stress to beyond turgor loss point. Spatially explicit modeling of leaf water transport pointed to a role for reduced membrane conductivity consistent with published data for cells and tissues. Plant-scale modeling suggested that outside-xylem hydraulic vulnerability can protect the xylem from tensions that would induce embolism and disruption of water transport under mild to moderate soil and atmospheric droughts. These findings pinpoint outside-xylem tissues as a central locus for the control of leaf and plant water transport during progressive drought. PMID:28049739
Structure-function constraints of tracheid-based xylem: a comparison of conifers and ferns.
Pittermann, Jarmila; Limm, Emily; Rico, Christopher; Christman, Mairgareth A
2011-10-01
The ferns comprise one of the most ancient tracheophytic plant lineages, and occupy habitats ranging from tundra to deserts and the equatorial tropics. Like their nearest relatives the conifers, modern ferns possess tracheid-based xylem but the structure-function relationships of fern xylem are poorly understood. Here, we sampled the fronds (megaphylls) of 16 species across the fern phylogeny, and examined the relationships among hydraulic transport, drought-induced cavitation resistance, the xylem anatomy of the stipe, and the gas-exchange response of the pinnae. For comparison, the results are presented alongside a similar suite of conifer data. Fern xylem is as resistant to cavitation as conifer xylem, but exhibits none of the hydraulic or structural trade-offs associated with resistance to cavitation. On a conduit diameter basis, fern xylem can exhibit greater hydraulic efficiency than conifer and angiosperm xylem. In ferns, wide and long tracheids compensate in part for the lack of secondary xylem and allow ferns to exhibit transport rates on a par with those of conifers. We suspect that it is the arrangement of the primary xylem, in addition to the intrinsic traits of the conduits themselves, that may help explain the broad range of cavitation resistance in ferns. © 2011 The Authors. New Phytologist © 2011 New Phytologist Trust.
An annual pattern of native embolism in upper branches of four tall conifer species
Katherine A. McCulloh; Daniel M. Johnson; Frederick C. Meinzer; Barbara Lachenbruch
2011-01-01
The Pacific Northwest of North America experiences relatively mild winters and dry summers. For the tall coniferous trees that grow in this region, we predicted that loss in the hydraulic conductivity of uppermost branches would be avoided because of difficulty reversing accumulated emboli in xylem that is always under negative pressure. To test this hypothesis, we...
Anderegg, William R L; Klein, Tamir; Bartlett, Megan; Sack, Lawren; Pellegrini, Adam F A; Choat, Brendan; Jansen, Steven
2016-05-03
Drought-induced tree mortality has been observed globally and is expected to increase under climate change scenarios, with large potential consequences for the terrestrial carbon sink. Predicting mortality across species is crucial for assessing the effects of climate extremes on forest community biodiversity, composition, and carbon sequestration. However, the physiological traits associated with elevated risk of mortality in diverse ecosystems remain unknown, although these traits could greatly improve understanding and prediction of tree mortality in forests. We performed a meta-analysis on species' mortality rates across 475 species from 33 studies around the globe to assess which traits determine a species' mortality risk. We found that species-specific mortality anomalies from community mortality rate in a given drought were associated with plant hydraulic traits. Across all species, mortality was best predicted by a low hydraulic safety margin-the difference between typical minimum xylem water potential and that causing xylem dysfunction-and xylem vulnerability to embolism. Angiosperms and gymnosperms experienced roughly equal mortality risks. Our results provide broad support for the hypothesis that hydraulic traits capture key mechanisms determining tree death and highlight that physiological traits can improve vegetation model prediction of tree mortality during climate extremes.
Hierarchical statistical modeling of xylem vulnerability to cavitation.
Ogle, Kiona; Barber, Jarrett J; Willson, Cynthia; Thompson, Brenda
2009-01-01
Cavitation of xylem elements diminishes the water transport capacity of plants, and quantifying xylem vulnerability to cavitation is important to understanding plant function. Current approaches to analyzing hydraulic conductivity (K) data to infer vulnerability to cavitation suffer from problems such as the use of potentially unrealistic vulnerability curves, difficulty interpreting parameters in these curves, a statistical framework that ignores sampling design, and an overly simplistic view of uncertainty. This study illustrates how two common curves (exponential-sigmoid and Weibull) can be reparameterized in terms of meaningful parameters: maximum conductivity (k(sat)), water potential (-P) at which percentage loss of conductivity (PLC) =X% (P(X)), and the slope of the PLC curve at P(X) (S(X)), a 'sensitivity' index. We provide a hierarchical Bayesian method for fitting the reparameterized curves to K(H) data. We illustrate the method using data for roots and stems of two populations of Juniperus scopulorum and test for differences in k(sat), P(X), and S(X) between different groups. Two important results emerge from this study. First, the Weibull model is preferred because it produces biologically realistic estimates of PLC near P = 0 MPa. Second, stochastic embolisms contribute an important source of uncertainty that should be included in such analyses.
NASA Astrophysics Data System (ADS)
Allen, S. T.; Kirchner, J. W.; Braun, S.; Siegwolf, R. T.; Goldsmith, G. R.
2017-12-01
Xylem water isotopic composition can reveal how water moves through soil and is subsequently taken up by plants. By examining how xylem water isotopes vary across distinct climates and soils, we test how these site characteristics control critical-zone water movement and tree uptake. Xylem water was collected from over 900 trees at 191 sites across Switzerland during a 10-day period in mid-summer 2015. Sites contained oak, beech and/or spruce trees and ranged in elevation from 260 to 1870 m asl with mean annual precipitation from 700 to 2060 mm. Xylem water samples were analyzed for 2H and 18O using isotope ratio mass spectrometry. Patterns in the temporal origin of xylem water showed regional differences. For example, trees in the southern and alpine regions had xylem water isotopic signatures that more closely resembled summer precipitation. The isotopic spatial range observed for mid-summer xylem waters was similar to the seasonal range of precipitation; that is, mid-summer xylem water at some sites resembled summer precipitation, and at other sites resembled winter precipitation. Xylem water from spruces, oaks, and beeches at the same sites did not differ from each other, despite these species having different rooting habits. Across all sites and species, precipitation amount correlated positively with xylem δ18O. In higher-precipitation areas, summer rain apparently displaces or mixes with older (winter) stored waters, thus reducing the winter-water isotopic signal in xylem water. Alternatively, in areas with limited precipitation, xylem water more closely matched winter water, indicating greater use of older stored water. We conclude that regional variations in precipitation deficits determine variations in the turnover rate of plant-available soil water and storage.
Outside-Xylem Vulnerability, Not Xylem Embolism, Controls Leaf Hydraulic Decline during Dehydration.
Scoffoni, Christine; Albuquerque, Caetano; Brodersen, Craig R; Townes, Shatara V; John, Grace P; Bartlett, Megan K; Buckley, Thomas N; McElrone, Andrew J; Sack, Lawren
2017-02-01
Leaf hydraulic supply is crucial to maintaining open stomata for CO 2 capture and plant growth. During drought-induced dehydration, the leaf hydraulic conductance (K leaf ) declines, which contributes to stomatal closure and, eventually, to leaf death. Previous studies have tended to attribute the decline of K leaf to embolism in the leaf vein xylem. We visualized at high resolution and quantified experimentally the hydraulic vulnerability of xylem and outside-xylem pathways and modeled their respective influences on plant water transport. Evidence from all approaches indicated that the decline of K leaf during dehydration arose first and foremost due to the vulnerability of outside-xylem tissues. In vivo x-ray microcomputed tomography of dehydrating leaves of four diverse angiosperm species showed that, at the turgor loss point, only small fractions of leaf vein xylem conduits were embolized, and substantial xylem embolism arose only under severe dehydration. Experiments on an expanded set of eight angiosperm species showed that outside-xylem hydraulic vulnerability explained 75% to 100% of K leaf decline across the range of dehydration from mild water stress to beyond turgor loss point. Spatially explicit modeling of leaf water transport pointed to a role for reduced membrane conductivity consistent with published data for cells and tissues. Plant-scale modeling suggested that outside-xylem hydraulic vulnerability can protect the xylem from tensions that would induce embolism and disruption of water transport under mild to moderate soil and atmospheric droughts. These findings pinpoint outside-xylem tissues as a central locus for the control of leaf and plant water transport during progressive drought. © 2017 The author(s). All Rights Reserved.
Hydraulics of high-yield orchard trees: a case study of three Malus domestica cultivars.
Beikircher, Barbara; De Cesare, Chiara; Mayr, Stefan
2013-12-01
The drought tolerance of three economically important apple cultivars, Golden Delicious, Braeburn and Red Delicious, was analysed. The work offers insights into the hydraulics of these high-yield trees and indicates a possible hydraulic limitation of carbon gain. The hydraulic safety and efficiency of branch xylem and leaves were quantified, drought tolerance of living tissues was measured and stomatal regulation, turgor-loss point and osmotic potential at full turgor were analysed. Physiological measurements were correlated with anatomical parameters, such as conduit diameter, cell-wall reinforcement, stomatal density and stomatal pore length. Hydraulic safety differed considerably between the three cultivars with Golden Delicious being significantly less vulnerable to drought-induced embolism than Braeburn and Red Delicious. In Golden Delicious, leaves were less resistant than branch xylem, while in the other cultivars leaves were more resistant than branch xylem. Hydraulic efficiency and xylem anatomical measurements indicate differences in pit properties, which may also be responsible for variations in hydraulic safety. In all three cultivars, full stomatal closure occurred at water potentials where turgor had already been lost and severe loss of hydraulic conductivity as well as damage to living cells had been induced. The consequential negative safety margins pose a risk for hydraulic failure but facilitate carbon gain, which is further improved by the observed high stomatal conductance. Maximal stomatal conductance was clearly seen to be related to stomatal density and size. Based on our results, these three high-yield Malus domestica Borkh. cultivars span a wide range of drought tolerances, appear optimized for maximal carbon gain and, thus, all perform best under well-managed growing conditions.
Hefer, Charles A; Mizrachi, Eshchar; Myburg, Alexander A; Douglas, Carl J; Mansfield, Shawn D
2015-06-01
Wood formation is a complex developmental process governed by genetic and environmental stimuli. Populus and Eucalyptus are fast-growing, high-yielding tree genera that represent ecologically and economically important species suitable for generating significant lignocellulosic biomass. Comparative analysis of the developing xylem and leaf transcriptomes of Populus trichocarpa and Eucalyptus grandis together with phylogenetic analyses identified clusters of homologous genes preferentially expressed during xylem formation in both species. A conserved set of 336 single gene pairs showed highly similar xylem preferential expression patterns, as well as evidence of high functional constraint. Individual members of multi-gene orthologous clusters known to be involved in secondary cell wall biosynthesis also showed conserved xylem expression profiles. However, species-specific expression as well as opposite (xylem versus leaf) expression patterns observed for a subset of genes suggest subtle differences in the transcriptional regulation important for xylem development in each species. Using sequence similarity and gene expression status, we identified functional homologs likely to be involved in xylem developmental and biosynthetic processes in Populus and Eucalyptus. Our study suggests that, while genes involved in secondary cell wall biosynthesis show high levels of gene expression conservation, differential regulation of some xylem development genes may give rise to unique xylem properties. © 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.
Vinya, Royd; Malhi, Yadvinder; Brown, Nick D; Fisher, Joshua B; Brodribb, Timothy; Aragão, Luiz E O C
2018-06-15
Water availability has frequently been linked to seasonal leaf display in seasonally dry ecosystems, but there have been few ecohydrological investigations of this link. Miombo woodland is a dominant seasonally dry tropical forest ecosystem type in southern Africa; however, there are few data on the relationship between seasonal dynamics in plant-water relations and patterns of leaf display for Miombo woodland. Here we investigate this relationship among nine key Miombo woodland tree species differing in drought tolerance ability and leaf phenology. Results of this study showed that seasonal patterns of leaf phenology varied significantly with seasonal changes in stem water relations among the nine species. Leaf shedding coincided with the attainment of seasonal minimum stem water potential. Leaf flush occurred following xylem rehydration at the peak of the dry season suggesting that endogenous plant factors play a pivotal role in seasonal leaf display in this forest type. Drought-tolerant deciduous species suffered significantly higher seasonal losses in xylem hydraulic conductivity than the drought-intolerant semi-evergreen tree species (P < 0.05). There was a significant and positive correlation between species drought tolerance index and species' seasonal loss in hydraulic conductivity (P < 0.05), confirming the ecological role of long-distance xylem transport in this seasonally dry tropical forest. Our results reveal that water stress in seasonally dry tropical forests selects for water conservative traits that protect the vulnerable xylem transport system. Therefore, seasonal rhythms in xylem transport dictate patterns of leaf display in seasonally dry tropical forests.
Chen, Hong; Wang, Haiyang; Liu, Yanfang; Dong, Li
2013-01-01
1. The terminal shoot (or current-year shoot), as one of the most active parts on a woody plant, is a basic unit determining plant height and is potentially influenced by a variety of environmental factors. It has been predicted that tissues amount and their allocation in plant stems may play a critical role in determining plant size in alpine regions. The primary structure in terminal shoots is a key to our understanding treeline formation. The existing theories on treeline formation, however, are still largely lacking of evidence at the species level, much less from anatomy for the terminal shoot. 2. The primary structures within terminal shoot were measured quantitatively for 100 species from four elevation zones along the eastern slope of Gongga Mountain, southwestern China; one group was sampled from above the treeline. An allometric approach was employed to examine scaling relationships interspecifically, and a principal components analysis (PCA) was performed to test the relation among primary xylem, ground tissue, species growth form and altitude. 3. The results showed that xylem tissue size was closely correlated with ground tissue size isometrically across species, while undergoing significant y- or/and x-intercept shift in response to altitudinal belts. Further, a conspicuous characteristic of terminal shoot was its allocation of contrasting tissues between primary xylem and ground tissues with increasing elevation. The result of the PCA showed correlations between anatomical variation, species growth form/height classes and environment. 4. The current study presents a comparative assessment of the allocation of tissue in terminal shoot across phylogenically and ecologically diverse species, and analyzes tissue, function and climate associations with plant growth forms and height classes among species. The interspecific connection between primary xylem ratio and plant size along an elevation gradient suggests the importance of primary xylem in explaining the treeline formation.
Chen, Hong; Wang, Haiyang; Liu, Yanfang; Dong, Li
2013-01-01
1. The terminal shoot (or current-year shoot), as one of the most active parts on a woody plant, is a basic unit determining plant height and is potentially influenced by a variety of environmental factors. It has been predicted that tissues amount and their allocation in plant stems may play a critical role in determining plant size in alpine regions. The primary structure in terminal shoots is a key to our understanding treeline formation. The existing theories on treeline formation, however, are still largely lacking of evidence at the species level, much less from anatomy for the terminal shoot. 2. The primary structures within terminal shoot were measured quantitatively for 100 species from four elevation zones along the eastern slope of Gongga Mountain, southwestern China; one group was sampled from above the treeline. An allometric approach was employed to examine scaling relationships interspecifically, and a principal components analysis (PCA) was performed to test the relation among primary xylem, ground tissue, species growth form and altitude. 3. The results showed that xylem tissue size was closely correlated with ground tissue size isometrically across species, while undergoing significant y- or/and x-intercept shift in response to altitudinal belts. Further, a conspicuous characteristic of terminal shoot was its allocation of contrasting tissues between primary xylem and ground tissues with increasing elevation. The result of the PCA showed correlations between anatomical variation, species growth form/height classes and environment. 4. The current study presents a comparative assessment of the allocation of tissue in terminal shoot across phylogenically and ecologically diverse species, and analyzes tissue, function and climate associations with plant growth forms and height classes among species. The interspecific connection between primary xylem ratio and plant size along an elevation gradient suggests the importance of primary xylem in explaining the treeline formation. PMID:23658621
Xylem and phloem phenology in co-occurring conifers exposed to drought.
Swidrak, Irene; Gruber, Andreas; Oberhuber, Walter
2014-01-01
Variability in xylem and phloem phenology among years and species is caused by contrasting temperatures prevailing at the start of the growing season and species-specific sensitivity to drought. The focus of this study was to determine temporal dynamics of xylem and phloem formation in co-occurring deciduous and evergreen coniferous species in a dry inner Alpine environment (750 m a.s.l., Tyrol, Austria). By repeated micro-sampling of the stem, timing of key phenological dates of xylem and phloem formation was compared among mature Pinus sylvestris , Larix decidua and Picea abies during two consecutive years. Xylem formation in P. sylvestris started in mid and late April 2011 and 2012, respectively, and in both years about 2 week later in P. abies and L. decidua . Phloem formation preceded xylem formation on average by 3 week in P. sylvestris , and c . 5 week in P. abies and L. decidua . Based on modeled cell number increase, tracheid production peaked between early through late May 2011 and late May through mid-June 2012. Phloem formation culminated between late April and mid-May in 2011 and in late May 2012. Production of xylem and phloem cells continued for about 4 and 5-6 months, respectively. High variability in xylem increment among years and species is related to exogenous control by climatic factors and species-specific sensitivity to drought, respectively. On the other hand, production of phloem cells was quite homogenous and showed asymptotic decrease with respect to xylem cells indicating endogenous control. Results indicate that onset and culmination of xylem and phloem formation are controlled by early spring temperature, whereby strikingly advanced production of phloem compared to xylem cells suggests lower temperature requirement for initiation of the former.
Martin, Barry A.; Tolbert, N. E.
1983-01-01
Phosphate in the xylem exudate of tomato (Lycopersicon esculentum) plants was 70 to 98% inorganic phosphate (Pi), 2 to 30% P-choline, and less than 1% P-ethanolamine. Upon adding 32Pi to the nutrient, Pi in xylem exudate had the same specific activity within 4 hours. P-choline and P-ethanolamine reached the same specific activity only after 96 hours. The amount of Pi in xylem exudate was dependent on Pi concentration in the nutrient and decreased from 1700 to 170 micromolar when Pi in the nutrient decreased from 50 to 2 micromolar. The flux of 0.4 nmoles organic phosphate per minute per gram fresh weight root into the xylem exudate was not affected by the Pi concentration in the nutrient solution unless it was below 1 micromolar. During 7 days of Pi starvation, Pi in the xylem exudate decreased from 1400 to 130 micromolar while concentrations of the two phosphate esters remained unchanged. The concentration of phosphate esters in the xylem exudate was increased by addition of choline or ethanolamine to the nutrient solution, but Pi remained unchanged. Upon adding [14C]choline to the nutrient, 10 times more [14C]P-choline than [14C]choline was in the xylem exudate and 85 to 90% of the ester phosphate was P-choline. When [14C]ethanolamine was added, [14C]P-ethanolamine and [14C]ethanolamine in the xylem sap were equal in amount. P-choline and P-ethanolamine accumulated in leaves of whole plants at the same time and the same proportion as observed for their flux into the xylem exudate. No relationship between the transport of P-choline and Pi in the xylem was established. Rather, the amount of choline in xylem exudate and its incorporation into phosphatidylcholine in the leaf suggest that the root is a site of synthesis of P-choline and P-ethanolamine for phospholipid synthesis in tomato leaves. PMID:16663240
NASA Astrophysics Data System (ADS)
Ewers, B. E.; Pendall, E.; Reed, D. E.; Barnard, H. R.; Whitehouse, F.; Frank, J. M.; Massman, W. J.; Brooks, P. D.; Biederman, J. A.; Harpold, A. A.; Naithani, K. J.; Mitra, B.; Mackay, D. S.; Norton, U.; Borkhuu, B.
2011-12-01
While mountainous areas are critical for providing numerous ecosystem benefits at the regional scale, the strong gradients in environmental controls make predictions difficult. A key part of the problem is quantifying and predicting the feedback between mountain gradients and plant function which then controls ecosystem cycling. The emerging theory of plant hydraulics provides a rigorous yet simple platform from which to generate testable hypotheses and predictions of ecosystem pools and fluxes. Plant hydraulic theory predicts that plant controls over carbon, water, energy and nutrient fluxes can be derived from the limitation of plant water transport from the soil through xylem and out of stomata. In addition, the limit to plant water transport can be predicted by combining plant structure (e.g. xylem diameters or root-to-shoot ratios) and plant function (response of stomatal conductance to vapor pressure deficit or root vulnerability to cavitation). We evaluate the predictions of the plant hydraulic theory by testing it against data from a mountain gradient encompassing sagebrush steppe through subalpine forests (2700 to 3400 m). We further test the theory by predicting the carbon, water and nutrient exchanges from several coniferous trees in the same gradient that are dying from xylem dysfunction caused by blue-stain fungi carried by bark beetles. The common theme of both of these data sets is a change in water limitation caused by either changing precipitation along the mountainous gradient or lack of access to soil water from xylem-occluding fungi. Across all of the data sets which range in scale from individual plants to hillslopes, the data fit the predictions of plant hydraulic theory. Namely, there was a proportional tradeoff between the reference canopy stomatal conductance to water vapor and the sensitivity of that conductance to vapor pressure deficit that quantitatively fits the predictions of plant hydraulic theory. Incorporating this result into whole plant mass and energy exchange models allows prediction of plant carbon, energy and nitrogen exchange that fits recently collected data including plant sap flux, leaf gas exchange, eddy covariance towers and stand and watershed-scale biogeochemistry measurements. The results of our work will allow the next generation of ecosystem to regional scale coupled-biogeochemistry models to incorporate a simple plant hydraulic mechanism that will enable defensible predictions of carbon, water, energy and nutrient cycling with changing climate and land use.
Niu, Junfang; Chen, Fanjun; Mi, Guohua; Li, Chunjian; Zhang, Fusuo
2007-01-01
Background and Aims The influence of two nitrogen (N) levels on growth, water relations, and N uptake and flow was investigated in two different inbred lines of maize (N-efficient Zi330 and N-inefficient Chen94-11) to analyse the differences in N uptake and cycling within a plant. Methods Xylem sap from different leaves of the inbred lines cultured in quartz sand was collected by application of pressure to the root system. Plant transpiration was measured on a daily basis by weighing five pots of each of the treatments. Key Results N-efficient Zi330 had a higher relative growth rate and water-use efficiency at both high (4 mm) and low (0·08 mm) N levels. At a high N level, the amount of N taken up was similar for the two inbred lines; the amount of N transported in the xylem and retranslocated in the phloem was slight greater in Chen94-11 than in Zi330. At a low N level, however, the total amount of N taken up, transported in the xylem and retranslocated in the phloem of Zi330 was 2·2, 2·7 and 2·7 times more, respectively, than that of Chen94-11. Independent of inbred line and N level, the amounts of N transported in the xylem and cycled in the phloem were far more than that taken up by roots at the same time. Low N supply shifted NO3−1 reduction towards the roots. The major nitrogenous compound in the xylem sap was NO3−1, when plants grew at the high N level, while amino acid-N was predominant when plants grew at the low N level. Conclusions The N-efficient maize inbred line Zi330 had a higher ability to take up N and cycle N within the plant than N-inefficient Chen94-11 when grown under N-deficiency. PMID:17088295
Federal Register 2010, 2011, 2012, 2013, 2014
2013-12-24
... DEPARTMENT OF COMMERCE Foreign-Trade Zones Board [B-106-2013] Notification of Proposed Production Activity, Xylem Water Systems USA LLC, Subzone 37D, (Centrifugal, Submersible Pumps and Related Components), Auburn, New York Xylem Water Systems USA LLC (Xylem), operator of Subzone 37D, submitted a notification of proposed production activity to...
The watering of tall trees--embolization and recovery.
Gouin, Henri
2015-03-21
We can propound a thermo-mechanical understanding of the ascent of sap to the top of tall trees thanks to a comparison between experiments associated with the cohesion-tension theory and the disjoining pressure concept for liquid thin-films. When a segment of xylem is tight-filled with crude sap, the liquid pressure can be negative although the pressure in embolized vessels remains positive. Examples are given that illustrate how embolized vessels can be refilled and why the ascent of sap is possible even in the tallest trees avoiding the problem due to cavitation. However, the maximum height of trees is limited by the stability domain of liquid thin-films. Copyright © 2015 Elsevier Ltd. All rights reserved.
Rico, Christopher; Pittermann, Jarmila; Polley, H Wayne; Aspinwall, Michael J; Fay, Phillip A
2013-09-01
Plant gas exchange is regulated by stomata, which coordinate leaf-level water loss with xylem transport. Stomatal opening responds to internal concentrations of CO₂ in the leaf, but changing CO₂ can also lead to changes in stomatal density that influence transpiration. Given that stomatal conductance increases under subambient concentrations of CO₂ and, conversely, that plants lose less water at elevated concentrations, can downstream effects of atmospheric CO₂ be observed in xylem tissue? We approached this problem by evaluating leaf stomatal density, xylem transport, xylem anatomy and resistance to cavitation in Helianthus annuus plants grown under three CO₂ regimes ranging from pre-industrial to elevated concentrations. Xylem transport, conduit size and stomatal density all increased at 290 ppm relative to ambient and elevated CO₂ concentrations. The shoots of the 290-ppm-grown plants were most vulnerable to cavitation, whereas xylem cavitation resistance did not differ in 390- and 480-ppm-grown plants. Our data indicate that, even as an indirect driver of water loss, CO₂ can affect xylem structure and water transport by coupling stomatal and xylem hydraulic functions during plant development. This plastic response has implications for plant water use under variable concentrations of CO₂, as well as the evolution of efficient xylem transport. © 2013 The Authors. New Phytologist © 2013 New Phytologist Trust.
A broad survey of hydraulic and mechanical safety in the xylem of conifers
Pauline S. Bouche; Maximilien Larter; Jean-Christophe Domec; Regis Burlett; Peter Gasson; Steven Jansen; Sylvain Delzon
2014-01-01
Drought-induced forest dieback has been widely reported over the last decades, and the evidence for a direct causal link between survival and hydraulic failure (xylem cavitation) is now well known. Because vulnerability to cavitation is intimately linked to the anatomy of the xylem, the main objective of this study was to better understand the xylem anatomical...
Xylem formation can be modeled statistically as a function of primary growth and cambium activity.
Huang, Jian-Guo; Deslauriers, Annie; Rossi, Sergio
2014-08-01
Primary (budburst, foliage and shoot) growth and secondary (cambium and xylem) growth of plants play a vital role in sequestering atmospheric carbon. However, their potential relationships have never been mathematically quantified and the underlying physiological mechanisms are unclear. We monitored primary and secondary growth in Picea mariana and Abies balsamea on a weekly basis from 2010 to 2013 at four sites over an altitudinal gradient (25-900 m) in the eastern Canadian boreal forest. We determined the timings of onset and termination through the fitted functions and their first derivative. We quantified the potential relationships between primary growth and secondary growth using the mixed-effects model. We found that xylem formation of boreal conifers can be modeled as a function of cambium activity, bud phenology, and shoot and needle growth, as well as species- and site-specific factors. Our model reveals that there may be an optimal mechanism to simultaneously allocate the photosynthetic products and stored nonstructural carbon to growth of different organs at different times in the growing season. This mathematical link can bridge phenological modeling, forest ecosystem productivity and carbon cycle modeling, which will certainly contribute to an improved prediction of ecosystem productivity and carbon equilibrium. © 2014 The Authors. New Phytologist © 2014 New Phytologist Trust.
Lateral Movement of Water and Sugar Across Xylem in Sugarcane Stalks
Bull, T. A.; Gayler, K. R.; Glasziou, K. T.
1972-01-01
Laterally connected vascular bundles in the nodes of sugarcane (Saccharum species cv. Pindar) stalks allow a rapid redistribution of water across the stalk should the vascular continuity be partly disrupted. Tritiated water supplied to the roots exchanged rapidly between the xylem and storage tissue so that net movement up the stalk was slow. The half-time for exchange in a labeled stalk was about 4 hours so that the entire water content of a sugarcane stalk can turn over at least once in a single day. No rapid flux of sugar between xylem and phloem or xylem and storage tissue was detected. Functional xylem contained only low sugar concentrations: less than 0.3% w/v in the stalk and less than 0.02% w/v in the leaf. Previous reports of high sugar levels (9% w/v) in sugarcane stalk xylem reflect some degree of xylem blockage followed by a slow equilibration with free space sugars in the storage tissue. PMID:16658067
Kunert, Norbert
2016-10-20
Daily xylem sap flux values (daily J s ) and maximum xylem sap flux values (max J s ) from 125 tropical trees from different study sites in the Neotropics were compared. A cross species and study site relationship was found between daily and maximum values. The relationship can be expressed as daily J s =6.5x max J s . The geometrical relationship between the maximum xylem sap flux of a given day is thus defining the daily xylem sap flux rates. Assuming a bell-shaped diurnal sap flux course and a relatively constant day length the maximum xylem sap flux is the only possible changing variable to define daily fluxes. Further, this relationship is showing the inertia of the xylem sap flux as a physical object and highlights the delayed response to environmental changes and its subsequent inevitable susceptibility under environmental stress to hydraulic failure. Copyright © 2016 Elsevier GmbH. All rights reserved.
Lens, Frederic; Vos, Rutger A.; Charrier, Guillaume; van der Niet, Timo; Merckx, Vincent; Baas, Pieter; Aguirre Gutierrez, Jesus; Jacobs, Bart; Chacon Dória, Larissa; Smets, Erik; Delzon, Sylvain; Janssens, Steven B.
2016-01-01
Background and Aims Angiosperms with simple vessel perforations have evolved many times independently of species having scalariform perforations, but detailed studies to understand why these transitions in wood evolution have happened are lacking. We focus on the striking difference in wood anatomy between two closely related genera of Adoxaceae, Viburnum and Sambucus, and link the anatomical divergence with climatic and physiological insights. Methods After performing wood anatomical observations, we used a molecular phylogenetic framework to estimate divergence times for 127 Adoxaceae species. The conditions under which the genera diversified were estimated using ancestral area reconstruction and optimization of ancestral climates, and xylem-specific conductivity measurements were performed. Key Results Viburnum, characterized by scalariform vessel perforations (ancestral), diversified earlier than Sambucus, having simple perforations (derived). Ancestral climate reconstruction analyses point to cold temperate preference for Viburnum and warm temperate for Sambucus. This is reflected in the xylem-specific conductivity rates of the co-occurring species investigated, showing that Viburnum lantana has rates much lower than Sambucus nigra. Conclusions The lack of selective pressure for high conductive efficiency during early diversification of Viburnum and the potentially adaptive value of scalariform perforations in frost-prone cold temperate climates have led to retention of the ancestral vessel perforation type, while higher temperatures during early diversification of Sambucus have triggered the evolution of simple vessel perforations, allowing more efficient long-distance water transport. PMID:27498812
Xylem development in prunus flower buds and the relationship to deep supercooling.
Ashworth, E N
1984-04-01
Xylem development in eight Prunus species was examined and the relationship to deep supercooling assessed. Dormant buds of six species, P. armeniaca, P. avium, P. cerasus, P. persica, P. salicina, and P. sargentii deep supercooled. Xylem vessel elements were not observed within the dormant floral primordia of these species. Instead, discrete bundles containing procambial cells were observed. Vascular differentiation resumed and xylem continuity was established during the time that the capacity to deep supercool was lost. In P. serotina and P. virginiana, two species which do not supercool, xylem vessels ran the length of the inflorescence and presumably provided a conduit for the spread of ice into the bud. The results support the hypothesis that the lack of xylem continuity is an important feature of buds which deep supercool.
Helfter, Carole; Shephard, Jonathon D; Martinez-Vilalta, Jordi; Mencuccini, Maurizio; Hand, Duncan P
2007-02-01
Over the past 70 years, heat has been widely used as a tracer for estimating the flow of water in woody and herbaceous plants. However, most commercially available techniques for monitoring whole plant water use are invasive and the measurements are potentially flawed because of wounding of the xylem tissue. The study of photosynthate transport in the phloem remains in its infancy, and little information about phloem transport rates is available owing to the fragility of the vascular tissue. The aim of our study was to develop a compact, stand-alone non-invasive system allowing for direct detection of phloem and xylem sap movement. The proposed method uses a heat pulse as a tracer for sap flow. Heat is applied to the surface of the stem with a near-infrared laser source, and heat propagation is monitored externally by means of an infrared camera. Heat pulse velocities are determined from the thermometric data and related to the more useful quantity, mass flow rate. Simulation experiments on the xylem tissue of severed silver birch (Betula pendula Roth.) branch segments were performed to assess the feasibility of the proposed approach, highlight the characteristics of the technique and outline calibration strategies. Good agreement between imposed and measured flow rates was achieved leading to experimentation with live silver birch and oak (Quercus robur L.) saplings. It was demonstrated that water flow through xylem vessels can be monitored non-invasively on an intact stem with satisfactory accuracy despite simultaneous sugar transport in the phloem. In addition, it was demonstrated that the technique allows for unequivocal detection of phloem flow velocities.
Rapid systemic up-regulation of genes after heat-wounding and electrical stimulation
NASA Technical Reports Server (NTRS)
Davies, E.; Vian, A.; Vian, C.; Stankovic, B.
1997-01-01
When one leaf of a tomato plant is electrically-stimulated or heat-wounded, proteinase inhibitor genes are rapidly up-regulated in distant leaves. The identity of the systemic wound signal(s) is not yet known, but major candidates include hormones transmitted via the phloem or the xylem, the electrically-stimulated self-propagating electrical signal in the phloem (the action potential, AP), or the heat-wound-induced surge in hydraulic pressure in the xylem evoking a local change in membrane potential in adjacent living cells (the variation potential, VP). In order to discriminate between these signals we have adopted two approaches. The first approach involves applying stimuli that evoke known signals and determining whether these signals have similar effects on the "model" transcripts for proteinase inhibitors (pin) and calmodulin (cal). Here we show that a heat wound almost invariably evokes a VP, while an electrical stimulation occasionally evokes an AP, and both of these signals induce accumulation of transcripts encoding proteinase inhibitors. The second approach involves identifying the array of genes turned on by heat-wounding. To this end, we have constructed a subtractive library for heat-wounded tissue, isolated over 800 putatively up-regulated clones, and shown that all but two of the fifty that we have analyzed by Northern hybridization are, indeed, up-regulated. Here we show the early kinetics of up-regulation of three of these transcripts in the terminal (4th) leaf in response to heat-wounding the 3rd leaf, about 5 cm away. Even though these transcripts show somewhat different time courses of induction, with one peaking at 30 min, another at 15 min, and another at 5 min after flaming of a distant leaf, they all exhibit a similar pattern, i.e., a transient period of transcript accumulation preceding a period of transcript decrease, followed by a second period of transcript accumulation.
NASA Astrophysics Data System (ADS)
Martín Gómez, Paula; Aguilera, Mònica; Pemán, Jesús; Gil Pelegrín, Eustaquio; Ferrio, Juan Pedro
2014-05-01
As a general rule, no isotopic fractionation occurs during water uptake and water transport, thus, xylem water reflects source water. However, this correspondence does not always happen. Isotopic enrichment of xylem water has been found in several cases and has been either associated to 'stem processes' like cuticular evaporation 1 and xylem-phloem communication under water stress 2,3 or to 'soil processes' such as species-specific use of contrasting water sources retained at different water potential forces in soil. In this regard, it has been demonstrated that mobile and tightly-bound water may show different isotopic signature 4,5. However, standard cryogenic distillation does not allow to separate different water pools within soil samples. Here, we carried out a study in a mixed adult forest (Pinus sylvestris, Quercus subpyrenaica and Buxus sempervirens) growing in a relatively deep loamy soil in the Pre-Pyrenees. During one year, we sampled xylem from twigs and soil at different depths (10, 30 and 50 cm). We also sampled xylem from trunk and bigger branches to assess whether xylem water was enriched in the distal parts of the tree. We found average deviations in the isotopic signature from xylem to soil of 4o 2o and 2.4o in δ18O and 18.3o 7.3o and 8.9o in δ2H, for P.sylvestris, Q.subpyrenaica and B.sempervirens respectively. Xylem water was always enriched compared to soil. In contrast, we did not find clear differences in isotopic composition between xylem samples along the tree. Declining the hypothesis that 'stem processes' would cause these uncoupling between soil and xylem isotopic values, we tested the possibility to separate mobile and tightly-bound water by centrifugation. Even though we could separate two water fractions in soils close to saturation, we could not recover a mobile fraction in drier soils. In this regard, we welcome suggestions on alternatives to separate different soil fractions in order to find the correspondence between soil and xylem water. References 1. Dawson, T. E. & Ehleringer, J. R. Isotopic enrichment of water in the 'woody' tissues of plants: Implications for plant water source, water uptake, and other studies which use the stable isotopic composition of cellulose. (1993). 2. Cernusak, L. a, Farquhar, G. D. & Pate, J. S. Environmental and physiological controls over oxygen and carbon isotope composition of Tasmanian blue gum, Eucalyptus globulus. Tree Physiol. 25, 129-46 (2005). 3. Bertrand, G. et al. Determination of spatiotemporal variability of tree water uptake using stable isotopes (δ 18 O, δ 2 H) in an alluvial system supplied by a high-altitude watershed, Pfyn forest, Switzerland. Ecohydrology (2012). doi:10.1002/eco.1347 4. Tang, K. & Feng, X. The effect of soil hydrology on the oxygen and hydrogen isotopic compositions of plants ' source water. 185, (2001). 5. Brooks, J. R., Barnard, H. R., Coulombe, R. & McDonnell, J. J. Ecohydrologic separation of water between trees and streams in a Mediterranean climate. Nat. Geosci. 3, 100-104 (2009). Acknowledgements This study was funded by RESILFOR project (AGL 2012-40039-C02-02) and FPU fellowship from the Spanish Ministry of Science and Innovation (FPU12/00648). We thank Instituto de Formación Agroambiental de Jaca and Unidad de Salud de los Bosques de Aragón for their support on field work and we feel very grateful to Miguel Ángel Lázaro for climbing the studied trees, José María Alcaire for one year of rain collection and Pilar Sopeña and Ma Josep Pau for laboratory analysis. Helpful comments by Jordi Voltas on statistical analysis have improved the quality of the work.
4th International Plant Biomechanics Conference Proceedings (Abstracts)
DOE Office of Scientific and Technical Information (OSTI.GOV)
Frank W. Telewski; Lothar H. Koehler; Frank W. Ewers
2003-07-20
The 4th International Plant Biomechanics Conference facilitated an interdisciplinary exchange between scientists, engineers, and educators addressing the major questions encountered in the field of Plant Biomechanics. Subjects covered by the conference include: Evolution; Ecology; Mechanoreception; Cell Walls; Genetic Modification; Applied Biomechanics of Whole Plants, Plant Products, Fibers & Composites; Fluid Dynamics; Wood & Trees; Fracture Mechanics; Xylem Pressure & Water Transport; Modeling; and Introducing Plant Biomechanics in Secondary School Education.
Zhang, Yong-Jiang; Rockwell, Fulton E.; Wheeler, James K.; Holbrook, N. Michele
2014-01-01
Declines in leaf hydraulic conductance (Kleaf) with increasing water stress have been attributed to cavitation of the leaf xylem. However, in the leaves of conifers, the reversible collapse of transfusion tracheids may provide an alternative explanation. Using Taxus baccata, a conifer species without resin, we developed a modified rehydration technique that allows the separation of declines in Kleaf into two components: one reversible and one irreversible upon relaxation of water potential to −1 MPa. We surveyed leaves at a range of water potentials for evidence of cavitation using cryo-scanning electron microscopy and quantified dehydration-induced structural changes in transfusion tracheids by cryo-fluorescence microscopy. Irreversible declines in Kleaf did not occur until leaf water potentials were more negative than −3 MPa. Declines in Kleaf between −2 and −3 MPa were reversible and accompanied by the collapse of transfusion tracheids, as evidenced by cryo-fluorescence microscopy. Based on cryo-scanning electron microscopy, cavitation of either transfusion or xylem tracheids did not contribute to declines in Kleaf in the reversible range. Moreover, the deformation of transfusion tracheids was quickly reversible, thus acting as a circuit breaker regulating the flux of water through the leaf vasculature. As transfusion tissue is present in all gymnosperms, the reversible collapse of transfusion tracheids may be a general mechanism in this group for the protection of leaf xylem from excessive loads generated in the living leaf tissue. PMID:24948828
Protein and metabolite composition of xylem sap from field-grown soybeans (Glycine max).
Krishnan, Hari B; Natarajan, Savithiry S; Bennett, John O; Sicher, Richard C
2011-05-01
The xylem, in addition to transporting water, nutrients and metabolites, is also involved in long-distance signaling in response to pathogens, symbionts and environmental stresses. Xylem sap has been shown to contain a number of proteins including metabolic enzymes, stress-related proteins, signal transduction proteins and putative transcription factors. Previous studies on xylem sap have mostly utilized plants grown in controlled environmental chambers. However, plants in the field are subjected to high light and to environmental stress that is not normally found in growth chambers. In this study, we have examined the protein and metabolite composition of xylem sap from field-grown cultivated soybean plants. One-dimensional gel electrophoresis of xylem sap from determinate, indeterminate, nodulating and non-nodulating soybean cultivars revealed similar protein profiles consisting of about 8-10 prominent polypeptides. Two-dimensional gel electrophoresis of soybean xylem sap resulted in the visualization of about 60 distinct protein spots. A total of 38 protein spots were identified using MALDI-TOF MS and LC-MS/MS. The most abundant proteins present in the xylem sap were identified as 31 and 28 kDa vegetative storage proteins. In addition, several proteins that are conserved among different plant species were also identified. Diurnal changes in the metabolite profile of xylem sap collected during a 24-h cycle revealed that asparagine and aspartate were the two predominant amino acids irrespective of the time collected. Pinitol (D-3-O-methyl-chiro-inositol) was the most abundant carbohydrate present. The possible roles of xylem sap proteins and metabolites as nutrient reserves for sink tissue and as an indicator of biotic stress are also discussed.
Choat, Brendan; Brodersen, Craig R; McElrone, Andrew J
2015-02-01
The formation of emboli in xylem conduits can dramatically reduce hydraulic capacity and represents one of the principal mechanisms of drought-induced mortality in woody plants. However, our understanding of embolism formation and repair is constrained by a lack of tools to directly and nondestructively measure these processes at high spatial resolution. Using synchrotron-based microcomputed tomography (microCT), we examined embolism in the xylem of coast redwood (Sequoia sempervirens) saplings that were subjected to cycles of drought and rewatering. Embolism formation was observed occurring by three different mechanisms: as tracheids embolizing in wide tangential bands; as isolated tracheids in seemingly random events; and as functional groups connected to photosynthetic organs. Upon rewatering, stem water potential recovered to predrought stress levels within 24 h; however, no evidence of embolism repair was observed even after a further 2 wk under well-watered conditions. The results indicate that intertracheid air seeding is the primary mechanism by which embolism spreads in the xylem of S. sempervirens, but also show that a small number of tracheids initially become gas-filled via another mechanism. The inability of S. sempervirens saplings to reverse drought-induced embolism is likely to have important ecological impacts on this species. © 2014 The Authors. New Phytologist © 2014 New Phytologist Trust.
Uroic, M. Kalle; Salaün, Pascal; Raab, Andrea; Feldmann, Jörg
2012-01-01
Arsenic uptake and translocation studies on xylem sap focus generally on the concentration and speciation of arsenic in the xylem. Arsenic impact on the xylem sap metabolite profile and its production during short term exposure has not been reported in detail. To investigate this, cucumbers were grown hydroponically and arsenate (AsV) and DMA were used for plant treatment for 24 h. Total arsenic and arsenic speciation in xylem sap was analyzed including a metabolite profiling under AsV stress. Produced xylem sap was quantified and absolute arsenic transported was determined. AsV exposure had a significant impact on the metabolite profile of xylem sap. Four m/z values corresponding to four compounds were up-regulated, one compound down-regulated by AsV exposure. The compound down-regulated was identified to be isoleucine. Furthermore, AsV exposure had a significant influence on sap production, leading to a reduction of up to 96% sap production when plants were exposed to 1000 μg kg−1 AsV. No difference to control plants was observed when plants were exposed to 1000 μg kg−1 DMA. Absolute arsenic amount in xylem sap was the lowest at high AsV exposure. These results show that AsV has a significant impact on the production and metabolite profile of xylem sap. The physiological importance of isoleucine needs further attention. PMID:22536187
Mobile Gibberellin Directly Stimulates Arabidopsis Hypocotyl Xylem Expansion[W][OA
Ragni, Laura; Nieminen, Kaisa; Pacheco-Villalobos, David; Sibout, Richard; Schwechheimer, Claus; Hardtke, Christian S.
2011-01-01
Secondary growth of the vasculature results in the thickening of plant structures and continuously produces xylem tissue, the major biological carbon sink. Little is known about the developmental control of this quantitative trait, which displays two distinct phases in Arabidopsis thaliana hypocotyls. The later phase of accelerated xylem expansion resembles the secondary growth of trees and is triggered upon flowering by an unknown, shoot-derived signal. We found that flowering-dependent hypocotyl xylem expansion is a general feature of herbaceous plants with a rosette growth habit. Flowering induction is sufficient to trigger xylem expansion in Arabidopsis. By contrast, neither flower formation nor elongation of the main inflorescence is required. Xylem expansion also does not depend on any particular flowering time pathway or absolute age. Through analyses of natural genetic variation, we found that ERECTA acts locally to restrict xylem expansion downstream of the gibberellin (GA) pathway. Investigations of mutant and transgenic plants indicate that GA and its signaling pathway are both necessary and sufficient to directly trigger enhanced xylogenesis. Impaired GA signaling did not affect xylem expansion systemically, suggesting that it acts downstream of the mobile cue. By contrast, the GA effect was graft transmissible, suggesting that GA itself is the mobile shoot-derived signal. PMID:21498678
Knipfer, Thorsten; Cuneo, Italo F; Brodersen, Craig R; McElrone, Andrew J
2016-06-01
Gas embolisms formed during drought can disrupt long-distance water transport through plant xylem vessels, but some species have the ability to remove these blockages. Despite evidence suggesting that embolism removal is linked to the presence of vessel-associated parenchyma, the underlying mechanism remains controversial and is thought to involve positive pressure generated by roots. Here, we used in situ x-ray microtomography on excised grapevine stems to determine if embolism removal is possible without root pressure, and if the embolism formation/removal affects vessel functional status after sample excision. Our data show that embolism removal in excised stems was driven by water droplet growth and was qualitatively identical to refilling in intact plants. When stem segments were rehydrated with H2O after excision, vessel refilling occurred rapidly (<1 h). The refilling process was substantially slower when polyethylene glycol was added to the H2O source, thereby providing new support for an osmotically driven refilling mechanism. In contrast, segments not supplied with H2O showed no refilling and increased embolism formation. Dynamic changes in liquid/wall contact angles indicated that the processes of embolism removal (i.e. vessel refilling) by water influx and embolism formation by water efflux were directly linked to the activity of vessel-associated living tissue. Overall, our results emphasize that root pressure is not required as a driving force for vessel refilling, and care should be taken when performing hydraulics measurements on excised plant organs containing living vessel-associated tissue, because the vessel behavior may not be static. © 2016 American Society of Plant Biologists. All Rights Reserved.
Knipfer, Thorsten; Cuneo, Italo F.; Brodersen, Craig R.; McElrone, Andrew J.
2016-01-01
Gas embolisms formed during drought can disrupt long-distance water transport through plant xylem vessels, but some species have the ability to remove these blockages. Despite evidence suggesting that embolism removal is linked to the presence of vessel-associated parenchyma, the underlying mechanism remains controversial and is thought to involve positive pressure generated by roots. Here, we used in situ x-ray microtomography on excised grapevine stems to determine if embolism removal is possible without root pressure, and if the embolism formation/removal affects vessel functional status after sample excision. Our data show that embolism removal in excised stems was driven by water droplet growth and was qualitatively identical to refilling in intact plants. When stem segments were rehydrated with H2O after excision, vessel refilling occurred rapidly (<1 h). The refilling process was substantially slower when polyethylene glycol was added to the H2O source, thereby providing new support for an osmotically driven refilling mechanism. In contrast, segments not supplied with H2O showed no refilling and increased embolism formation. Dynamic changes in liquid/wall contact angles indicated that the processes of embolism removal (i.e. vessel refilling) by water influx and embolism formation by water efflux were directly linked to the activity of vessel-associated living tissue. Overall, our results emphasize that root pressure is not required as a driving force for vessel refilling, and care should be taken when performing hydraulics measurements on excised plant organs containing living vessel-associated tissue, because the vessel behavior may not be static. PMID:27208267
Hatcher, P.G.; Lerch, H. E.; Verheyen, T.V.
1990-01-01
It is generally recognized that xylem from trees that are buried in peat swamps is transformed first to huminite macerals in brown coal and then to vitrinite macerals in bituminous coal by processes collectively known as coalification. In order to understand the chemical nature of coalification of xylem and the chemical structures that eventually evolve in coal, we examined a series of gymnospermous xylem samples coalified to varying degrees. The samples included modern fresh xylem, modern degraded xylem in peat, and xylem coalified to ranks of brown coal (lignite B), lignite A, and subbituminous coal. The organic geochemical methods used in this study included solid-state 13C nuclear magnetic resonance (NMR) and pyrolysis/gas chromatography/mass spectrometry. The NMR method provided average compositional information, and the pyrolysis provided detailed molecular information. Although the samples examined include different plants of different geologic ages, they all share a common feature in that they are gymnospermous and presumably have or had a similar kind of lignin. The data obtained in this study provide enough details to allow delineation of specific coalification pathway for the xylem is microbial degradation in peat (peatification), leading to selective removal of cellulosic components. These components constitute a large fraction of the total mass of xylem, usually greater than 50%. Although cellulosic components can survive degradation under certain conditions, their loss during microbial degradation is the rule rather than exception during peatification. As these components of xylem are degraded and lost, lignin, another major component of xylem, is selectively enriched because it is more resistant to microbial degradation than the cellulosic components. Thus, lignin survives peatification in a practically unaltered state and becomes the major precursor of coalified xylem. During its transformation to brown coal and lignite A, lignin in xylem is altered by two important processes. The first involves loss of methoxyl groups, primarily by demethylation (Fig. 1A). The end products of demethylation are catechol-like structures as shown below in Figure 1B. The second transformation process involves increased cross-linking of the aromatic rings. This cross-linking induces increased carbon substitution of the aromatic rings such that the lignin-derived structures become more highly condensed. During its conversion to coalified xylem in subbituminous coal, lignitic xylem, composed primarily of condensed catechol-like structures, is transformed to a macromolecular material primarily composed of phenol-like structures. The catechol-like structures of lignitic xylem loose a hydroxyl group, which is replaced by a hydrogen to form the phenol-like structure as shown in the example in Figure 1B. The pyrolysis data provided only a few clues as to the fate of the C3-side chain of lignin during coalification. However, the NMR data suggest that this side chain is altered, probably by loss of the hydroxyl groups that are attached in modern lignin. Interference in the NMR analysis by aliphatic components of wood, such as resins, precludes definitive determinations of the fate of the C3-side chain during coalification. ?? 1990.
Xylem Development in Prunus Flower Buds and the Relationship to Deep Supercooling
Ashworth, Edward N.
1984-01-01
Xylem development in eight Prunus species was examined and the relationship to deep supercooling assessed. Dormant buds of six species, P. armeniaca, P. avium, P. cerasus, P. persica, P. salicina, and P. sargentii deep supercooled. Xylem vessel elements were not observed within the dormant floral primordia of these species. Instead, discrete bundles containing procambial cells were observed. Vascular differentiation resumed and xylem continuity was established during the time that the capacity to deep supercool was lost. In P. serotina and P. virginiana, two species which do not supercool, xylem vessels ran the length of the inflorescence and presumably provided a conduit for the spread of ice into the bud. The results support the hypothesis that the lack of xylem continuity is an important feature of buds which deep supercool. Images Fig. 1 Fig. 2 Fig. 3 Fig. 4 Fig. 5 Fig. 6 PMID:16663523
Hajek, Peter; Kurjak, Daniel; von Wühlisch, Georg; Delzon, Sylvain; Schuldt, Bernhard
2016-01-01
In angiosperms, many studies have described the inter-specific variability of hydraulic-related traits and little is known at the intra-specific level. This information is however mandatory to assess the adaptive capacities of tree populations in the context of increasing drought frequency and severity. Ten 20-year old European beech (Fagus sylvatica L.) provenances representing the entire distribution range throughout Europe and differing significantly in aboveground biomass increment (ABI) by a factor of up to four were investigated for branch wood anatomical, hydraulic, and foliar traits in a provenance trial located in Northern Europe. We quantified to which extend xylem hydraulic and leaf traits are under genetic control and tested whether the xylem hydraulic properties (hydraulic efficiency and safety) trades off with yield and wood anatomical and leaf traits. Our results showed that only three out of 22 investigated ecophysiological traits showed significant genetic differentiations between provenances, namely vessel density (VD), the xylem pressure causing 88% loss of hydraulic conductance and mean leaf size. Depending of the ecophysiological traits measured, genetic differentiation between populations explained 0–14% of total phenotypic variation, while intra-population variability was higher than inter-population variability. Most wood anatomical traits and some foliar traits were additionally related to the climate of provenance origin. The lumen to sapwood area ratio, vessel diameter, theoretical specific conductivity and theoretical leaf-specific conductivity as well as the C:N-ratio increased with climatic aridity at the place of origin while the carbon isotope signature (δ13C) decreased. Contrary to our assumption, none of the wood anatomical traits were related to embolism resistance but were strong determinants of hydraulic efficiency. Although ABI was associated with both VD and δ13C, both hydraulic efficiency and embolism resistance were unrelated, disproving the assumed trade-off between hydraulic efficiency and safety. European beech seems to compensate increasing water stress with growing size mainly by adjusting vessel number and not vessel diameter. In conclusion, European beech has a high potential capacity to cope with climate change due to the high degree of intra-population genetic variability. PMID:27379112
Hajek, Peter; Kurjak, Daniel; von Wühlisch, Georg; Delzon, Sylvain; Schuldt, Bernhard
2016-01-01
In angiosperms, many studies have described the inter-specific variability of hydraulic-related traits and little is known at the intra-specific level. This information is however mandatory to assess the adaptive capacities of tree populations in the context of increasing drought frequency and severity. Ten 20-year old European beech (Fagus sylvatica L.) provenances representing the entire distribution range throughout Europe and differing significantly in aboveground biomass increment (ABI) by a factor of up to four were investigated for branch wood anatomical, hydraulic, and foliar traits in a provenance trial located in Northern Europe. We quantified to which extend xylem hydraulic and leaf traits are under genetic control and tested whether the xylem hydraulic properties (hydraulic efficiency and safety) trades off with yield and wood anatomical and leaf traits. Our results showed that only three out of 22 investigated ecophysiological traits showed significant genetic differentiations between provenances, namely vessel density (VD), the xylem pressure causing 88% loss of hydraulic conductance and mean leaf size. Depending of the ecophysiological traits measured, genetic differentiation between populations explained 0-14% of total phenotypic variation, while intra-population variability was higher than inter-population variability. Most wood anatomical traits and some foliar traits were additionally related to the climate of provenance origin. The lumen to sapwood area ratio, vessel diameter, theoretical specific conductivity and theoretical leaf-specific conductivity as well as the C:N-ratio increased with climatic aridity at the place of origin while the carbon isotope signature (δ(13)C) decreased. Contrary to our assumption, none of the wood anatomical traits were related to embolism resistance but were strong determinants of hydraulic efficiency. Although ABI was associated with both VD and δ(13)C, both hydraulic efficiency and embolism resistance were unrelated, disproving the assumed trade-off between hydraulic efficiency and safety. European beech seems to compensate increasing water stress with growing size mainly by adjusting vessel number and not vessel diameter. In conclusion, European beech has a high potential capacity to cope with climate change due to the high degree of intra-population genetic variability.
Redefining plant functional types for forests based on plant traits
NASA Astrophysics Data System (ADS)
Wei, L.; Xu, C.; Christoffersen, B. O.; McDowell, N. G.; Zhou, H.
2016-12-01
Our ability to predict forest mortality is limited by the simple plant functional types (PFTs) in current generations of Earth System models (ESMs). For example, forests were formerly separated into PFTs only based on leaf form and phenology across different regions (arctic, temperate, and tropic areas) in the Community Earth System Model (CESM). This definition of PFTs ignored the large variation in vulnerability of species to drought and shade tolerance within each PFT. We redefined the PFTs for global forests based on plant traits including phenology, wood density, leaf mass per area, xylem-specific conductivity, and xylem pressure at 50% loss of conductivity. Species with similar survival strategies were grouped into the same PFT. New PFTs highlighted variation in vulnerability and physiological adaptation to drought and shade. New PFTs were better clustered than old ones in the two-dimensional plane of the first two principle components in a principle component analysis. We expect that the new PFTs will strengthen ESMs' ability on predicting drought-induced mortality in the future.
Zheng, Huanquan
2015-01-01
Plant viruses move systemically in plants through the phloem. They move as virions or as ribonucleic protein complexes, although it is not clear what these complexes are made of. The approximately 10-kb RNA genome of Turnip mosaic virus (TuMV) encodes a membrane protein, known as 6K2, that induces endomembrane rearrangements for the formation of viral replication factories. These factories take the form of vesicles that contain viral RNA (vRNA) and viral replication proteins. In this study, we report the presence of 6K2-tagged vesicles containing vRNA and the vRNA-dependent RNA polymerase in phloem sieve elements and in xylem vessels. Transmission electron microscopy observations showed the presence in the xylem vessels of vRNA-containing vesicles that were associated with viral particles. Stem-girdling experiments, which leave xylem vessels intact but destroy the surrounding tissues, confirmed that TuMV could establish a systemic infection of the plant by going through xylem vessels. Phloem sieve elements and xylem vessels from Potato virus X-infected plants also contained lipid-associated nonencapsidated vRNA, indicating that the presence of membrane-associated ribonucleic protein complexes in the phloem and xylem may not be limited to TuMV. Collectively, these studies indicate that viral replication factories could end up in the phloem and the xylem. PMID:25717035
Does turgor limit growth in tall trees?
D.R. Woodruff; B.J. Bond; F.C. Meinzer
2004-01-01
The gravitational component of water potential contributes a standing 0.01 MPa m1 to the xylem tension gradient in plants. In tall trees, this contribution can significantly reduce the water potential near the tree tops. The turgor of cells in buds and leaves is expected to decrease in direct proportion with leaf water potential along a height gradient unless osmotic...
NASA Astrophysics Data System (ADS)
Mirfenderesgi, G.; Bohrer, G.; Matheny, A. M.; Fatichi, S.; Frasson, R. P. M.; Schafer, K. V.
2016-12-01
The Finite-difference Ecosystem-scale Tree-Crown Hydrodynamics model version 2 (FETCH2) is a novel tree-scale hydrodynamic model of transpiration. The FETCH2 model employs a finite difference numerical methodology and a simplified single-beam conduit system and simulates water flow through the tree as a continuum of porous media conduits. It explicitly resolves xylem water potential throughout the tree's vertical extent. Empirical equations relate water potential within the stem to stomatal conductance of the leaves at each height throughout the crown. While highly simplified, this approach brings additional realism to the simulation of transpiration by linking stomatal responses to stem water potential rather than directly to soil moisture, as is currently the case in the majority of land-surface models. FETCH2 accounts for plant hydraulic traits, such as the degree of anisohydric/isohydric response of stomata, maximal xylem conductivity, vertical distribution of leaf area, and maximal and minimal stemwater content. We used FETCH2 along with sap flow and eddy covariance data sets collected from a mixed plot of two genera (oak/pine) in Silas Little Experimental Forest, NJ, USA, to conduct an analysis of the inter-genera variation of hydraulic strategies and their effects on diurnal and seasonal transpiration dynamics. We define these strategies through the parameters that describe the genus-level transpiration and xylem conductivity responses to changes in stem water potential. A virtual experiment showed that the model was able to capture the effect of hydraulic strategies such as isohydric/anisohydric behavior on stomatal conductance under different soil-water availability conditions. Our evaluation revealed that FETCH2 considerably improved the simulation of ecosystem transpiration and latent heat flux than more conventional models.
Xylem- and phloem-based transport of CuO nanoparticles in maize (Zea mays L.).
Wang, Zhenyu; Xie, Xiaoyan; Zhao, Jian; Liu, Xiaoyun; Feng, Wenqiang; White, Jason C; Xing, Baoshan
2012-04-17
This work reports on the toxicity of CuO nanoparticles (NPs) to maize (Zea mays L.) and their transport and redistribution in the plant. CuO NPs (100 mg L(-1)) had no effect on germination, but inhibited the growth of maize seedlings; in comparison the dissolved Cu(2+) ions and CuO bulk particles had no obvious effect on maize growth. CuO NPs were present in xylem sap as examined by transmission electron microscopy (TEM) and energy dispersive spectroscopy (EDS), showing that CuO NPs were transported from roots to shoots via xylem. Split-root experiments and high-resolution TEM observation further showed that CuO NPs could translocate from shoots back to roots via phloem. During this translocation, CuO NPs could be reduced from Cu (II) to Cu (I). To our knowledge, this is the first report of root-shoot-root redistribution of CuO NPs within maize. The current study provides direct evidence for the bioaccumulation and biotransformation of CuO NPs (20-40 nm) in maize, which has significant implications on the potential risk of NPs and food safety.
Sisterson, Mark S; Wallis, Christopher M; Stenger, Drake C
2017-04-01
Glassy-winged sharpshooters must feed as adults to produce mature eggs. Cowpea and sunflower are both readily accepted by the glassy-winged sharpshooter for feeding, but egg production on sunflower was reported to be lower than egg production on cowpea. To better understand the role of adult diet in egg production, effects of xylem-sap chemistry on glassy-winged sharpshooter egg maturation was compared for females confined to cowpea and sunflower. Females confined to cowpea consumed more xylem-sap than females held on sunflower. In response, females held on cowpea produced more eggs, had heavier bodies, and greater lipid content than females held on sunflower. Analysis of cowpea and sunflower xylem-sap found that 17 of 19 amino acids were more concentrated in cowpea xylem-sap than in sunflower xylem-sap. Thus, decreased consumption of sunflower xylem-sap was likely owing to perceived lower quality, with decreased egg production owing to a combination of decreased feeding and lower return per unit volume of xylem-sap consumed. Examination of pairwise correlation coefficients among amino acids indicated that concentrations of several amino acids within a plant species were correlated. Principal component analyses identified latent variables describing amino acid composition of xylem-sap. For females held on cowpea, egg maturation was affected by test date, volume of excreta produced, and principal components describing amino acid composition of xylem-sap. Principal component analyses aided in identifying amino acids that were positively or negatively associated with egg production, although determining causality with respect to key nutritional requirements for glassy-winged sharpshooter egg production will require additional testing. Published by Oxford University Press on behalf of Entomological Society of America 2017. This work is written by US Government employees and is in the public domain in the US.
Causes and Effects of Changes in Xylem Functionality in Apple Fruit
DRAŽETA, LAZAR; LANG, ALEXANDER; HALL, ALISTAIR J.; VOLZ, RICHARD K.; JAMESON, PAULA E.
2004-01-01
• Background and Aims The xylem in fruit of a number of species becomes dysfunctional as the fruit develops, resulting in a reduction of xylem inflow to the fruit. Such a reduction may have consequential effects on the mineral balance of the fruit. The aim of this study was to elucidate the dynamics and nature of xylem failure in developing apples (Malus domestica) showing differing susceptibilities to bitter pit, a calcium‐related disorder. • Methods Developmental changes in xylem functionality of the fruit were investigated in ‘Braeburn’ and ‘Granny Smith’ apples by using a dye infusion technique, to stain the vasculature along the path of dye movement. The vascular bundles were clearly visible in transverse section when fruit were sectioned equatorially. The intensity of staining of the vascular bundles in the fruit was recorded at regular intervals throughout the season. Tissue containing dysfunctional bundles was fixed and embedded in wax for subsequent sectioning and examination. • Key Results As the season progressed, an increasing proportion of vascular bundles failed to show any staining, with the most marked change occurring in the primary bundles, and in nearly all bundles with increasing distance from the stalk end of the fruit. Decreased conductance in the primary bundles of ‘Braeburn’ occurred earlier than in ‘Granny Smith’. Microscopy revealed that the xylem in vascular bundles of the fruit suffered substantial damage, indicating that the mode of dysfunction was via the physical disruption of the xylem caused by expansion of the flesh. • Conclusions Results support the view that the relative calcium deficiency of apple fruit is due to a progressive breakdown of xylem conductance caused by growth‐induced damage to the xylem strand in the bundle. The earlier onset of xylem dysfunction in the cultivar more susceptible to bitter pit suggests that the relative growth dynamics of the fruit may control the occurrence of calcium‐related disorders. PMID:14988096
Gruber, A; Pirkebner, D; Oberhuber, W
2013-10-01
Recent studies on non-structural carbohydrate (NSC) reserves in trees focused on xylem NSC reserves, while still little is known about changes in phloem carbohydrate pools, where NSC charging might be significantly different. To gain insight on NSC dynamics in xylem and phloem, we monitored NSC concentrations in stems and roots of Pinus cembra (L.) and Larix decidua (Mill.) growing at the alpine timberline throughout 2011. Species-specific differences affected tree phenology and carbon allocation during the course of the year. After a delayed start in spring, NSC concentrations in L. decidua were significantly higher in all sampled tissues from August until the end of growing season. In both species, NSC concentrations were five to seven times higher in phloem than that in xylem. However, significant correlations between xylem and phloem starch content found for both species indicate a close linkage between long-term carbon reserves in both tissues. In L. decidua also, free sugar concentrations in xylem and phloem were significantly correlated throughout the year, while a lack of correlation between xylem and phloem free sugar pools in P. cembra indicate a decline of phloem soluble carbohydrate pools during periods of high sink demand.
Seasonal dynamics of mobile carbohydrate pools in phloem and xylem of two alpine timberline conifers
GRUBER, A.; PIRKEBNER, D.; OBERHUBER, W.
2016-01-01
Recent studies on non-structural carbohydrate (NSC) reserves in trees focused on xylem NSC reserves, while still little is known about changes in phloem carbohydrate pools, where NSC charging might be significantly different. To gain insight on NSC dynamics in xylem and phloem, we monitored NSC concentrations in stems and roots of Pinus cembra and Larix decidua growing at the alpine timberline throughout 2011. Species-specific differences affected tree phenology and carbon allocation in the course of the year. After a delayed start in spring, NSC concentrations in Larix decidua were significantly higher in all sampled tissues from August until end of growing season. In both species NSC concentrations were five to seven times higher in phloem than in xylem. However, significant correlations between xylem and phloem starch content found for both species indicate a close linkage between long term carbon reserves in both tissues. In Larix decidua also free sugar concentrations in xylem and phloem were significantly correlated throughout the year, while missing correlations between xylem and phloem free sugar pools in Pinus cembra indicate a decline of phloem soluble carbohydrate pools during periods of high sink demand. PMID:24186941
Zhu, Junqi; Dai, Zhanwu; Vivin, Philippe; Gambetta, Gregory A; Henke, Michael; Peccoux, Anthony; Ollat, Nathalie; Delrot, Serge
2017-12-23
Predicting both plant water status and leaf gas exchange under various environmental conditions is essential for anticipating the effects of climate change on plant growth and productivity. This study developed a functional-structural grapevine model which combines a mechanistic understanding of stomatal function and photosynthesis at the leaf level (i.e. extended Farqhuhar-von Caemmerer-Berry model) and the dynamics of water transport from soil to individual leaves (i.e. Tardieu-Davies model). The model included novel features that account for the effects of xylem embolism (fPLC) on leaf hydraulic conductance and residual stomatal conductance (g0), variable root and leaf hydraulic conductance, and the microclimate of individual organs. The model was calibrated with detailed datasets of leaf photosynthesis, leaf water potential, xylem sap abscisic acid (ABA) concentration and hourly whole-plant transpiration observed within a soil drying period, and validated with independent datasets of whole-plant transpiration under both well-watered and water-stressed conditions. The model well captured the effects of radiation, temperature, CO2 and vapour pressure deficit on leaf photosynthesis, transpiration, stomatal conductance and leaf water potential, and correctly reproduced the diurnal pattern and decline of water flux within the soil drying period. In silico analyses revealed that decreases in g0 with increasing fPLC were essential to avoid unrealistic drops in leaf water potential under severe water stress. Additionally, by varying the hydraulic conductance along the pathway (e.g. root and leaves) and changing the sensitivity of stomatal conductance to ABA and leaf water potential, the model can produce different water use behaviours (i.e. iso- and anisohydric). The robust performance of this model allows for modelling climate effects from individual plants to fields, and for modelling plants with complex, non-homogenous canopies. In addition, the model provides a basis for future modelling efforts aimed at describing the physiology and growth of individual organs in relation to water status. © The Author(s) 2017. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.
Burgess, Stephen S O; Pittermann, Jarmila; Dawson, Todd E
2006-02-01
The hydraulic limitation hypothesis of Ryan & Yoder (1997, Bioscience 47, 235-242) suggests that water supply to leaves becomes increasingly difficult with increasing tree height. Within the bounds of this hypothesis, we conjectured that the vertical hydrostatic gradient which gravity generates on the water column in tall trees would cause a progressive increase in xylem 'safety' (increased resistance to embolism and implosion) and a concomitant decrease in xylem 'efficiency' (decreased hydraulic conductivity). We based this idea on the historically recognized concept of a safety-efficiency trade-off in xylem function, and tested it by measuring xylem conductivity and vulnerability to embolism of Sequoia sempervirens branches collected at a range of heights. Measurements of resistance of branch xylem to embolism did indeed show an increase in 'safety' with height. However, the expected decrease in xylem 'efficiency' was not observed. Instead, sapwood-specific hydraulic conductivities (Ks) of branches increased slightly, while leaf-specific hydraulic conductivities increased dramatically, with height. The latter could be largely explained by strong vertical gradients in specific leaf area. The increase in Ks with height corresponded to a decrease in xylem wall fraction (a measure of wall thickness), an increase in percentage of earlywood and slight increases in conduit diameter. These changes are probably adaptive responses to the increased transport requirements of leaves growing in the upper canopy where evaporative demand is greater. The lack of a safety-efficiency tradeoff may be explained by opposing height trends in the pit aperture and conduit diameter of tracheids and the major and semi-independent roles these play in determining xylem safety and efficiency, respectively.
Changes in the Proteome of Xylem Sap in Brassica oleracea in Response to Fusarium oxysporum Stress
Pu, Zijing; Ino, Yoko; Kimura, Yayoi; Tago, Asumi; Shimizu, Motoki; Natsume, Satoshi; Sano, Yoshitaka; Fujimoto, Ryo; Kaneko, Kentaro; Shea, Daniel J.; Fukai, Eigo; Fuji, Shin-Ichi; Hirano, Hisashi; Okazaki, Keiichi
2016-01-01
Fusarium oxysporum f.sp. conlutinans (Foc) is a serious root-invading and xylem-colonizing fungus that causes yellowing in Brassica oleracea. To comprehensively understand the interaction between F. oxysporum and B. oleracea, composition of the xylem sap proteome of the non-infected and Foc-infected plants was investigated in both resistant and susceptible cultivars using liquid chromatography-tandem mass spectrometry (LC-MS/MS) after in-solution digestion of xylem sap proteins. Whole genome sequencing of Foc was carried out and generated a predicted Foc protein database. The predicted Foc protein database was then combined with the public B. oleracea and B. rapa protein databases downloaded from Uniprot and used for protein identification. About 200 plant proteins were identified in the xylem sap of susceptible and resistant plants. Comparison between the non-infected and Foc-infected samples revealed that Foc infection causes changes to the protein composition in B. oleracea xylem sap where repressed proteins accounted for a greater proportion than those of induced in both the susceptible and resistant reactions. The analysis on the proteins with concentration change > = 2-fold indicated a large portion of up- and down-regulated proteins were those acting on carbohydrates. Proteins with leucine-rich repeats and legume lectin domains were mainly induced in both resistant and susceptible system, so was the case of thaumatins. Twenty-five Foc proteins were identified in the infected xylem sap and 10 of them were cysteine-containing secreted small proteins that are good candidates for virulence and/or avirulence effectors. The findings of differential response of protein contents in the xylem sap between the non-infected and Foc-infected samples as well as the Foc candidate effectors secreted in xylem provide valuable insights into B. oleracea-Foc interactions. PMID:26870056
NASA Technical Reports Server (NTRS)
Hilaire, E.; Young, S. A.; Willard, L. H.; McGee, J. D.; Sweat, T.; Chittoor, J. M.; Guikema, J. A.; Leach, J. E.
2001-01-01
The rice bacterial blight pathogen Xanthomonas oryzae pv. oryzae is a vascular pathogen that elicits a defensive response through interaction with metabolically active rice cells. In leaves of 12-day-old rice seedlings, the exposed pit membrane separating the xylem lumen from the associated parenchyma cells allows contact with bacterial cells. During resistant responses, the xylem secondary walls thicken within 48 h and the pit diameter decreases, effectively reducing the area of pit membrane exposed for access by bacteria. In susceptible interactions and mock-inoculated controls, the xylem walls do not thicken within 48 h. Xylem secondary wall thickening is developmental and, in untreated 65-day-old rice plants, the size of the pit also is reduced. Activity and accumulation of a secreted cationic peroxidase, PO-C1, were previously shown to increase in xylem vessel walls and lumen. Peptide-specific antibodies and immunogold-labeling were used to demonstrate that PO-C1 is produced in the xylem parenchyma and secreted to the xylem lumen and walls. The timing of the accumulation is consistent with vessel secondary wall thickening. The PO-C1 gene is distinct but shares a high level of similarity with previously cloned pathogen-induced peroxidases in rice. PO-C1 gene expression was induced as early as 12 h during resistant interactions and peaked between 18 and 24 h after inoculation. Expression during susceptible interactions was lower than that observed in resistant interactions and was undetectable after infiltration with water, after mechanical wounding, or in mature leaves. These data are consistent with a role for vessel secondary wall thickening and peroxidase PO-C1 accumulation in the defense response in rice to X. oryzae pv. oryzae.
Changes in the Proteome of Xylem Sap in Brassica oleracea in Response to Fusarium oxysporum Stress.
Pu, Zijing; Ino, Yoko; Kimura, Yayoi; Tago, Asumi; Shimizu, Motoki; Natsume, Satoshi; Sano, Yoshitaka; Fujimoto, Ryo; Kaneko, Kentaro; Shea, Daniel J; Fukai, Eigo; Fuji, Shin-Ichi; Hirano, Hisashi; Okazaki, Keiichi
2016-01-01
Fusarium oxysporum f.sp. conlutinans (Foc) is a serious root-invading and xylem-colonizing fungus that causes yellowing in Brassica oleracea. To comprehensively understand the interaction between F. oxysporum and B. oleracea, composition of the xylem sap proteome of the non-infected and Foc-infected plants was investigated in both resistant and susceptible cultivars using liquid chromatography-tandem mass spectrometry (LC-MS/MS) after in-solution digestion of xylem sap proteins. Whole genome sequencing of Foc was carried out and generated a predicted Foc protein database. The predicted Foc protein database was then combined with the public B. oleracea and B. rapa protein databases downloaded from Uniprot and used for protein identification. About 200 plant proteins were identified in the xylem sap of susceptible and resistant plants. Comparison between the non-infected and Foc-infected samples revealed that Foc infection causes changes to the protein composition in B. oleracea xylem sap where repressed proteins accounted for a greater proportion than those of induced in both the susceptible and resistant reactions. The analysis on the proteins with concentration change > = 2-fold indicated a large portion of up- and down-regulated proteins were those acting on carbohydrates. Proteins with leucine-rich repeats and legume lectin domains were mainly induced in both resistant and susceptible system, so was the case of thaumatins. Twenty-five Foc proteins were identified in the infected xylem sap and 10 of them were cysteine-containing secreted small proteins that are good candidates for virulence and/or avirulence effectors. The findings of differential response of protein contents in the xylem sap between the non-infected and Foc-infected samples as well as the Foc candidate effectors secreted in xylem provide valuable insights into B. oleracea-Foc interactions.
Hatcher, P.G.; Lerch, H. E.; Verheyen, Vincent T.
1989-01-01
Organic geochemical investigations of peatified and coalified xylem from gymnosperms have provided useful information on the organic transformational processes collectively known as coalification. The combined use of solid-state 13C nuclear magnetic resonance (NMR) and pyrolysis/gas chromatography/mass spectrometry (py/gc/ms) has allowed us to examine the organic composition of peatified and coalified xylem on both a bulk (average) compositional basis and on a detailed molecular basis. We conclude from our studies that coalification of gymnospermous xylem involves the following processes: 1. (1) early selective removal of cellulosic materials so that lignin, a primary constituent of xylem, is transformed to macromolecular aromatic components in coal; 2. (2) modification of gymnospermous lignin by demethylation to form catechol-like structures, and by condensation reactions to induce a high level of cross-linking at an early stage of coalification; and 3. (3) dehydroxylation during increasing coalification to subbituminous coal, the resultant xylem becomes more phenolic in character as the catechol-like structures decrease. ?? 1989.
Lauzière, Isabelle; Sheather, Simon; Mitchell, Forrest
2008-08-01
A survey of xylem fluid-feeding insects (Hemiptera) exhibiting potential for transmission of Xylella fastidiosa, the bacterium causing Pierce's disease of grapevine, was conducted from 2004 to 2006 in the Hill Country grape growing region of central Texas. Nineteen insect species were collected from yellow sticky traps. Among these, two leafhoppers and one spittlebug comprised 94.57% of the xylem specialists caught in this region. Homalodisca vitripennis (Germar), Graphocephala versuta (Say), and Clastoptera xanthocephala Germar trap catches varied significantly over time, with greatest counts usually recorded between May or June and August and among localities. A comparison of insect counts from traps placed inside and outside vineyards indicated that G. versuta is always more likely captured on the vegetation adjacent to the vineyard. C. xanthocephala was caught inside the vineyard during the summer. Between October and December, the natural habitat offers more suitable host plants, and insects were absent from the vineyards after the first freezes. H. vitripennis was caught in higher numbers inside the vineyards throughout the grape vegetative season. However, insects were also caught in the habitat near the affected crop throughout the year, and residual populations overwintering near vineyards were also recorded. This study shed new light on the fauna of xylem fluid-feeding insects of Texas. These results also provide critical information to vineyard managers for timely applications of insecticides before insect feeding and vectoring to susceptible grapevines.
Inside out: efflux of carbon dioxide from leaves represents more than leaf metabolism.
Stutz, Samantha S; Anderson, Jeremiah; Zulick, Rachael; Hanson, David T
2017-05-17
High concentrations of inorganic carbon in the xylem, produced from root, stem, and branch respiration, travel via the transpiration stream and eventually exit the plant through distant tissues as CO2. Unlike previous studies that focused on the efflux of CO2 from roots and woody tissues, we focus on efflux from leaves and the potential effect on leaf respiration measurements. We labeled transported inorganic carbon, spanning reported xylem concentrations, with 13C and then manipulated transpiration rates in the dark in order to vary the rates of inorganic carbon supply to cut leaves from Brassica napus and Populus deltoides. We used tunable diode laser absorbance spectroscopy to directly measure the rate of gross 13CO2 efflux, derived from inorganic carbon supplied from outside of the leaf, relative to gross 12CO2 efflux generated from leaf cells. These experiemnts showed that 13CO2 efflux was dependent upon the rate of inorganic carbon supply to the leaf and the rate of transpiration. Our data show that the gross leaf efflux of xylem-transported CO2 is likely small in the dark when rates of transpiration are low. However, gross leaf efflux of xylem-transported CO2 could approach half the rate of leaf respiration in the light when transpiration rates and branch inorganic carbon concentrations are high, irrespective of the grossly different petiole morphologies in our experiment. © The Author 2017. Published by Oxford University Press on behalf of the Society for Experimental Biology.
Secchi, Francesca; Zwieniecki, Maciej A.
2012-01-01
It is assumed that the refilling of drought-induced embolism requires the creation of an osmotic gradient between xylem parenchyma cells and vessel lumens to generate the water efflux needed to fill the void. To assess the mechanism of embolism repair, it is crucial to determine if plants can up-regulate the efflux of osmotically active substances into embolized vessels and identify the major components of the released osmoticum. Here, we introduce a new approach of sap collection designed to separate water from nonembolized (functional) and embolized (nonfunctional) vessels. This new approach made possible the chemical analysis of liquid collected from both types of vessels in plants subjected to different levels of water stress. The technique also allowed us to determine the water volumes in nonfunctional vessels as a function of stress level. Overall, with the increase of water stress in plants, the osmotic potential of liquid collected from nonfunctional vessels increased while its volume decreased. These results revealed the presence of both sugars and ions in nonfunctional vessels at elevated levels in comparison with liquid collected from functional vessels, in which only traces of sugars were found. The increased sugar concentration was accompanied by decreased xylem sap pH. These results provide new insight into the biology of refilling, underlining the role of sugar and sugar transporters, and imply that a large degree of hydraulic compartmentalization must exist in the xylem during the refilling process. PMID:22837359
Umebayashi, Toshihiro; Morita, Toshimitsu; Utsumi, Yasuhiro; Kusumoto, Dai; Yasuda, Yuko; Haishi, Tomoyuki; Fukuda, Kenji
2016-10-01
Although previous studies have suggested that branch dieback and whole-plant death due to drought stress occur at 50-88% loss of stem hydraulic conductivity (P 50 and P 88 , respectively), the dynamics of catastrophic failure in the water-conducting pathways in whole plants subjected to drought remain poorly understood. We examined the dynamics of drought stress tolerance in 3-year-old Japanese black pine (Pinus thunbergii Parl.). We nondestructively monitored (i) the spatial distribution of drought-induced embolisms in the stem at greater than P 50 and (ii) recovery from embolisms following rehydration. Stem water distributions were visualized by cryo-scanning electron microscopy. The percentages of both embolized area and loss of hydraulic conductivity showed similar patterns of increase, although the water loss in xylem increased markedly at -5.0 MPa or less. One seedling that had reached 72% loss of the water-conducting area survived and the xylem water potential recovered to -0.3 MPa. We concluded that Japanese black pines may need to maintain water-filled tracheids within earlywood of the current-year xylem under natural conditions to avoid disconnection of water movement between the stem and the tops of branches. It is necessary to determine the spatial distribution of embolisms around the point of the lethal threshold to gain an improved understanding of plant survival under conditions of drought. © The Author 2016. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.
2012-01-01
Background Cost-efficient generation of second-generation biofuels requires plant biomass that can easily be degraded into sugars and further fermented into fuels. However, lignocellulosic biomass is inherently recalcitrant toward deconstruction technologies due to the abundant lignin and cross-linked hemicelluloses. Furthermore, lignocellulosic biomass has a high content of pentoses, which are more difficult to ferment into fuels than hexoses. Engineered plants with decreased amounts of xylan in their secondary walls have the potential to render plant biomass a more desirable feedstock for biofuel production. Results Xylan is the major non-cellulosic polysaccharide in secondary cell walls, and the xylan deficient irregular xylem (irx) mutants irx7, irx8 and irx9 exhibit severe dwarf growth phenotypes. The main reason for the growth phenotype appears to be xylem vessel collapse and the resulting impaired transport of water and nutrients. We developed a xylan-engineering approach to reintroduce xylan biosynthesis specifically into the xylem vessels in the Arabidopsis irx7, irx8 and irx9 mutant backgrounds by driving the expression of the respective glycosyltransferases with the vessel-specific promoters of the VND6 and VND7 transcription factor genes. The growth phenotype, stem breaking strength, and irx morphology was recovered to varying degrees. Some of the plants even exhibited increased stem strength compared to the wild type. We obtained Arabidopsis plants with up to 23% reduction in xylose levels and 18% reduction in lignin content compared to wild-type plants, while exhibiting wild-type growth patterns and morphology, as well as normal xylem vessels. These plants showed a 42% increase in saccharification yield after hot water pretreatment. The VND7 promoter yielded a more complete complementation of the irx phenotype than the VND6 promoter. Conclusions Spatial and temporal deposition of xylan in the secondary cell wall of Arabidopsis can be manipulated by using the promoter regions of vessel-specific genes to express xylan biosynthetic genes. The expression of xylan specifically in the xylem vessels is sufficient to complement the irx phenotype of xylan deficient mutants, while maintaining low overall amounts of xylan and lignin in the cell wall. This engineering approach has the potential to yield bioenergy crop plants that are more easily deconstructed and fermented into biofuels. PMID:23181474
NASA Astrophysics Data System (ADS)
Pleban, J. R.; Mackay, D. S.; Aston, T.; Ewers, B. E.; Wienig, C.
2013-12-01
Quantifying the drought tolerance of crop species and genotypes is essential in order to predict how water stress may impact agricultural productivity. As climate models predict an increase in both frequency and severity of drought corresponding plant hydraulic and biochemical models are needed to accurately predict crop drought tolerance. Drought can result in cavitation of xylem conduits and related loss of plant hydraulic conductivity. This study tested the hypothesis that a model incorporating a plants vulnerability to cavitation would best assess drought tolerance in Brassica rapa. Four Brassica genotypes were subjected to drought conditions at a field site in Laramie, WY. Concurrent leaf gas exchange, volumetric soil moisture content and xylem pressure measurements were made during the drought period. Three models were used to access genotype specific drought tolerance. All 3 models rely on the Farquhar biochemical/biophysical model of leaf level photosynthesis, which is integrated into the Terrestrial Regional Ecosystem Exchange Simulator (TREES). The models differ in how TREES applies the environmental driving data and plant physiological mechanisms; specifically how water availability at the site of photosynthesis is derived. Model 1 established leaf water availability from a modeled soil moisture content; Model 2 input soil moisture measurements directly to establish leaf water availability; Model 3 incorporated the Sperry soil-plant transport model, which calculates flows and pressure along the soil-plant water transport pathway to establish leaf water availability. This third model incorporated measured xylem pressures thus constraining leaf water availability via genotype specific vulnerability curves. A multi-model intercomparison was made using a Bayesian approach, which assessed the interaction between uncertainty in model results and data. The three models were further evaluated by assessing model accuracy and complexity via deviance information criteria (DIC). Results suggest that model 1 was unable to model soil moisture accurately and thus did not effectively characterize drought tolerance. Models 2 and 3 were both effective at characterizing drought tolerance; model 3 preformed best in genotypes with the highest vulnerability to cavitation. By identifying through both Bayesian and DIC analyses models that best characterize drought tolerance future investigations into the interaction between crop productivity and water use can be informed by hypothesis testing using models prior to experimentation.
CLIMAITE - a three factor climate change ecosystem manipulation experiment.
NASA Astrophysics Data System (ADS)
Mikkelsen, T. N.; Beier, C.; Albert, K.; Ro-Poulsen, H.
2007-12-01
The Danish multi factorial climate change effects on vegetation experiment (Climaite) have now been conducted for two years on semi-natural grassland. The day time [CO2], night time temperature and precipitation (drought) have been altered, according to a regional climate change model for the year 2075, in a full factorial split plot design. The manipulated area for each treatment is 7 m2 and it is replicated 6 times. The CO2 and temperature treatments have been conducted continuously except for periods with snow cover. The CO2 is enhanced to 510 ppm via a FACE system based on concentrated CO2 released upwind under pressure. The control of the [CO2] varies with wind speed and irradiation, but during 50 percent of the fumigation period the target concentration was kept within +/-5 percent. The temperature treatment is conducted via infrared reflective curtains covering the plots during night time, and the warming of plants and soil depends of the day irradiation, night time wind speed and factors related to seasonality. In general, the air temperature is increased during night time with 1-2 C° and negligible during the day. The soil temperature in 5 cm depth is enhanced to 0.3 - 0.6 C° during night and day. The artificial summer droughts lasted about one month and differences in soil water content were developed over time. By the end of the treatment the èv content in the soil was as low as 0.06 m3 m-3 compared to 0.20 m3 m-3 in the control. Numerous physical and biological parameters in the grassland ecosystem have been measured and several are responding to the changed environment. After 9 months of exposure enhanced [CO2] stimulated the net photosynthesis (based on dry weight) in both of the domination plant species Calluna Vulgaris and Deschampsia flexuosa. When the plants were exposed to short term saturated [CO2] during gasexchange measurements the long term CO2 treated plants also had the highest photosynthesis rate, meaning that the plants were not physiological down regulated. Calluna Vulgaris exposed to both enhanced [CO2] and temperature in combination with drought showed a lowered photosynthesis during short term saturated [CO2] gasexchange measurements. After one and a half year of manipulation the xylem water potential in the plants were measured during drought treatment. Drought treatment generally decreased the xylem water potential, and the enhanced [CO2] diminished the drought effect for both of the species except for Calluna exposed to all treatments in combination, here the xylem water potential dropped to the lowest level measured.
Guo, Huijuan; Sun, Yucheng; Peng, Xinhong; Wang, Qinyang; Harris, Marvin; Ge, Feng
2016-01-01
The activation of the abscisic acid (ABA) signaling pathway reduces water loss from plants challenged by drought stress. The effect of drought-induced ABA signaling on the defense and nutrition allocation of plants is largely unknown. We postulated that these changes can affect herbivorous insects. We studied the effects of drought on different feeding stages of pea aphids in the wild-type A17 of Medicago truncatula and ABA signaling pathway mutant sta-1. We examined the impact of drought on plant water status, induced plant defense signaling via the abscisic acid (ABA), jasmonic acid (JA), and salicylic acid (SA) pathways, and on the host nutritional quality in terms of leaf free amino acid content. During the penetration phase of aphid feeding, drought decreased epidermis/mesophyll resistance but increased mesophyll/phloem resistance of A17 but not sta-1 plants. Quantification of transcripts associated with ABA, JA and SA signaling indicated that the drought-induced up-regulation of ABA signaling decreased the SA-dependent defense but increased the JA-dependent defense in A17 plants. During the phloem-feeding phase, drought had little effect on the amino acid concentrations and the associated aphid phloem-feeding parameters in both plant genotypes. In the xylem absorption stage, drought decreased xylem absorption time of aphids in both genotypes because of decreased water potential. Nevertheless, the activation of the ABA signaling pathway increased water-use efficiency of A17 plants by decreasing the stomatal aperture and transpiration rate. In contrast, the water potential of sta-1 plants (unable to close stomata) was too low to support xylem absorption activity of aphids; the aphids on sta-1 plants had the highest hemolymph osmolarity and lowest abundance under drought conditions. Taken together this study illustrates the significance of cross-talk between biotic-abiotic signaling pathways in plant-aphid interaction, and reveals the mechanisms leading to alter aphid fecundity in water stresses plants. PMID:26546578
Salt stress induces the formation of a novel type of 'pressure wood' in two Populus species.
Janz, Dennis; Lautner, Silke; Wildhagen, Henning; Behnke, Katja; Schnitzler, Jörg-Peter; Rennenberg, Heinz; Fromm, Jörg; Polle, Andrea
2012-04-01
• Salinity causes osmotic stress and limits biomass production of plants. The goal of this study was to investigate mechanisms underlying hydraulic adaptation to salinity. • Anatomical, ecophysiological and transcriptional responses to salinity were investigated in the xylem of a salt-sensitive (Populus × canescens) and a salt-tolerant species (Populus euphratica). • Moderate salt stress, which suppressed but did not abolish photosynthesis and radial growth in P. × canescens, resulted in hydraulic adaptation by increased vessel frequencies and decreased vessel lumina. Transcript abundances of a suite of genes (FLA, COB-like, BAM, XET, etc.) previously shown to be activated during tension wood formation, were collectively suppressed in developing xylem, whereas those for stress and defense-related genes increased. A subset of cell wall-related genes was also suppressed in salt-exposed P. euphratica, although this species largely excluded sodium and showed no anatomical alterations. Salt exposure influenced cell wall composition involving increases in the lignin : carbohydrate ratio in both species. • In conclusion, hydraulic stress adaptation involves cell wall modifications reciprocal to tension wood formation that result in the formation of a novel type of reaction wood in upright stems named 'pressure wood'. Our data suggest that transcriptional co-regulation of a core set of genes determines reaction wood composition. © 2011 The Authors. New Phytologist © 2011 New Phytologist Trust.
Herbaceous Angiosperms Are Not More Vulnerable to Drought-Induced Embolism Than Angiosperm Trees.
Lens, Frederic; Picon-Cochard, Catherine; Delmas, Chloé E L; Signarbieux, Constant; Buttler, Alexandre; Cochard, Hervé; Jansen, Steven; Chauvin, Thibaud; Doria, Larissa Chacon; Del Arco, Marcelino; Delzon, Sylvain
2016-10-01
The water transport pipeline in herbs is assumed to be more vulnerable to drought than in trees due to the formation of frequent embolisms (gas bubbles), which could be removed by the occurrence of root pressure, especially in grasses. Here, we studied hydraulic failure in herbaceous angiosperms by measuring the pressure inducing 50% loss of hydraulic conductance (P 50 ) in stems of 26 species, mainly European grasses (Poaceae). Our measurements show a large range in P 50 from -0.5 to -7.5 MPa, which overlaps with 94% of the woody angiosperm species in a worldwide, published data set and which strongly correlates with an aridity index. Moreover, the P 50 values obtained were substantially more negative than the midday water potentials for five grass species monitored throughout the entire growing season, suggesting that embolism formation and repair are not routine and mainly occur under water deficits. These results show that both herbs and trees share the ability to withstand very negative water potentials without considerable embolism formation in their xylem conduits during drought stress. In addition, structure-function trade-offs in grass stems reveal that more resistant species are more lignified, which was confirmed for herbaceous and closely related woody species of the daisy group (Asteraceae). Our findings could imply that herbs with more lignified stems will become more abundant in future grasslands under more frequent and severe droughts, potentially resulting in lower forage digestibility. © 2016 American Society of Plant Biologists. All Rights Reserved.
Matsuzaki, Jun; Masumori, Masaya; Tange, Takeshi
2007-05-01
Active phototropic bending of non-elongating and radially growing portion of stems (woody stems) has not been previously documented, whereas negative gravitropic bending is well known. We found phototropic bending in woody stems and searched for the underlying mechanism. We inclined 1-year-old Quercus crispula Blume seedlings and unilaterally illuminated them from a horizontal direction perpendicular to ('normal' illumination) or parallel to ('parallel' illumination) the inclination azimuth. With normal illumination, active phototropic bending and xylem formation could be evaluated separately from the negative gravitropic response and vertical deflection resulting from the weight of the seedlings. One-year-old stems with normal illumination bent significantly, with asymmetrical xylem formation towards the illuminated upper surface and side of the stem, whereas those with parallel illumination showed non-significant lateral bending, with asymmetrical xylem formation only on the upper side. A mechanical model was built on the assumption that a bending moment resulted from the asymmetrical xylem formation during phototropic bending of the woody stems. The model fitted the relationship between the observed spatial distributions of the xylem and the observed lateral bending, and thus supported the hypothesis that phototropic bending of woody stems results from asymmetrical xylem formation, as such occurs during gravitropism.
Xylem phenology and wood production: resolving the chicken-or-egg dilemma.
Lupi, Carlo; Morin, Hubert; Deslauriers, Annie; Rossi, Sergio
2010-10-01
Delays in the start of the growing season reduce the period available for growth and the amount of xylem production. However, a higher number of developing tracheids could prolong cell differentiation and, consequently, lengthen the growing season. The relationship between the amount and duration of cell production in the xylem remains an unresolved issue. The aim of this study was to resolve the chicken-or-egg causality dilemma about duration of growth and cell production through simple- and double-cause models. This was achieved by (1) analysing the intra-annual growth dynamics of the xylem in Picea mariana (Mill.) BSP during 2006-2009 in two contrasting sites of the boreal forest of Quebec, Canada, and (2) extracting the dates of onset and ending of xylem formation and the number of radial cells along the tree ring. A higher number of cells was linked to an earlier onset (r=0.74) and later ending (r=0.61) of cell differentiation. The absence of a relationship between the residuals of the onset and ending of xylogenesis (r(p)=-0.06) indicated that cell production influenced the correlation between the two phenophases of the xylem. These results demonstrated that a higher number of cells produced delay the ending of xylem maturation, so extending the duration of wood formation. © 2010 Blackwell Publishing Ltd.
Effects of Age and Size on Xylem Phenology in Two Conifers of Northwestern China.
Zeng, Qiao; Rossi, Sergio; Yang, Bao
2017-01-01
The climatic signals that directly affect the trees can be registered by xylem during its growth. If the timings and duration of xylem formation change, xylogenesis can occur under different environmental conditions and subsequently be subject to different climatic signals. An experimental design was applied in the field to disentangle the effects of age and size on xylem phenology, and it challenges the hypothesis that the timings and dynamics of xylem growth are size-dependent. Intra-annual dynamics of xylem formation were monitored weekly during the growing seasons 2013 and 2014 in Chinese pine ( Pinus tabulaeformis ) and Qilian juniper ( Juniperus przewalskii ) with different sizes and ages in a semi-arid region of northwestern China. Cell differentiation started 3 weeks earlier in 2013 and terminated 1 week later in 2014 in small-young pines than in big-old pines. However, differences in the timings of growth reactivation disappeared when comparing the junipers with different sizes but similar age. Overall, 77 days were required for xylem differentiation to take place, but timings were shorter for older trees, which also exhibited smaller cell production. Results from this study suggest that tree age does play an important role in timings and duration of growth. The effect of age should also be considered to perform reliable responses of trees to climate.
Secchi, Francesca; Pagliarani, Chiara; Zwieniecki, Maciej A
2017-06-01
Xylem parenchyma cells [vessel associated cells (VACs)] constitute a significant fraction of the xylem in woody plants. These cells are often closely connected with xylem vessels or tracheids via simple pores (remnants of plasmodesmata fields). The close contact and biological activity of VACs during times of severe water stress and recovery from stress suggest that they are involved in the maintenance of xylem transport capacity and responsible for the restoration of vessel/tracheid functionality following embolism events. As recovery from embolism requires the transport of water across xylem parenchyma cell membranes, an understanding of stem-specific aquaporin expression patterns, localization and activity is a crucial part of any biological model dealing with embolism recovery processes in woody plants. In this review, we provide a short overview of xylem parenchyma cell biology with a special focus on aquaporins. In particular we address their distributions and activity during the development of drought stress, during the formation of embolism and the subsequent recovery from stress that may result in refilling. Complemented by the current biological model of parenchyma cell function during recovery from stress, this overview highlights recent breakthroughs on the unique ability of long-lived perennial plants to undergo cycles of embolism-recovery related to drought/rewetting or freeze/thaw events. © 2016 John Wiley & Sons Ltd.
Trueba, Santiago; Pouteau, Robin; Lens, Frederic; Feild, Taylor S; Isnard, Sandrine; Olson, Mark E; Delzon, Sylvain
2017-02-01
Increases in drought-induced tree mortality are being observed in tropical rain forests worldwide and are also likely to affect the geographical distribution of tropical vegetation. However, the mechanisms underlying the drought vulnerability and environmental distribution of tropical species have been little studied. We measured vulnerability to xylem embolism (P 50 ) of 13 woody species endemic to New Caledonia and with different xylem conduit morphologies. We examined the relation between P 50 , along with other leaf and xylem functional traits, and a range of habitat variables. Selected species had P 50 values ranging between -4.03 and -2.00 MPa with most species falling in a narrow range of resistance to embolism above -2.7 MPa. Embolism vulnerability was significantly correlated with elevation, mean annual temperature and percentage of species occurrences located in rain forest habitats. Xylem conduit type did not explain variation in P 50 . Commonly used functional traits such as wood density and leaf traits were not related to embolism vulnerability. Xylem embolism vulnerability stands out among other commonly used functional traits as a major driver of species environmental distribution. Drought-induced xylem embolism vulnerability behaves as a physiological trait closely associated with the habitat occupation of rain forest woody species. © 2016 John Wiley & Sons Ltd.
Effects of environmental conditions on onset of xylem growth in Pinus sylvestris under drought.
Swidrak, Irene; Gruber, Andreas; Kofler, Werner; Oberhuber, Walter
2011-05-01
We determined the influence of environmental factors (air and soil temperature, precipitation, photoperiod) on onset of xylem growth in Scots pine (Pinus sylvestris L.) within a dry inner Alpine valley (750 m a.s.l., Tyrol, Austria) by repeatedly sampling micro-cores throughout 2007-10 at two sites (xeric and dry-mesic) at the start of the growing season. Temperature sums were calculated in degree-days (DD) ≥5 °C from 1 January and 20 March, i.e., spring equinox, to account for photoperiodic control of release from winter dormancy. Threshold temperatures at which xylogenesis had a 0.5 probability of being active were calculated by logistic regression. Onset of xylem growth, which was not significantly different between the xeric and dry-mesic sites, ranged from mid-April in 2007 to early May in 2008. Among most study years, statistically significant differences (P<0.05) in onset of xylem growth were detected. Mean air temperature sums calculated from 1 January until onset of xylem growth were 230 ± 44 DD (mean ± standard deviation) at the xeric site and 205 ± 36 DD at the dry-mesic site. Temperature sums calculated from spring equinox until onset of xylem growth showed somewhat less variability during the 4-year study period, amounting to 144 ± 10 and 137 ± 12 DD at the xeric and dry-mesic sites, respectively. At both sites, xylem growth was active when daily minimum, mean and maximum air temperatures were 5.3, 10.1 and 16.2 °C, respectively. Soil temperature thresholds and DD until onset of xylem growth differed significantly between sites, indicating minor importance of root-zone temperature for onset of xylem growth. Although spring precipitation is known to limit radial growth in P. sylvestris exposed to a dry inner Alpine climate, the results of this study revealed that (i) a daily minimum air temperature threshold for onset of xylem growth in the range 5-6 °C exists and (ii) air temperature sum rather than precipitation or soil temperature triggers start of xylem growth. Based on these findings, we suggest that drought stress forces P. sylvestris to draw upon water reserves in the stem for enlargement of first tracheids after cambial resumption in spring. © The Author 2011. Published by Oxford University Press. All rights reserved.
Effects of environmental conditions on onset of xylem growth in Pinus sylvestris under drought
Swidrak, Irene; Gruber, Andreas; Kofler, Werner; Oberhuber, Walter
2012-01-01
Summary We determined influence of environmental factors (air and soil temperature, precipitation, photoperiod) on onset of xylem growth in Scots pine (Pinus sylvestris L.) within a dry inner Alpine valley (750 m a.s.l., Tyrol, Austria) by repeatedly sampling micro-cores throughout 2007-2010 at two sites (xeric and dry-mesic) at the start of the growing season. Temperature sums were calculated in degree-days (DD) ≥ 5 °C from 1 January and 20 March, i.e. spring equinox, to account for photoperiodic control of release from winter dormancy. Threshold temperatures at which xylogenesis had a 0.5 probability of being active were calculated by logistic regression. Onset of xylem growth, which was not significantly different between the xeric and dry-mesic site, ranged from mid-April in 2007 to early May in 2008. Among most study years statistically significant differences (P < 0.05) in onset of xylem growth were detected. Mean air temperature sums calculated from 1 January until onset of xylem growth were 230 ± 44 DD (mean ± standard deviation) at the xeric and 205 ± 36 DD at the dry-mesic site. Temperature sums calculated from spring equinox until onset of xylem growth showed quite less variability during the four year study period amounting to 144 ± 10 and 137 ± 12 DD at the xeric and dry-mesic site, respectively. At both sites xylem growth was active when daily minimum, mean and maximum air temperatures were 5.3, 10.1 and 16.2 °C, respectively. Soil temperature thresholds and DD until onset of xylem growth differed significantly between sites indicating minor importance of root-zone temperature for onset of xylem growth. Although spring precipitation is known to limit radial growth in P. sylvestris exposed to dry inner Alpine climate, results of this study revealed that (i) a daily minimum air temperature threshold for onset of xylem growth in the range of 5-6 °C exists and (ii) air temperature sum rather than precipitation or soil temperature triggers start of xylem growth. Based on these findings we suggest that drought stress forces P. sylvestris to draw upon water reserves in the stem for enlargement of first tracheids after cambial resumption in spring. PMID:21593011
Bailey, Karen J.
2016-01-01
Measurements of amino acids in the guttation fluid and in the xylem exudates of cut leaves from intact plants provide evidence of the remarkable efficiency with which these nitrogenous compounds are reabsorbed from the xylem sap. This could be achieved by mechanisms involving intercellular transport and/or metabolism. Developmental changes in transcripts and protein showed that transcripts for phosphoenolpyruvate carboxykinase (PEPCK) increased from the base to the leaf tip, and were markedly increased by supplying asparagine. Supplying amino acids also increased the amounts of protein of PEPCK and, to a lesser extent, of pyruvate, Pi dikinase. PEPCK is present in the hydathodes, stomata and vascular parenchyma of rice leaves. Evidence for the role of PEPCK was obtained by using 3-mercaptopicolinic acid (MPA), a specific inhibitor of PEPCK, and by using an activation-tagged rice line that had an increase in PEPCK activity, to show that activation of PEPCK resulted in a decrease in N in the guttation fluid and that treatment by MPA resulted in an increase in amino acids in the guttation fluid and xylem sap towards the leaf tip. Furthermore, increasing PEPCK activity decreased the amount of guttation fluid, whereas decreasing PEPCK activity increased the amount of xylem sap or guttation fluid towards the leaf tip. The findings suggest the following hypotheses: (i) both metabolism and transport are involved in xylem recycling and (ii) excess N is the signal involved in modulating xylem hydraulics, perhaps via nutrient regulation of water-transporting aquaporins. Water relations and vascular metabolism and transport are thus intimately linked. PMID:27053722
In vivo dynamic analysis of water refilling in embolized xylem vessels of intact Zea mays leaves
Ryu, Jeongeun; Hwang, Bae Geun; Lee, Sang Joon
2016-01-01
Background and Aims The refilling of embolized xylem vessels under tension is a major issue in water transport among vascular plants. However, xylem embolism and refilling remain poorly understood because of technical limitations. Direct observation of embolism repair in intact plants is essential to understand the biophysical aspects of water refilling in embolized xylem vessels. This paper reports on details of the water refilling process in leaves of the intact herbaceous monocot plant Zea mays and its refilling kinetics obtained by a direct visualization technique. Methods A synchrotron X-ray micro-imaging technique was used to monitor water refilling in embolized xylem vessels of intact maize leaves. Xylem embolism was artificially induced by using a glass capillary; real-time images of water refilling dynamics were consecutively captured at a frame rate of 50 f.p.s. Key Results Water supply in the radial direction initiates droplet formation on the wall of embolized xylem vessels. Each droplet grows into a water column; this phenomenon shows translation motion or continuous increase in water column volume. In some instances, water columns merge and form one large water column. Water refilling in the radial direction causes rapid recovery from embolism in several minutes. The average water refilling velocity is approx. 1 μm s−1. Conclusions Non-destructive visualization of embolized xylem vessels demonstrates rapid water refilling and gas bubble removal as key elements of embolism repair in a herbaceous monocot species. The refilling kinetics provides new insights into the dynamic mechanism of water refilling phenomena. PMID:27539601
USDA-ARS?s Scientific Manuscript database
Pierce’s disease (PD), caused by the xylem-dwelling pathogen Xylella fastidiosa (X.f.), is a serious threat to grape production. The effects of X.f. infection six months post-inoculation on defense-associated proteins and phenolic compounds found in xylem sap and tissue were evaluated. Defense-assoc...
USDA-ARS?s Scientific Manuscript database
Pierce’s disease of grapevine is a serious threat to grape production and is caused by the xylem-dwelling bacterial pathogen Xylella fastidiosa. Microscopy studies have documented morphological changes to grapevine xylem due to infection by X. fastidiosa. Comparatively, less is known about the bi...
Analysis of HRCT-derived xylem network reveals reverse flow in some vessels
USDA-ARS?s Scientific Manuscript database
Flow in xylem vessels is modeled based on constructions of three dimensional xylem networks derived from High Resolution Computed Tomography (HRCT) images of grapevine (Vitis vinifera) stems. Flow in 6-14% of the vessels was found to be oriented in the opposite direction to the bulk flow under norma...
USDA-ARS?s Scientific Manuscript database
Certain hemipteran insects such as the glassy-winged sharpshooter, Homalodisca vitripennis, subsist entirely on xylem fluid, notwithstanding the poor nutrition of such food. Among many adaptations enabling xylem-feeding are aspects of the insect’s salivation that may also allow these insects to tra...
Will, Rodney E; Wilson, Stuart M; Zou, Chris B; Hennessey, Thomas C
2013-10-01
Tree species growing along the forest-grassland ecotone are near the moisture limit of their range. Small increases in temperature can increase vapor pressure deficit (VPD) which may increase tree water use and potentially hasten mortality during severe drought. We tested a 40% increase in VPD due to an increase in growing temperature from 30 to 33°C (constant dewpoint 21°C) on seedlings of 10 tree species common to the forest-grassland ecotone in the southern Great Plains, USA. Measurement at 33 vs 30°C during reciprocal leaf gas exchange measurements, that is, measurement of all seedlings at both growing temperatures, increased transpiration for seedlings grown at 30°C by 40% and 20% for seedlings grown at 33°C. Higher initial transpiration of seedlings in the 33°C growing temperature treatment resulted in more negative xylem water potentials and fewer days until transpiration decreased after watering was withheld. The seedlings grown at 33°C died 13% (average 2 d) sooner than seedlings grown at 30°C during terminal drought. If temperature and severity of droughts increase in the future, the forest-grassland ecotone could shift because low seedling survival rate may not sufficiently support forest regeneration and migration. © 2013 The Authors. New Phytologist © 2013 New Phytologist Trust.
Nolf, Markus; Creek, Danielle; Duursma, Remko; Holtum, Joseph; Mayr, Stefan; Choat, Brendan
2015-12-01
Coordination of stem and leaf hydraulic traits allows terrestrial plants to maintain safe water status under limited water supply. Tropical rain forests, one of the world's most productive biomes, are vulnerable to drought and potentially threatened by increased aridity due to global climate change. However, the relationship of stem and leaf traits within the plant hydraulic continuum remains understudied, particularly in tropical species. We studied within-plant hydraulic coordination between stems and leaves in three tropical lowland rain forest tree species by analyses of hydraulic vulnerability [hydraulic methods and ultrasonic emission (UE) analysis], pressure-volume relations and in situ pre-dawn and midday water potentials (Ψ). We found finely coordinated stem and leaf hydraulic features, with a strategy of sacrificing leaves in favour of stems. Fifty percent of hydraulic conductivity (P50 ) was lost at -2.1 to -3.1 MPa in stems and at -1.7 to -2.2 MPa in leaves. UE analysis corresponded to hydraulic measurements. Safety margins (leaf P50 - stem P50 ) were very narrow at -0.4 to -1.4 MPa. Pressure-volume analysis and in situ Ψ indicated safe water status in stems but risk of hydraulic failure in leaves. Our study shows that stem and leaf hydraulics were finely tuned to avoid embolism formation in the xylem. © 2015 John Wiley & Sons Ltd.
Transport and coordination in the coupled soil-root-xylem-phloem leaf system
NASA Astrophysics Data System (ADS)
Huang, C. W.; Katul, G. G.; Pockman, W.; Litvak, M. E.; Domec, J. C.; Palmroth, S.
2016-12-01
In response to varying environmental conditions, stomatal pores act as biological valves that dynamically adjust their size thereby determining the rate of CO2 assimilation and water loss (i.e., transpiration) to the dry atmosphere. Although the significance of this biotic control on gas exchange is rarely disputed, representing parsimoniously all the underlying mechanisms responsible for stomatal kinetics remain a subject of some debate. It has been conjectured that stomatal control in seed plants (i.e., angiosperm and gymnosperm) represents a compromise between biochemical demand for CO2 and prevention of excessive water loss. This view has been amended at the whole-plant level, where xylem hydraulics and sucrose transport efficiency in phloem appear to impose additional constraints on gas exchange. If such additional constraints impact stomatal opening and closure, then seed plants may have evolved coordinated photosynthetic-hydraulic-sugar transporting machinery that confers some competitive advantages in fluctuating environmental conditions. Thus, a stomatal optimization model that explicitly considers xylem hydraulics and maximum sucrose transport is developed to explore this coordination in the leaf-xylem-phloem system. The model is then applied to progressive drought conditions. The main findings from the model calculations are that (1) the predicted stomatal conductance from the conventional stomatal optimization theory at the leaf and the newly proposed models converge, suggesting a tight coordination in the leaf-xylem-phloem system; (2) stomatal control is mainly limited by the water supply function of the soil-xylem hydraulic system especially when the water flux through the transpiration stream is significantly larger than water exchange between xylem and phloem; (3) thus, xylem limitation imposed on the supply function can be used to differentiate species with different water use strategy across the spectrum of isohydric to anisohydric behavior. Keywords: leaf-level gas exchange, stomatal control, sucrose transport in phloem, xylem hydraulics
Gravitropism of cucumber hypocotyls: biophysical mechanism of altered growth
NASA Technical Reports Server (NTRS)
Cosgrove, D. J.
1990-01-01
The biophysical basis for the changes in cell elongation rate during gravitropism was examined in aetiolated cucumber (Cucumis sativus L.) hypocotyls. Bulk osmotic pressures on the two sides of the stem and in the epidermal cells were not altered during the early time course of gravitropism. By the pressure-probe technique, a small increase in turgor (0.3 bar, 30 kPa) was detected on the upper (inhibited) side, whereas there was a negligible decrease in turgor on the lower (stimulated) side. These small changes in turgor and water potential appeared to be indirect, passive consequences of the altered growth and the small resistance for water movement from the xylem, and indicated that the change in growth was principally due to changes in wall properties. The results indicate that the hydraulic conductance of the water-transport pathway was large (.25 h-1 bar-1) and the water potential difference supporting cell expansion was no greater than 0.3 bar (30 kPa). From pressure-block experiments, it appeared that upon gravitropic stimulation (1) the yield threshold of the lower half of the stem did not decrease and (2) the wall on the upper side of the stem was not made more rigid by a cross-linking process. Mechanical measurements of the stress/strain properties of the walls showed that the initial development of gravitropism did not involve an alteration of the mechanical behaviour of the isolated walls. Thus, gravitropism in cucumber hypocotyls occurs principally by an alteration of the wall relaxation process, without a necessary change in wall mechanical properties.
Petit, Giai; von Arx, Georg; Kiorapostolou, Natasa; Lechthaler, Silvia; Prendin, Angela Luisa; Anfodillo, Tommaso; Caldeira, Maria C; Cochard, Hervé; Copini, Paul; Crivellaro, Alan; Delzon, Sylvain; Gebauer, Roman; Gričar, Jožica; Grönholm, Leila; Hölttä, Teemu; Jyske, Tuula; Lavrič, Martina; Lintunen, Anna; Lobo-do-Vale, Raquel; Peltoniemi, Mikko; Peters, Richard L; Robert, Elisabeth M R; Roig Juan, Sílvia; Senfeldr, Martin; Steppe, Kathy; Urban, Josef; Van Camp, Janne; Sterck, Frank
2018-06-01
Trees scale leaf (A L ) and xylem (A X ) areas to couple leaf transpiration and carbon gain with xylem water transport. Some species are known to acclimate in A L : A X balance in response to climate conditions, but whether trees of different species acclimate in A L : A X in similar ways over their entire (continental) distributions is unknown. We analyzed the species and climate effects on the scaling of A L vs A X in branches of conifers (Pinus sylvestris, Picea abies) and broadleaved (Betula pendula, Populus tremula) sampled across a continental wide transect in Europe. Along the branch axis, A L and A X change in equal proportion (isometric scaling: b ˜ 1) as for trees. Branches of similar length converged in the scaling of A L vs A X with an exponent of b = 0.58 across European climates irrespective of species. Branches of slow-growing trees from Northern and Southern regions preferentially allocated into new leaf rather than xylem area, with older xylem rings contributing to maintaining total xylem conductivity. In conclusion, trees in contrasting climates adjust their functional balance between water transport and leaf transpiration by maintaining biomass allocation to leaves, and adjusting their growth rate and xylem production to maintain xylem conductance. © 2018 The Authors. New Phytologist © 2018 New Phytologist Trust.
Effects of Age and Size on Xylem Phenology in Two Conifers of Northwestern China
Zeng, Qiao; Rossi, Sergio; Yang, Bao
2018-01-01
The climatic signals that directly affect the trees can be registered by xylem during its growth. If the timings and duration of xylem formation change, xylogenesis can occur under different environmental conditions and subsequently be subject to different climatic signals. An experimental design was applied in the field to disentangle the effects of age and size on xylem phenology, and it challenges the hypothesis that the timings and dynamics of xylem growth are size-dependent. Intra-annual dynamics of xylem formation were monitored weekly during the growing seasons 2013 and 2014 in Chinese pine (Pinus tabulaeformis) and Qilian juniper (Juniperus przewalskii) with different sizes and ages in a semi-arid region of northwestern China. Cell differentiation started 3 weeks earlier in 2013 and terminated 1 week later in 2014 in small-young pines than in big-old pines. However, differences in the timings of growth reactivation disappeared when comparing the junipers with different sizes but similar age. Overall, 77 days were required for xylem differentiation to take place, but timings were shorter for older trees, which also exhibited smaller cell production. Results from this study suggest that tree age does play an important role in timings and duration of growth. The effect of age should also be considered to perform reliable responses of trees to climate. PMID:29379517
Lowe-Power, Tiffany M; Hendrich, Connor G; von Roepenack-Lahaye, Edda; Li, Bin; Wu, Dousheng; Mitra, Raka; Dalsing, Beth L; Ricca, Patrizia; Naidoo, Jacinth; Cook, David; Jancewicz, Amy; Masson, Patrick; Thomma, Bart; Lahaye, Thomas; Michael, Anthony J; Allen, Caitilyn
2018-04-01
Ralstonia solanacearum thrives in plant xylem vessels and causes bacterial wilt disease despite the low nutrient content of xylem sap. We found that R. solanacearum manipulates its host to increase nutrients in tomato xylem sap, enabling it to grow better in sap from infected plants than in sap from healthy plants. Untargeted GC/MS metabolomics identified 22 metabolites enriched in R. solanacearum-infected sap. Eight of these could serve as sole carbon or nitrogen sources for R. solanacearum. Putrescine, a polyamine that is not a sole carbon or nitrogen source for R. solanacearum, was enriched 76-fold to 37 µM in R. solanacearum-infected sap. R. solanacearum synthesized putrescine via a SpeC ornithine decarboxylase. A ΔspeC mutant required ≥ 15 µM exogenous putrescine to grow and could not grow alone in xylem even when plants were treated with putrescine. However, co-inoculation with wildtype rescued ΔspeC growth, indicating R. solanacearum produced and exported putrescine to xylem sap. Intriguingly, treating plants with putrescine before inoculation accelerated wilt symptom development and R. solanacearum growth and systemic spread. Xylem putrescine concentration was unchanged in putrescine-treated plants, so the exogenous putrescine likely accelerated disease indirectly by affecting host physiology. These results indicate that putrescine is a pathogen-produced virulence metabolite. © 2017 Society for Applied Microbiology and John Wiley & Sons Ltd.
Martin, Diane; Tholl, Dorothea; Gershenzon, Jonathan; Bohlmann, Jörg
2002-01-01
Norway spruce (Picea abies L. Karst) produces an oleoresin characterized by a diverse array of terpenoids, monoterpenoids, sesquiterpenoids, and diterpene resin acids that can protect conifers against potential herbivores and pathogens. Oleoresin accumulates constitutively in resin ducts in the cortex and phloem (bark) of Norway spruce stems. De novo formation of traumatic resin ducts (TDs) is observed in the developing secondary xylem (wood) after insect attack, fungal elicitation, and mechanical wounding. Here, we characterize the methyl jasmonate-induced formation of TDs in Norway spruce by microscopy, chemical analyses of resin composition, and assays of terpenoid biosynthetic enzymes. The response involves tissue-specific differentiation of TDs, terpenoid accumulation, and induction of enzyme activities of both prenyltransferases and terpene synthases in the developing xylem, a tissue that constitutively lacks axial resin ducts in spruce. The induction of a complex defense response in Norway spruce by methyl jasmonate application provides new avenues to evaluate the role of resin defenses for protection of conifers against destructive pests such as white pine weevils (Pissodes strobi), bark beetles (Coleoptera, Scolytidae), and insect-associated tree pathogens. PMID:12114556
Ultrasonic emissions during ice nucleation and propagation in plant xylem.
Charrier, Guillaume; Pramsohler, Manuel; Charra-Vaskou, Katline; Saudreau, Marc; Améglio, Thierry; Neuner, Gilbert; Mayr, Stefan
2015-08-01
Ultrasonic acoustic emission analysis enables nondestructive monitoring of damage in dehydrating or freezing plant xylem. We studied acoustic emissions (AE) in freezing stems during ice nucleation and propagation, by combining acoustic and infrared thermography techniques and controlling the ice nucleation point. Ultrasonic activity in freezing samples of Picea abies showed two distinct phases: the first on ice nucleation and propagation (up to 50 AE s(-1) ; reversely proportional to the distance to ice nucleation point), and the second (up to 2.5 AE s(-1) ) after dissipation of the exothermal heat. Identical patterns were observed in other conifer and angiosperm species. The complex AE patterns are explained by the low water potential of ice at the ice-liquid interface, which induced numerous and strong signals. Ice propagation velocities were estimated via AE (during the first phase) and infrared thermography. Acoustic activity ceased before the second phase probably because the exothermal heating and the volume expansion of ice caused decreasing tensions. Results indicate cavitation events at the ice front leading to AE. Ultrasonic emission analysis enabled new insights into the complex process of xylem freezing and might be used to monitor ice propagation in natura. © 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.
USDA-ARS?s Scientific Manuscript database
Purified cell-wall constituents or grape xylem sap added to media affected in vitro growth, biofilm formation, cell aggregation and gene expression of Xylella fastidiosa. Media containing xylem sap from Pierce’s disease (PD)-susceptible plants provided better support for bacterial growth and biofil...
Surface tension phenomena in the xylem sap of three diffuse porous temperate tree species
K. K. Christensen-Dalsgaard; M. T. Tyree; P. G. Mussone
2011-01-01
In plant physiology models involving bubble nucleation, expansion or elimination, it is typically assumed that the surface tension of xylem sap is equal to that of pure water, though this has never been tested. In this study we collected xylem sap from branches of the tree species Populus tremuloides, Betula papyrifera and Sorbus...
Unique growth strategy in the Earth's first trees revealed in silicified fossil trunks from China.
Xu, Hong-He; Berry, Christopher M; Stein, William E; Wang, Yi; Tang, Peng; Fu, Qiang
2017-11-07
Cladoxylopsida included the earliest large trees that formed critical components of globally transformative pioneering forest ecosystems in the Mid- and early Late Devonian (ca. 393-372 Ma). Well-known cladoxylopsid fossils include the up to ∼1-m-diameter sandstone casts known as Eospermatopteris from Middle Devonian strata of New York State. Cladoxylopsid trunk structure comprised a more-or-less distinct cylinder of numerous separate cauline xylem strands connected internally with a network of medullary xylem strands and, near the base, externally with downward-growing roots, all embedded within parenchyma. However, the means by which this complex vascular system was able to grow to a large diameter is unknown. We demonstrate-based on exceptional, up to ∼70-cm-diameter silicified fossil trunks with extensive preservation of cellular anatomy from the early Late Devonian (Frasnian, ca. 374 Ma) of Xinjiang, China-that trunk expansion is associated with a cylindrical zone of diffuse secondary growth within ground and cortical parenchyma and with production of a large amount of wood containing both rays and growth increments concentrically around individual xylem strands by normal cambia. The xylem system accommodates expansion by tearing of individual strand interconnections during secondary development. This mode of growth seems indeterminate, capable of producing trees of large size and, despite some unique features, invites comparison with secondary development in some living monocots. Understanding the structure and growth of cladoxylopsids informs analysis of canopy competition within early forests with the potential to drive global processes. Published under the PNAS license.
Knipfer, Thorsten; Barrios-Masias, Felipe H; Cuneo, Italo F; Bouda, Martin; Albuquerque, Caetano P; Brodersen, Craig R; Kluepfel, Daniel A; McElrone, Andrew J
2018-05-30
A germplasm collection containing varied Juglans genotypes holds potential to improve drought resistance of plant materials for commercial production. We used X-ray computed microtomography to evaluate stem xylem embolism susceptibility/repair in relation to vessel anatomical features (size, arrangement, connectivity and pit characteristics) in 2-year-old saplings of three Juglans species. In vivo analysis revealed interspecific variations in embolism susceptibility among Juglans microcarpa, J. hindsii (both native to arid habitats) and J. ailantifolia (native to mesic habitats). Stem xylem of J. microcarpa was more resistant to drought-induced embolism as compared with J. hindsii and J. ailantifolia (differences in embolism susceptibility among older and current year xylem were not detected in any species). Variations in most vessel anatomical traits were negligible among the three species; however, we detected substantial interspecific differences in intervessel pit characteristics. As compared with J. hindsii and J. ailantifolia, low embolism susceptibility in J. microcarpa was associated with smaller pit size in larger diameter vessels, a smaller area of the shared vessel wall occupied by pits, lower pit frequency and no changes in pit characteristics as vessel diameters increased. Changes in amount of embolized vessels following 40 days of re-watering were minor in intact saplings of all three species highlighting that an embolism repair mechanism did not contribute to drought recovery. In conclusion, our data indicate that interspecific variations in drought-induced embolism susceptibility are associated with species-specific pit characteristics, and these traits may provide a future target for breeding efforts aimed at selecting walnut germplasm with improved drought resistance.
Conifer species adapt to low-rainfall climates by following one of two divergent pathways.
Brodribb, Timothy J; McAdam, Scott A M; Jordan, Gregory J; Martins, Samuel C V
2014-10-07
Water stress is one of the primary selective forces in plant evolution. There are characters often cited as adaptations to water stress, but links between the function of these traits and adaptation to drying climates are tenuous. Here we combine distributional, climatic, and physiological evidence from 42 species of conifers to show that the evolution of drought resistance follows two distinct pathways, both involving the coordinated evolution of tissues regulating water supply (xylem) and water loss (stomatal pores) in leaves. Only species with very efficient stomatal closure, and hence low minimum rates of water loss, inhabit dry habitats, but species diverged in their apparent mechanism for maintaining closed stomata during drought. An ancestral mechanism found in Pinaceae and Araucariaceae species relies on high levels of the hormone abscisic acid (ABA) to close stomata during water stress. A second mechanism, found in the majority of Cupressaceae species, uses leaf desiccation rather than high ABA levels to close stomata during sustained water stress. Species in the latter group were characterized by xylem tissues with extreme resistance to embolism but low levels of foliar ABA after 30 d without water. The combination of low levels of ABA under stress with cavitation-resistant xylem enables these species to prolong stomatal opening during drought, potentially extending their photosynthetic activity between rainfall events. Our data demonstrate a surprising simplicity in the way conifers evolved to cope with water shortage, indicating a critical interaction between xylem and stomatal tissues during the process of evolution to dry climates.
Choat, Brendan; Gambetta, Greg A.; Shackel, Kenneth A.; Matthews, Mark A.
2009-01-01
During the latter stages of development in fleshy fruit, water flow through the xylem declines markedly and the requirements of transpiration and further expansion are fulfilled primarily by the phloem. We evaluated the hypothesis that cessation of water transport through the xylem results from disruption or occlusion of pedicel and berry xylem conduits (hydraulic isolation). Xylem hydraulic resistance (Rh) was measured in developing fruit of grape (Vitis vinifera ‘Chardonnay’) 20 to 100 d after anthesis (DAA) and compared with observations of xylem anatomy by light and cryo-scanning electron microscopy and expression of six plasma membrane intrinsic protein (PIP) aquaporin genes (VvPIP1;1, VvPIP1;2, VvPIP1;3, VvPIP2;1, VvPIP2;2, VvPIP2;3). There was a significant increase in whole berry Rh and receptacle Rh in the latter stages of ripening (80–100 DAA), which was associated with deposition of gels or solutes in many receptacle xylem conduits. Peaks in the expression of some aquaporin isoforms corresponded to lower whole berry Rh 60 to 80 DAA, and the increase in Rh beginning at 80 DAA correlated with decreases in the expression of the two most predominantly expressed PIP genes. Although significant, the increase in berry Rh was not great enough, and occurred too late in development, to explain the decline in xylem flow that occurs at 60 to 75 DAA. The evidence suggests that the fruit is not hydraulically isolated from the parent plant by xylem occlusion but, rather, is “hydraulically buffered” by water delivered via the phloem. PMID:19741048
Kitin, Peter; Voelker, Steven L.; Meinzer, Frederick C.; Beeckman, Hans; Strauss, Steven H.; Lachenbruch, Barbara
2010-01-01
Of 14 transgenic poplar genotypes (Populus tremula × Populus alba) with antisense 4-coumarate:coenzyme A ligase that were grown in the field for 2 years, five that had substantial lignin reductions also had greatly reduced xylem-specific conductivity compared with that of control trees and those transgenic events with small reductions in lignin. For the two events with the lowest xylem lignin contents (greater than 40% reduction), we used light microscopy methods and acid fuchsin dye ascent studies to clarify what caused their reduced transport efficiency. A novel protocol involving dye stabilization and cryo-fluorescence microscopy enabled us to visualize the dye at the cellular level and to identify water-conducting pathways in the xylem. Cryo-fixed branch segments were planed in the frozen state on a sliding cryo-microtome and observed with an epifluorescence microscope equipped with a cryo-stage. We could then distinguish clearly between phenolic-occluded vessels, conductive (stain-filled) vessels, and nonconductive (water- or gas-filled) vessels. Low-lignin trees contained areas of nonconductive, brown xylem with patches of collapsed cells and patches of noncollapsed cells filled with phenolics. In contrast, phenolics and nonconductive vessels were rarely observed in normal colored wood of the low-lignin events. The results of cryo-fluorescence light microscopy were supported by observations with a confocal microscope after freeze drying of cryo-planed samples. Moreover, after extraction of the phenolics, confocal microscopy revealed that many of the vessels in the nonconductive xylem were blocked with tyloses. We conclude that reduced transport efficiency of the transgenic low-lignin xylem was largely caused by blockages from tyloses and phenolic deposits within vessels rather than by xylem collapse. PMID:20639405
Bailey, Karen J; Leegood, Richard C
2016-04-01
Measurements of amino acids in the guttation fluid and in the xylem exudates of cut leaves from intact plants provide evidence of the remarkable efficiency with which these nitrogenous compounds are reabsorbed from the xylem sap. This could be achieved by mechanisms involving intercellular transport and/or metabolism. Developmental changes in transcripts and protein showed that transcripts for phosphoenolpyruvate carboxykinase (PEPCK) increased from the base to the leaf tip, and were markedly increased by supplying asparagine. Supplying amino acids also increased the amounts of protein of PEPCK and, to a lesser extent, of pyruvate, Pi dikinase. PEPCK is present in the hydathodes, stomata and vascular parenchyma of rice leaves. Evidence for the role of PEPCK was obtained by using 3-mercaptopicolinic acid (MPA), a specific inhibitor of PEPCK, and by using an activation-tagged rice line that had an increase in PEPCK activity, to show that activation of PEPCK resulted in a decrease in N in the guttation fluid and that treatment by MPA resulted in an increase in amino acids in the guttation fluid and xylem sap towards the leaf tip. Furthermore, increasing PEPCK activity decreased the amount of guttation fluid, whereas decreasing PEPCK activity increased the amount of xylem sap or guttation fluid towards the leaf tip. The findings suggest the following hypotheses: (i) both metabolism and transport are involved in xylem recycling and (ii) excess N is the signal involved in modulating xylem hydraulics, perhaps via nutrient regulation of water-transporting aquaporins. Water relations and vascular metabolism and transport are thus intimately linked. © The Author 2016. Published by Oxford University Press on behalf of the Society for Experimental Biology.
NASA Astrophysics Data System (ADS)
Katul, Gabriel; Huang, Cheng-Wei
2017-04-01
In response to varying environmental conditions, stomatal pores act as biological valves that dynamically adjust their size thereby determining the rate of CO2 assimilation and water loss (i.e., transpiration) to the atmosphere. Although the significance of this biotic control on gas exchange is rarely disputed, representing parsimoniously all the underlying mechanisms responsible for stomatal kinetics remain a subject of some debate. It has been conjectured that stomatal control in seed plants (i.e., angiosperm and gymnosperm) represents a compromise between biochemical demand for CO2 and prevention of excessive water loss. This view has been amended at the whole-plant level, where xylem hydraulics and sucrose transport efficiency in phloem appear to impose additional constraints on gas exchange. If such additional constraints impact stomatal opening and closure, then seed plants may have evolved coordinated photosynthetic-hydraulic-sugar transporting machinery that confers some competitive advantages in fluctuating environmental conditions. Thus, a stomatal optimization model that explicitly considers xylem hydraulics and maximum sucrose transport is developed to explore this coordination in the leaf-xylem-phloem system. The model is then applied to progressive drought conditions. The main findings from the model calculations are that (1) the predicted stomatal conductance from the conventional stomatal optimization theory at the leaf and the newly proposed models converge, suggesting a tight coordination in the leaf-xylem-phloem system; (2) stomatal control is mainly limited by the water supply function of the soil-xylem hydraulic system especially when the water flux through the transpiration stream is significantly larger than water exchange between xylem and phloem; (3) thus, xylem limitation imposed on the supply function can be used to differentiate species with different water use strategy across the spectrum of isohydric to anisohydric behavior.
Dynamic analysis on cavitation and embolization in vascular plants under tension
NASA Astrophysics Data System (ADS)
Ryu, Jeongeun; Hwang, Bae Geun; Kim, Yangmin; Lee, Sang Joon
2014-11-01
Plants can transport sap water from the soil to the tip of their leaves using the tensile forces created by leaf transpiration without any mechanical pumps. However, the high tension adversely induces a thermodynamically metastable state in sap water with negative pressure and gas bubbles are prone to be formed in xylem vessels. Cavitation easily breaks down continuous water columns and grows into embolization, which limits water transport through xylem vessels. Meanwhile, the repair process of embolization is closely related to water management and regulation of sap flow in plants. In this study, the cavitation and embolization phenomena of liquid water in vascular plants and a physical model system are experimentally and theoretically investigated in detail under in vivo and in vitro conditions. This study will not only shed light on the understanding of these multiphase flows under tension but also provide a clue to solve cavitation problems in micro-scale conduits and microfluidic network systems. This work was supported by the National Research Foundation of Korea (NRF) grant funded by the Korea government (MSIP) (No. 2008-0061991).
Pérez-Donoso, Alonso G.; Sun, Qiang; Roper, M. Caroline; Greve, L. Carl; Kirkpatrick, Bruce; Labavitch, John M.
2010-01-01
The pit membrane (PM) is a primary cell wall barrier that separates adjacent xylem water conduits, limiting the spread of xylem-localized pathogens and air embolisms from one conduit to the next. This paper provides a characterization of the size of the pores in the PMs of grapevine (Vitis vinifera). The PM porosity (PMP) of stems infected with the bacterium Xylella fastidiosa was compared with the PMP of healthy stems. Stems were infused with pressurized water and flow rates were determined; gold particles of known size were introduced with the water to assist in determining the size of PM pores. The effect of introducing trans-1,2-diaminocyclohexane-N,N,N′,N′-tetraacetic acid (CDTA), oligogalacturonides, and polygalacturonic acid into stems on water flux via the xylem was also measured. The possibility that cell wall-degrading enzymes could alter the pore sizes, thus facilitating the ability of X. fastidiosa to cross the PMs, was tested. Two cell wall-degrading enzymes likely to be produced by X. fastidiosa (polygalactuoronase and endo-1,4- β -glucanase) were infused into stems, and particle passage tests were performed to check for changes in PMP. Scanning electron microscopy of control and enzyme-infused stem segments revealed that the combination of enzymes opened holes in PMs, probably explaining enzyme impacts on PMP and how a small X. fastidiosa population, introduced into grapevines by insect vectors, can multiply and spread throughout the vine and cause Pierce's disease. PMID:20107028
USDA-ARS?s Scientific Manuscript database
Xylem flow into the fruit decline at the onset of ripening (i.e. veraison) in grapes, and current literature suggests that there is an increase in hydraulic resistance in the pedicel at this time. However, it is unknown how pedicel hydraulic properties change developmentally in relation to xylem an...
Bae, Chungyun; Han, Sang Wook; Song, Yu-Rim; Kim, Bo-Young; Lee, Hyung-Jin; Lee, Je-Min; Yeam, Inhwa; Heu, Sunggi; Oh, Chang-Sik
2015-07-01
Disease resistance against xylem-colonizing pathogenic bacteria in crops. Plant pathogenic bacteria cause destructive diseases in many commercially important crops. Among these bacteria, eight pathogens, Ralstonia solanacearum, Xanthomonas oryzae pv. oryzae, X. campestris pv. campestris, Erwinia amylovora, Pantoea stewartii subsp. stewartii, Clavibacter michiganensis subsp. michiganensis, Pseudomonas syringae pv. actinidiae, and Xylella fastidiosa, infect their host plants through different infection sites and paths and eventually colonize the xylem tissues of their host plants, resulting in wilting symptoms by blocking water flow or necrosis of xylem tissues. Noticeably, only a relatively small number of resistant cultivars in major crops against these vascular bacterial pathogens except X. oryzae pv. oryzae have been found or generated so far, although these pathogens threaten productivity of major crops. In this review, we summarize the lifestyles of major xylem-colonizing bacterial pathogens and then discuss the progress of current research on disease resistance controlled by qualitative disease resistance genes or quantitative trait loci against them. Finally, we propose infection processes of xylem-colonizing bacterial pathogens as one of possible reasons for why so few qualitative disease resistance genes against these pathogens have been developed or identified so far in crops.
Petit, Giai; Savi, Tadeja; Consolini, Martina; Anfodillo, Tommaso; Nardini, Andrea
2016-11-01
Efficient leaf water supply is fundamental for assimilation processes and tree growth. Renovating the architecture of the xylem transport system requires an increasing carbon investment while growing taller, and any deficiency of carbon availability may result in increasing hydraulic constraints to water flow. Therefore, plants need to coordinate carbon assimilation and biomass allocation to guarantee an efficient and safe long-distance transport system. We tested the hypothesis that reduced branch elongation rates together with carbon-saving adjustments of xylem anatomy hydraulically compensate for the reduction in biomass allocation to xylem. We measured leaf biomass, hydraulic and anatomical properties of wood segments along the main axis of branches in 10 slow growing (SG) and 10 fast growing (FG) Fraxinus ornus L. trees. Branches of SG trees had five times slower branch elongation rate (7 vs 35 cm year -1 ), and produced a higher leaf biomass (P < 0.0001) and thinner xylem rings with fewer but larger vessels (P < 0.0001). On the contrary, we found no differences between SG and FG trees in terms of leaf-specific conductivity (P > 0.05) and xylem safety (Ψ 50 ≈ -3.2 MPa). Slower elongation rate coupled with thinner annual rings and larger vessels allows the reduction of carbon costs associated with growth, while maintaining similar leaf-specific conductivity and xylem safety. © The Author 2016. Published by Oxford University Press. All rights reserved. For Permissions, please e-mail: journals.permissions@oup.com.
NASA Astrophysics Data System (ADS)
Gentine, P.; Guerin, M. F.; von Arx, G.; Martin-Benito, D.; Griffin, K. L.; McDowell, N.; Pockman, W.; Andreu-Hayles, L.
2017-12-01
Recent droughts in the Southwest US have resulted in extensive mortality in the pinion pine population (Pinus Edulis). An important factor for resiliency is the ability of a plant to maintain a functional continuum between soil and leaves, allowing water's motion to be sustained or resumed. During droughts, loss of functional tracheids happens through embolism, which can be partially mitigated by increasing the hydraulic safety of the xylem. However, higher hydraulic safety is usually achieved by building narrower tracheids with thicker walls, resulting in a reduction of the hydraulic efficiency of the xylem (conductivity per unit area). Reduced efficiency constrains water transport, limits photosynthesis and might delay recovery after the drought. Supporting existing research on safety-efficiency tradeoff, we test the hypothesis that under dry conditions, isohydric pinions grow xylem that favor efficiency over safety. Using a seven-year experiment with three watering treatments (drought, control, irrigated) in New Mexico, we investigate the effect of drought on the xylem anatomy of pinions' branches. We also compare the treatment effect with interannual variations in xylem structure. We measure anatomical variables - conductivities, cell wall thicknesses, hydraulic diameter, cell reinforcement and density - and preliminarily conclude that treatment has little effect on hydraulic efficiency while hydraulic safety is significantly reduced under dry conditions. Taking advantage of an extremely dry year occurrence during the experiment, we find a sharp increase in vulnerability for xylem tissues built the same year.
The tomato plastidic fructokinase SlFRK3 plays a role in xylem development.
Stein, Ofer; Damari-Weissler, Hila; Secchi, Francesca; Rachmilevitch, Shimon; German, Marcelo A; Yeselson, Yelena; Amir, Rachel; Schaffer, Arthur; Holbrook, N Michele; Aloni, Roni; Zwieniecki, Maciej A; Granot, David
2016-03-01
Plants have two kinds of fructokinases (FRKs) that catalyze the key step of fructose phosphorylation, cytosolic and plastidic. The major cytosolic tomato FRK, SlFRK2, is essential for the development of xylem vessels. In order to study the role of SlFRK3, which encodes the only plastidic FRK, we generated transgenic tomato (Solanum lycopersicon) plants with RNAi suppression of SlFRK3 as well as plants expressing beta-glucoronidase (GUS) under the SlFRK3 promoter. GUS staining indicated SlFRK3 expression in vascular tissues of the leaves and stems, including cambium, differentiating xylem, young xylem fibers and phloem companion cells. Suppression of SlFRK3 reduced the stem xylem area, stem and root water conductance, and whole-plant transpiration, with minor effects on plant development. However, suppression of SlFRK3 accompanied by partial suppression of SlFRK2 induced significant growth-inhibition effects, including the wilting of mature leaves. Grafting experiments revealed that these growth effects are imposed primarily by the leaves, whose petioles had unlignified, thin-walled xylem fibers with collapsed parenchyma cells around the vessels. A cross between the SlFRK2-antisense and SlFRK3-RNAi lines exhibited similar wilting and anatomical effects, confirming that these effects are the result of the combined suppression of SlFRK3 and SlFRK2. These results demonstrate a role of the plastidic SlFRK3 in xylem development and hydraulic conductance. © 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.
S.T. Michaletz; E.A. Johnson; M.T. Tyree
2012-01-01
It is widely assumed that post-fire tree mortality results from necrosis of phloem and vascular cambium in stems, despite strong evidence that reduced xylem conductivity also plays an important role. In this study, experiments with Populus balsamifera were used to demonstrate two mechanisms by which heat reduces the hydraulic conductivity of xylem:...
John R. Donnelly; John B. Shane; Paul G. Schaberg
1990-01-01
Development of Pb pollution histories using tree ring analyses has been troubled by possible mobility of Pb within stem xylem. In a 2-yr study, we exposed red spruce (Picea rubens Sarg.) seedlings to Pb during one growing season, with Pb excluded in either the previous or following growing season. Lead levels within xylem rings and bark were...
Anatomical and hydraulic properties of sorghum roots exposed to water deficit. [Sorghum bicolor
DOE Office of Scientific and Technical Information (OSTI.GOV)
Cruz, R.T.; Jordan, W.R.; Drew, M.C.
1991-05-01
The effects of a severe water stress in the upper 0-0.15 m rooting zone on development of the exodermis, endodermis and xylem and on radial (Lp) and axial (Ls) hydraulic conductances were studied for Sorghum bicolor. Lp and Lx were based on water flow rates obtained by applying a negative hydrostatic pressure to the proximal xylem ends of excised roots placed in aerated nutrient solution. The same roots were stained with fluorescent berberine and acid phloroglucinol to describe the development of the exodermal and endodermal cell walls from formation of the Casparian band (State I), to deposition of suberin lamellaemore » (State II), and lignification (State III). Lp of 1.5 {times} 10{sup {minus}11} m{sup 3}s{sup {minus}1}MPa{sup {minus}1} was 80% lower in stressed roots than in unstressed controls. At 0.01 and 0.07 m from the root apex, stressed roots were in State III while control roots were in States I and II, respectively. SEM-image analysis for stressed roots indicated that in the exodermis a greater proportion of the cross sectional area was occupied by lignified walls than in the endodermis. Cellufluor, an apoplastic tracer, was blocked at the lignified exodermis even at 0.01 m from the apex in stressed roots. Uranin, a symplastic tracer, was taken up only in the apical region in stressed roots but farther from the apex in the controls. Lx of 7.1 {times} 10{sup {minus}11}m{sup 3}s{sup {minus}1}MPa{sup {minus}1} was 90% lower in stressed roots compared with the controls. Cellufluor test and image analysis showed that although the protoxylem and early metaxylem were conductive in both treatments, stress caused more than a 50% reduction in the diameter of the xylem elements. Results suggest that lignification of the exodermis and endodermis to a large extent decreased apoplastic and symplastic flows and hence Lp in stressed roots. The low Lx in stressed roots was due to a decrease in the diameters of the conductive xylem elements.« less
Effect of long-term forest fertilization on Scots pine xylem quality and wood borer performance.
Heijari, Juha; Nerg, Anne-Marja; Kainulainen, Pirjo; Noldt, Uwe; Levula, Teuvo; Raitio, Hannu; Holopainen, Jarmo K
2008-01-01
We tested whether changes in long-term nutrient availability would affect the xylem quality and characteristics of Scots pine trees as a food source for the larvae of the xylophagous wood borer Hylotrupes bajulus L. (Cerambycidae). We looked for an effect of host plant growth and xylem structural traits on H. bajulus larval performance, and looked for delayed effects of long-term forest fertilization on xylem chemical quality. In general, larval performance was dependent on larval developmental stage. However, the growth of larvae also varied with host plant quality (increases in the concentration of nitrogen and carbon-based secondary compounds of xylem were correlated with a decrease in the larval growth rate). The greater annual growth of trees reduced tracheid length and correlated positively with second-instar H. bajulus growth rate. This is consistent with the hypothesis that intrinsic growth patterns of host plants influence the development of the xylophagous wood borer H. bajulus.
Spatial and temporal patterns of xylem sap pH derived from stems and twigs of Populus deltoides L.
Doug Aubrey; Justin Boyles; Laura Krysinsky; Robert Teskey
2011-01-01
Xylem sap pH (pHX) is critical in determining the quantity of inorganic carbon dissolved in xylem solution from gaseous [CO2] measurements. Studies of internal carbon transport have generally assumed that pHX derived from stems and twigs is similar and that pHX remains constant through time; however, no empirical studies have investigated these assumptions. If any of...
Basha, Sheikh M; Mazhar, Hifza; Vasanthaiah, Hemanth K N
2010-03-01
Pierce's disease (PD) is a destructive bacterial disease of grapes caused by Xylella fastidiosa which is xylem-confined. The tolerance level to this disease varies among Vitis species. Our research was aimed at identifying unique xylem sap proteins present in PD-tolerant Vitis species. The results showed wide variation in the xylem sap protein composition, where a set of polypeptides with pI between 4.5 and 4.7 and M(r) of 31 kDa were present in abundant amount in muscadine (Vitis rotundifolia, PD-tolerant), in reduced levels in Florida hybrid bunch (Vitis spp., PD-tolerant) and absent in bunch grapes (Vitis vinifera, PD-susceptible). Liquid chromatography/mass spectrometry/mass spectrometry analysis of these proteins revealed their similarity to beta-1, 3-glucanase, peroxidase, and a subunit of oxygen-evolving enhancer protein 1, which are known to play role in defense and oxygen generation. In addition, the amount of free amino acids and soluble sugars was found to be significantly lower in xylem sap of muscadine genotypes compared to V. vinifera genotypes, indicating that the higher nutritional value of bunch grape sap may be more suitable for Xylella growth. These data suggest that the presence of these unique proteins in xylem sap is vital for PD tolerance in muscadine and Florida hybrid bunch grapes.
Metzner, Ralf; Schneider, Heike Ursula; Breuer, Uwe; Thorpe, Michael Robert; Schurr, Ulrich; Schroeder, Walter Heinz
2010-01-01
Fluxes of mineral nutrients in the xylem are strongly influenced by interactions with the surrounding stem tissues and are probably regulated by them. Toward a mechanistic understanding of these interactions, we applied stable isotope tracers of magnesium, potassium, and calcium continuously to the transpiration stream of cut bean (Phaseolus vulgaris) shoots to study their radial exchange at the cell and tissue level with stem tissues between pith and phloem. For isotope localization, we combined sample preparation with secondary ion mass spectrometry in a completely cryogenic workflow. After 20 min of application, tracers were readily detectable to various degrees in all tissues. The xylem parenchyma near the vessels exchanged freely with the vessels, its nutrient elements reaching a steady state of strong exchange with elements in the vessels within 20 min, mainly via apoplastic pathways. A slow exchange between vessels and cambium and phloem suggested that they are separated from the xylem, parenchyma, and pith, possibly by an apoplastic barrier to diffusion for nutrients (as for carbohydrates). There was little difference in these distributions when tracers were applied directly to intact xylem via a microcapillary, suggesting that xylem tension had little effect on radial exchange of these nutrients and that their movement was mainly diffusive. PMID:19965970
Antonucci, Serena; Rossi, Sergio; Deslauriers, Annie; Lombardi, Fabio; Marchetti, Marco; Tognetti, Roberto
2015-10-01
Phenological synchronisms between apical and lateral meristems could clarify some aspects related to the physiological relationships among the different organs of trees. This study correlated the phenological phases of bud development and xylem differentiation during spring 2010-14 in balsam fir (Abies balsamea Mill.) and black spruce [(Picea mariana Mill. (BSP)] of the Monts-Valin National Park (Quebec, Canada) by testing the hypothesis that bud development occurs after the reactivation of xylem growth. From May to September, we conducted weekly monitoring of xylem differentiation using microcores and bud development with direct observations on terminal branches. Synchronism between the beginning of bud development and xylem differentiation was found in both species with significant correlations between the phases of bud and xylem phenology. Degree-day sum was more appropriate in assessing the date of bud growth resumption, while thermal thresholds were more suitable for cambium phenology. Our results provide new knowledge on the dynamics of spring phenology and novel information on the synchronisms between two meristems in coniferous trees. The study demonstrates the importance of precisely defining the phases of bud development in order to correctly analyse the relationships with xylem phenology. © The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.
Analysis of xylem formation in pine by cDNA sequencing
NASA Technical Reports Server (NTRS)
Allona, I.; Quinn, M.; Shoop, E.; Swope, K.; St Cyr, S.; Carlis, J.; Riedl, J.; Retzel, E.; Campbell, M. M.; Sederoff, R.;
1998-01-01
Secondary xylem (wood) formation is likely to involve some genes expressed rarely or not at all in herbaceous plants. Moreover, environmental and developmental stimuli influence secondary xylem differentiation, producing morphological and chemical changes in wood. To increase our understanding of xylem formation, and to provide material for comparative analysis of gymnosperm and angiosperm sequences, ESTs were obtained from immature xylem of loblolly pine (Pinus taeda L.). A total of 1,097 single-pass sequences were obtained from 5' ends of cDNAs made from gravistimulated tissue from bent trees. Cluster analysis detected 107 groups of similar sequences, ranging in size from 2 to 20 sequences. A total of 361 sequences fell into these groups, whereas 736 sequences were unique. About 55% of the pine EST sequences show similarity to previously described sequences in public databases. About 10% of the recognized genes encode factors involved in cell wall formation. Sequences similar to cell wall proteins, most known lignin biosynthetic enzymes, and several enzymes of carbohydrate metabolism were found. A number of putative regulatory proteins also are represented. Expression patterns of several of these genes were studied in various tissues and organs of pine. Sequencing novel genes expressed during xylem formation will provide a powerful means of identifying mechanisms controlling this important differentiation pathway.
Fichot, Régis; Laurans, Françoise; Monclus, Romain; Moreau, Alain; Pilate, Gilles; Brignolas, Franck
2009-12-01
Six Populus deltoides Bartr. ex Marsh. x P. nigra L. genotypes were selected to investigate whether stem xylem anatomy correlated with gas exchange rates, water-use efficiency (WUE) and growth performance. Clonal copies of the genotypes were grown in a two-plot common garden test under contrasting water regimes, with one plot maintained irrigated and the other one subjected to moderate summer water deficit. The six genotypes displayed a large range of xylem anatomy, mean vessel and fibre diameter varying from about 40 to 60 microm and from 7.5 to 10.5 microm, respectively. Decreased water availability resulted in a reduced cell size and an important rise in vessel density, but the extent of xylem plasticity was both genotype and trait dependent. Vessel diameter and theoretical xylem-specific hydraulic conductivity correlated positively with stomatal conductance, carbon isotope discrimination and growth performance-related traits and negatively with intrinsic WUE, especially under water deficit conditions. Vessel diameter and vessel density measured under water deficit conditions correlated with the relative losses in biomass production in response to water deprivation; this resulted from the fact that a more plastic xylem structure was generally accompanied by a larger loss in biomass production.
Bowling, David R.; Schulze, Emily S.; Hall, Steven J.
2016-10-14
We revisit a classic ecohydrological study that showed streamside riparian trees in a semiarid mountain catchment did not use perennial stream water. The original study suggested that mature individuals of Acer negundo, Acer grandidentatum, and other species were dependent on water from “deeper strata,” possibly groundwater. We used a dual stable isotope approach (δ 18O and δ 2H) to further examine the water sources of these trees. We tested the hypothesis that groundwater was the main tree water source, but found that neither groundwater nor stream water matched the isotope composition of xylem water during two growing seasons. Soil watermore » (0–1 m depth) was closest to and periodically overlapped with xylem water isotope composition, but overall, xylem water was isotopically enriched compared to all measured water sources. The “two water worlds” hypothesis postulates that soil water comprises isotopically distinct mobile and less mobile pools that do not mix, potentially explaining this disparity. We further hypothesized that isotopic effects during snowpack metamorphosis impart a distinct isotope signature to the less mobile soil water that supplies summer transpiration. Depth trends in water isotopes following snowmelt were consistent with the two water worlds hypothesis, but snow metamorphic isotope effects could not explain the highly enriched xylem water. Thus, the dual isotope approach did not unambiguously determine the water source(s) of these riparian trees. Further exploration of physical, geochemical, and biological mechanisms of water isotope fractionation and partitioning is necessary to resolve these data, highlighting critical challenges in the isotopic determination of plant water sources.« less
Krugner, Rodrigo; Backus, Elaine A
2014-02-01
ABSTRACT The glassy-winged sharpshooter, Homalodisca vitripennis (Germar) (Hemiptera: Cicadellidae), is a xylem fluid-ingesting leafhopper that transmits Xylella fastidiosa Wells et al., a plant-infecting bacterium that causes several plant diseases in the Americas. Although the role of plant water stress on the population density and dispersal ofH. vitripennis has been studied, nothing is known about the effects of plant water stress on the transmission of X. fastidiosa by H. vitripennis. A laboratory study was conducted to determine the influence of plant water stress on the sharpshooter stylet probing behaviors associated with the acquisition and inoculation of X. fastidiosa. Electrical penetration graph was used to monitor H. vitripennis feeding behaviors for 20-h periods on citrus [Citrus sinensis (L.) Osbeck] and almond [Prunus dulcis (Miller) D.A. Webb] plants subjected to levels of water stress. Adult H. vitripennis successfully located xylem vessels, then performed behaviors related to the evaluation of the xylem cell and fluid, and finally ingested xylem fluid from citrus and almond plants under the tested fluid tensions ranging from -5.5 to -33.0 bars and -6.0 to -24.5 bars, respectively. In general, long and frequent feeding events associated with the acquisition and inoculation of X. fastidiosa were observed only in fully irrigated plants (i.e., >-10 bars), which suggests that even low levels of plant water stress may reduce the spread of X. fastidiosa. Results provided insights to disease epidemiology and support the hypothesis that application of regulated deficit irrigation has the potential to reduce the incidence of diseases caused by X.fastidiosa by reducing the number of vectors and by decreasing pathogen transmission efficiency.
DOE Office of Scientific and Technical Information (OSTI.GOV)
Bowling, David R.; Schulze, Emily S.; Hall, Steven J.
We revisit a classic ecohydrological study that showed streamside riparian trees in a semiarid mountain catchment did not use perennial stream water. The original study suggested that mature individuals of Acer negundo, Acer grandidentatum, and other species were dependent on water from “deeper strata,” possibly groundwater. We used a dual stable isotope approach (δ 18O and δ 2H) to further examine the water sources of these trees. We tested the hypothesis that groundwater was the main tree water source, but found that neither groundwater nor stream water matched the isotope composition of xylem water during two growing seasons. Soil watermore » (0–1 m depth) was closest to and periodically overlapped with xylem water isotope composition, but overall, xylem water was isotopically enriched compared to all measured water sources. The “two water worlds” hypothesis postulates that soil water comprises isotopically distinct mobile and less mobile pools that do not mix, potentially explaining this disparity. We further hypothesized that isotopic effects during snowpack metamorphosis impart a distinct isotope signature to the less mobile soil water that supplies summer transpiration. Depth trends in water isotopes following snowmelt were consistent with the two water worlds hypothesis, but snow metamorphic isotope effects could not explain the highly enriched xylem water. Thus, the dual isotope approach did not unambiguously determine the water source(s) of these riparian trees. Further exploration of physical, geochemical, and biological mechanisms of water isotope fractionation and partitioning is necessary to resolve these data, highlighting critical challenges in the isotopic determination of plant water sources.« less
Esseili, Malak A.; Zhang, Zhenwen
2012-01-01
Noroviruses are the leading cause of food-borne outbreaks, including those that involve lettuce. The culturable porcine sapovirus (SaV) was used as a norovirus surrogate to study the persistence and the potential transfer of the virus from roots to leaves and from outer to inner leaves of lettuce plants. Treatment of lettuce with SaV was done through the roots of young plants, the soil, or the outer leaves of mature plants. Sampling of roots, xylem sap, and inner and outer leaves followed by RNA extraction and SaV-specific real-time reverse transcription (RT)-PCR was performed at 2 h and on postinoculation days (PID) 2, 5, 7, 14, and/or 28. When SaV was inoculated through the roots, viral RNA persisted on the roots and in the leaves until PID 28. When the virus was inoculated through the soil, viral RNA was detected on the roots and in the xylem sap until PID 14; viral RNA was detected in the leaves only until PID 2. No infectious virus was detected inside the leaves for either treatment. When SaV was inoculated through the outer leaves, viral RNA persisted on the leaves until PID 14; however, the virus did not transfer to inner leaves. Infectious viral particles on leaves were detected only at 2 h postinoculation. The milky sap (latex) of leaves, but not the roots' xylem sap, significantly decreased virus infectivity when tested in vitro. Collectively, our results showed the transfer of SaV from roots to leaves through the xylem system and the capacity of the sap of lettuce leaves to decrease virus infectivity in leaves. PMID:22752176
Keadtidumrongkul, Pornthep; Suttangkakul, Anongpat; Pinmanee, Phitsanu; Pattana, Kanokwan; Kittiwongwattana, Chokchai; Apisitwanich, Somsak; Vuttipongchaikij, Supachai
2017-08-01
The expression of cell-wall-targeted Carbohydrate Binding Modules (CBMs) can alter cell wall properties and modulate growth and development in plants such as tobacco and potato. CBM2a identified in xylanase 10A from Cellulomonas fimi is of particular interest for its ability to bind crystalline cellulose. However, its potential for promoting plant growth has not been explored. In this work, we tested the ability of CBM2a to promote growth when expressed using both CaMV35S and a vascular tissue-specific promoter derived from Arabidopsis expansin4 (AtEXP4) in three plant species: Arabidopsis, Nicotiana tabacum and Eucalyptus camaldulensis. In Arabidopsis, the expression of AtEXP4pro:CBM2a showed trends for growth promoting effects including the increase of root and hypocotyl lengths and the enlargements of the vascular xylem area, fiber cells and vessel cells. However, in N. tabacum, the expression of CBM2a under the control of either CaMV35S or AtEXP4 promoter resulted in subtle changes in the plant growth, and the thickness of secondary xylem and vessel and fiber cell sizes were generally reduced in the transgenic lines with AtEXP4pro:CBM2a. In Eucalyptus, while transgenics expressing CaMV35S:CBM2a showed very subtle changes compared to wild type, those transgenics with AtEXP4pro:CBM2a showed increases in plant height, enlargement of xylem areas and xylem fiber and vessel cells. These data provide comparative effects of expressing CBM2a protein in different plant species, and this finding can be applied for plant biomass improvement.
Guo, Huijuan; Sun, Yucheng; Peng, Xinhong; Wang, Qinyang; Harris, Marvin; Ge, Feng
2016-02-01
The activation of the abscisic acid (ABA) signaling pathway reduces water loss from plants challenged by drought stress. The effect of drought-induced ABA signaling on the defense and nutrition allocation of plants is largely unknown. We postulated that these changes can affect herbivorous insects. We studied the effects of drought on different feeding stages of pea aphids in the wild-type A17 of Medicago truncatula and ABA signaling pathway mutant sta-1. We examined the impact of drought on plant water status, induced plant defense signaling via the abscisic acid (ABA), jasmonic acid (JA), and salicylic acid (SA) pathways, and on the host nutritional quality in terms of leaf free amino acid content. During the penetration phase of aphid feeding, drought decreased epidermis/mesophyll resistance but increased mesophyll/phloem resistance of A17 but not sta-1 plants. Quantification of transcripts associated with ABA, JA and SA signaling indicated that the drought-induced up-regulation of ABA signaling decreased the SA-dependent defense but increased the JA-dependent defense in A17 plants. During the phloem-feeding phase, drought had little effect on the amino acid concentrations and the associated aphid phloem-feeding parameters in both plant genotypes. In the xylem absorption stage, drought decreased xylem absorption time of aphids in both genotypes because of decreased water potential. Nevertheless, the activation of the ABA signaling pathway increased water-use efficiency of A17 plants by decreasing the stomatal aperture and transpiration rate. In contrast, the water potential of sta-1 plants (unable to close stomata) was too low to support xylem absorption activity of aphids; the aphids on sta-1 plants had the highest hemolymph osmolarity and lowest abundance under drought conditions. Taken together this study illustrates the significance of cross-talk between biotic-abiotic signaling pathways in plant-aphid interaction, and reveals the mechanisms leading to alter aphid fecundity in water stresses plants. © The Author 2015. Published by Oxford University Press on behalf of the Society for Experimental Biology.
NASA Astrophysics Data System (ADS)
Poyatos, R.; Aguadé, D.; Gómez, M.; Mencuccini, M.; Martínez-Vilalta, J.
2013-12-01
Drought-induced defoliation has recently been associated with depletion of carbohydrate reserves and increased mortality risk in Scots pine (Pinus sylvestris L.) at its dry limit. Are defoliated pines hydraulically impaired compared to non-defoliated pines? Moreover, how do defoliated pines cope with potentially lethal droughts, as compared to non-defoliated pines in the same population? In order to address these questions, we measured the seasonal dynamics of sap flow and needle water potentials (2010-2012), hydraulic function and non-structural carbohydrates (NSC) (2012) in healthy and defoliated pines in the Prades mountains (NE Spain). The summer drought was mild in 2010, intense in 2012 and extremely long in 2011. Defoliated Scots pines showed higher sap flow per unit leaf area during spring, but they were more sensitive to summer drought (Figure 1). This pattern was associated with a steeper decline in soil-to-leaf hydraulic conductance, which could not be explained by differences in branch vulnerability to embolism across defoliation classes. Accordingly, the native loss of xylem conductivity in branches, measured in 2012, remained similar across defoliation classes and reached >65% at the peak of the drought. However, a steeper vulnerability curve was observed for root xylem of defoliated pines. Xylem diameter variations (2011-2012) will be used to further investigate possible differences in the aboveground/belowground partitioning of hydraulic resistance across defoliation classes. NSC levels varied across tree organs (leaves>branches>roots>trunk) and strongly declined with drought. Defoliated pines displayed reduced NSC levels throughout the study period, despite enhanced water transport capacity and increased gas exchange rates during spring. Overall, the defoliated vs. healthy status seems to be more associated to differences in carbohydrate storage and dynamics than to hydraulic differences per se. However, starch conversion to soluble sugars during drought also suggests that NSC may be actively involved in the maintenance of xylem and phloem transport. These results highlight the close connection between carbon and water relations in declining Scots pines. Seasonal course (2010-2012) of VPD (upper panel), soil moisture (mid panel) and sap flow per unit leaf area of defoliated and non defoliated Scots pines (lower panel).
NASA Astrophysics Data System (ADS)
Meunier, Félicien; Couvreur, Valentin; Draye, Xavier; Zarebanadkouki, Mohsen; Vanderborght, Jan; Javaux, Mathieu
2017-12-01
In 1978, Landsberg and Fowkes presented a solution of the water flow equation inside a root with uniform hydraulic properties. These properties are root radial conductivity and axial conductance, which control, respectively, the radial water flow between the root surface and xylem and the axial flow within the xylem. From the solution for the xylem water potential, functions that describe the radial and axial flow along the root axis were derived. These solutions can also be used to derive root macroscopic parameters that are potential input parameters of hydrological and crop models. In this paper, novel analytical solutions of the water flow equation are developed for roots whose hydraulic properties vary along their axis, which is the case for most plants. We derived solutions for single roots with linear or exponential variations of hydraulic properties with distance to root tip. These solutions were subsequently combined to construct single roots with complex hydraulic property profiles. The analytical solutions allow one to verify numerical solutions and to get a generalization of the hydric behaviour with the main influencing parameters of the solutions. The resulting flow distributions in heterogeneous roots differed from those in uniform roots and simulations led to more regular, less abrupt variations of xylem suction or radial flux along root axes. The model could successfully be applied to maize effective root conductance measurements to derive radial and axial hydraulic properties. We also show that very contrasted root water uptake patterns arise when using either uniform or heterogeneous root hydraulic properties in a soil-root model. The optimal root radius that maximizes water uptake under a carbon cost constraint was also studied. The optimal radius was shown to be highly dependent on the root hydraulic properties and close to observed properties in maize roots. We finally used the obtained functions for evaluating the impact of root maturation versus root growth on water uptake. Very diverse uptake strategies arise from the analysis. These solutions open new avenues to investigate for optimal genotype-environment-management interactions by optimization, for example, of plant-scale macroscopic hydraulic parameters used in ecohydrogolocial models.
Frost drought in conifers at the alpine timberline: xylem dysfunction and adaptations.
Mayr, Stefan; Hacke, Uwe; Schmid, Peter; Schwienbacher, Franziska; Gruber, Andreas
2006-12-01
Drought stress can cause xylem embolism in trees when the water potential (psi) in the xylem falls below specific vulnerability thresholds. At the alpine timberline, frost drought is known to cause excessive winter embolism unless xylem vulnerability or transpiration is sufficiently reduced to avoid critical psi. We compared annual courses of psi and embolism in Picea abies, Pinus cembra, Pinus mugo, Larix decidua, and Juniperus communis growing at the timberline vs. low altitude. In addition, vulnerability properties and related anatomical parameters as well as wood density (D(t)) and wall reinforcement (wall thickness related to conduit diameter) were studied. This allowed an estimate of stress intensities as well as a detection of adaptations that reduce embolism formation. At the alpine timberline, psi was lowest during winter with corresponding embolism rates of up to 100% in three of the conifers studied. Only Pinus cembra and Larix decidua avoided winter embolism due to moderate psi. Minor embolism was observed at low altitude where the water potentials of all species remained within a narrow range throughout the year. Within species, differences in psi50 (psi at 50% loss of conductivity) at high vs. low altitude were less than 1 MPa. In Picea abies and Pinus cembra, psi50 was more negative at the timberline while, in the other conifer species, psi50 was more negative at low altitude. Juniperus communis exhibited the lowest (-6.4 +/- 0.04 MPa; mean +/- SE) and Pinus mugo the highest psi50 (-3.34 +/- 0.03 MPa). In some cases, D(t) and tracheid wall reinforcement were higher than in previously established relationships of these parameters with psi50, possibly because of mechanical demands associated with the specific growing conditions. Conifers growing at the alpine timberline were exposed to higher drought stress intensities than individuals at low altitude. Frost drought during winter caused high embolism rates which were probably amplified by freeze-thaw stress. Although frost drought had a large effect on plant water transport, adaptations in hydraulic safety and related anatomical parameters were observed in only a few of the conifer species studied.
Fernández-de-Uña, Laura; Rossi, Sergio; Aranda, Ismael; Fonti, Patrick; González-González, Borja D.; Cañellas, Isabel; Gea-Izquierdo, Guillermo
2017-01-01
Climatic scenarios for the Mediterranean region forecast increasing frequency and intensity of drought events. Consequently, a reduction in Pinus sylvestris L. distribution range is projected within the region, with this species being outcompeted at lower elevations by more drought-tolerant taxa such as Quercus pyrenaica Willd. The functional response of these species to the projected shifts in water availability will partially determine their performance and, thus, their competitive success under these changing climatic conditions. We studied how the cambial and leaf phenology and xylem anatomy of these two species responded to a 3-year rainfall exclusion experiment set at their elevational boundary in Central Spain. Additionally, P. sylvestris leaf gas exchange, water potential and carbon isotope content response to the treatment were measured. Likewise, we assessed inter-annual variability in the studied functional traits under control and rainfall exclusion conditions. Prolonged exposure to drier conditions did not affect the onset of xylogenesis in either of the studied species, whereas xylem formation ceased 1–3 weeks earlier in P. sylvestris. The rainfall exclusion had, however, no effect on leaf phenology on either species, which suggests that cambial phenology is more sensitive to drought than leaf phenology. P. sylvestris formed fewer, but larger tracheids under dry conditions and reduced the proportion of latewood in the tree ring. On the other hand, Q. pyrenaica did not suffer earlywood hydraulic diameter changes under rainfall exclusion, but experienced a cumulative reduction in latewood width, which could ultimately challenge its hydraulic performance. The phenological and anatomical response of the studied species to drought is consistent with a shift in resource allocation under drought stress from xylem to other sinks. Additionally, the tighter stomatal control and higher intrinsic water use efficiency observed in drought-stressed P. sylvestris may eventually limit carbon uptake in this species. Our results suggest that both species are potentially vulnerable to the forecasted increase in drought stress, although P. sylvestris might experience a higher risk of drought-induced decline at its low elevational limit. PMID:28744292
Fernández-de-Uña, Laura; Rossi, Sergio; Aranda, Ismael; Fonti, Patrick; González-González, Borja D; Cañellas, Isabel; Gea-Izquierdo, Guillermo
2017-01-01
Climatic scenarios for the Mediterranean region forecast increasing frequency and intensity of drought events. Consequently, a reduction in Pinus sylvestris L. distribution range is projected within the region, with this species being outcompeted at lower elevations by more drought-tolerant taxa such as Quercus pyrenaica Willd. The functional response of these species to the projected shifts in water availability will partially determine their performance and, thus, their competitive success under these changing climatic conditions. We studied how the cambial and leaf phenology and xylem anatomy of these two species responded to a 3-year rainfall exclusion experiment set at their elevational boundary in Central Spain. Additionally, P. sylvestris leaf gas exchange, water potential and carbon isotope content response to the treatment were measured. Likewise, we assessed inter-annual variability in the studied functional traits under control and rainfall exclusion conditions. Prolonged exposure to drier conditions did not affect the onset of xylogenesis in either of the studied species, whereas xylem formation ceased 1-3 weeks earlier in P. sylvestris . The rainfall exclusion had, however, no effect on leaf phenology on either species, which suggests that cambial phenology is more sensitive to drought than leaf phenology. P. sylvestris formed fewer, but larger tracheids under dry conditions and reduced the proportion of latewood in the tree ring. On the other hand, Q. pyrenaica did not suffer earlywood hydraulic diameter changes under rainfall exclusion, but experienced a cumulative reduction in latewood width, which could ultimately challenge its hydraulic performance. The phenological and anatomical response of the studied species to drought is consistent with a shift in resource allocation under drought stress from xylem to other sinks. Additionally, the tighter stomatal control and higher intrinsic water use efficiency observed in drought-stressed P. sylvestris may eventually limit carbon uptake in this species. Our results suggest that both species are potentially vulnerable to the forecasted increase in drought stress, although P. sylvestris might experience a higher risk of drought-induced decline at its low elevational limit.
VULNERABILITY TO CAVITATION IN GRAPEVINES HAS BEEN OVERESTIMATED BY THE CENTRIFUGE TECHNIQUE
USDA-ARS?s Scientific Manuscript database
Grapevines are considered among the most vulnerable woody plant species to water stress-induced cavitation with embolism forming at slight tensions. However, we found that native embolism in stems of field grown Vitis vinifera cv. Chardonnay never exceeded 30% despite xylem water potentials ('x) rea...
Soil acidity and manganese in declining and nondeclining sugar maple stands in Pennsylvania.
Kogelmann, Wilhelm J; Sharpe, William E
2006-01-01
For decades, the hardwood forests of northern Pennsylvania have been subjected to chronic atmospheric loading of acidifying agents. On marginal, high-elevation, unglaciated sites, sugar maples (Acer saccharum Marsh.) have experienced severe decline symptoms and mortality. Accelerated soil acidification, base cation leaching, and increased availability of toxic metals have been suggested as predisposing factors contributing to this decline. Manganese, an essential micronutrient, is also a potentially phytotoxic metal that may be a factor associated with poor sugar maple health on soils vulnerable to acidification from anthropogenic sources. We measured Mn levels in four compartments of the soil-tree system (soil, foliage, xylem wood, and sap) on three sugar maple stands in northern Pennsylvania. Two stands were classified as declining and one was in good health. Negative correlations were found between soil pH and Mn levels in the soil, foliage, sap, and xylem wood. Levels of Mn in these pools were consistently higher on declining sites, which correspondingly exhibited lower levels of Ca and Mg. Species differences between red maple (Acer rubrum L.) and sugar maple at the two declining sites suggested different tolerances to excessive Mn. Molar ratios of Mg/Mn and Ca/Mn were different among sites and showed potential as indicators of soil acidification. Significant correlations among soil, sap, foliage, and xylem wood Mn were also noted. These results show clear Mn differences among sites and, when viewed with recent Mn toxicity experiments and other observational studies, suggest that excessive Mn may play a role in the observed decline and mortality of sugar maple.
Chatelet, David S.; Wistrom, Christina M.; Purcell, Alexander H.; Rost, Thomas L.; Matthews, Mark A.
2011-01-01
Background and Aims The bacterium Xylella fastidiosa (Xf), responsible for Pierce's disease (PD) of grapevine, colonizes the xylem conduits of vines, ultimately killing the plant. However, Vitis vinifera grapevine varieties differ in their susceptibility to Xf and numerous other plant species tolerate Xf populations without showing symptoms. The aim of this study was to examine the xylem structure of grapevines with different susceptibilities to Xf infection, as well as the xylem structure of non-grape plant species that support or limit movement of Xf to determine if anatomical differences might explain some of the differences in susceptibility to Xf. Methods Air and paint were introduced into leaves and stems to examine the connectivity between stem and leaves and the length distribution of their vessels. Leaf petiole and stem anatomies were studied to determine the basis for the free or restricted movement of Xf into the plant. Key Results There were no obvious differences in stem or petiole vascular anatomy among the grape varieties examined, nor among the other plant species that would explain differences in resistance to Xf. Among grape varieties, the more tolerant ‘Sylvaner’ had smaller stem vessel diameters and 20 % more parenchyma rays than the other three varieties. Alternative hosts supporting Xf movement had slightly longer open xylem conduits within leaves, and more connection between stem and leaves, when compared with alternative hosts that limit Xf movement. Conclusions Stem–leaf connectivity via open xylem conduits and vessel length is not responsible for differences in PD tolerance among grape varieties, or for limiting bacterial movement in the tolerant plant species. However, it was found that tolerant host plants had narrower vessels and more parenchyma rays, possibly restricting bacterial movement at the level of the vessels. The implications of xylem structure and connectivity for the means and regulation of bacterial movement are discussed. PMID:21546428
Liu, Li-Min; Qi, Hua; Luo, Xin-Lan; Zhang, Xuan
2008-09-01
Some important phenomena and behaviors concerned with the coordination effect between vapor water loss through plant stomata and liquid water supply in SPAC were discussed in this paper. A large amount of research results showed that plants show isohydric behavior when the plant hydraulic and chemical signals cooperate to promote the stomatal regulation of leaf water potential. The feedback response of stomata to the change of environmental humidity could be used to explain the midday depression of stomatal conductance and photosynthesis under drought condition, and also, to interpret the correlation between stomatal conductance and hydraulic conductance. The feed-forward response of stomata to the change of environmental humidity could be used to explain the hysteresis response of stomatal conductance to leaf-atmosphere vapor pressure deficit. The strategy for getting the most of xylem transport requires the rapid stomatal responses to avoid excess cavitation and the corresponding mechanisms for reversal of cavitation in short time.
Savi, Tadeja; Bertuzzi, Stefano; Branca, Salvatore; Tretiach, Mauro; Nardini, Andrea
2015-02-01
Urban trees help towns to cope with climate warming by cooling both air and surfaces. The challenges imposed by the urban environment, with special reference to low water availability due to the presence of extensive pavements, result in high rates of mortality of street trees, that can be increased by climatic extremes. We investigated the water relations and xylem hydraulic safety/efficiency of Quercus ilex trees growing at urban sites with different percentages of surrounding impervious pavements. Seasonal changes of plant water potential and gas exchange, vulnerability to cavitation and embolism level, and morpho-anatomical traits were measured. We found patterns of increasing water stress and vulnerability to drought at increasing percentages of impervious pavement cover, with a consequent reduction in gas exchange rates, decreased safety margins toward embolism development, and increased vulnerability to cavitation, suggesting the occurrence of stress-induced hydraulic deterioration. The amount of impermeable surface and chronic exposure to water stress influence the site-specific risk of drought-induced dieback of urban trees under extreme drought. Besides providing directions for management of green spaces in towns, our data suggest that xylem hydraulics is key to a full understanding of the responses of urban trees to global change. © 2014 The Authors. New Phytologist © 2014 New Phytologist Trust.
Photosynthetic, hydraulic and biomechanical responses of Juglans californica shoots to wildfire.
Utsumi, Yasuhiro; Bobich, Edward G; Ewers, Frank W
2010-10-01
Leaf gas exchange and stem xylem hydraulic and mechanical properties were studied for unburned adults and resprouting burned Juglans californica (southern California black walnut) trees 1 year after a fire to explore possible trade-offs between mechanical and hydraulic properties of plants. The CO(2) uptake rates and stomatal conductance were 2-3 times greater for resprouting trees than for unburned adults. Both predawn and midday water potentials were more negative for unburned adult trees, indicating that the stems were experiencing greater water stress than the stems of resprouting trees. In addition, the xylem specific conductivity was similar in the two growth forms, even though the stems of resprouting trees were less vulnerable to water-stress-induced embolism than similar diameter, but older, stems of adult trees. The reduced vulnerability may have been due to less cavitation fatigue in stems of resprouts. The modulus of elasticity, modulus of rupture and xylem density were all greater for resprouts, indicating that resprouts have greater mechanical strength than do adult trees. The data suggest that there is no trade-off between stem mechanical strength and shoot hydraulic and photosynthetic efficiency in resprouts, which may have implications for the success of this species in the fire-prone plant communities of southern California.
A comprehensive strategy for identifying long-distance mobile peptides in xylem sap.
Okamoto, Satoru; Suzuki, Takamasa; Kawaguchi, Masayoshi; Higashiyama, Tetsuya; Matsubayashi, Yoshikatsu
2015-11-01
There is a growing awareness that secreted pemediate organ-to-organ communication in higher plants. Xylem sap peptidomics is an effective but challenging approach for identifying long-distance mobile peptides. In this study we developed a simple, gel-free purification system that combines o-chlorophenol extraction with HPLC separation. Using this system, we successfully identified seven oligopeptides from soybean xylem sap exudate that had one or more post-transcriptional modifications: glycosylation, sulfation and/or hydroxylation. RNA sequencing and quantitative PCR analyses showed that the peptide-encoding genes are expressed in multiple tissues. We further analyzed the long-distance translocation of four of the seven peptides using gene-encoding peptides with single amino acid substitutions, and identified these four peptides as potential root-to-shoot mobile oligopeptides. Promoter-GUS analysis showed that all four peptide-encoding genes were expressed in the inner tissues of the root endodermis. Moreover, we found that some of these peptide-encoding genes responded to biotic and/or abiotic factors. These results indicate that our purification system provides a comprehensive approach for effectively identifying endogenous small peptides and reinforce the concept that higher plants employ various peptides in root-to-shoot signaling. © 2015 The Authors The Plant Journal © 2015 John Wiley & Sons Ltd.
Heavy browsing affects the hydraulic capacity of Ceanothus rigidus (Rhamnaceae).
Pittermann, Jarmila; Lance, Jonathan; Poster, Lauren; Baer, Alex; Fox, Laurel R
2014-07-01
Defoliation by herbivores can reduce carbon assimilation, change plant water relations, and even shift the biotic structure of plant communities. In this study, we took advantage of a long-term deer exclosure experiment to examine the consequences of persistent deer herbivory on plant water relations and the xylem structure-function relationships in Ceanothus rigidus, a maritime chaparral shrub in coastal California. Browsed plants had thicker stems with many intertwined short distal twigs, and significantly higher sapwood-to-leaf area ratios than their non-browsed counterparts. Leaf area-specific hydraulic conductivity was similar in both browsed and non-browsed plants, but xylem area-specific conductivity was significantly lower in the browsed plants. Vessel diameters were equivalent in both plant groups, but the number of vessels on a transverse area basis was nearly 40% lower in the browsed plants, accounting for their lower transport efficiency. Mid-day in situ water potentials and losses of hydraulic conductivity due to embolism were similar in both groups of plants but stomatal conductance was higher in the browsed shrubs in the early part of the growing season. We discuss our findings in the context of whole-plant ecophysiology, and explore the consequences of herbivory on hormonal signals, wood anatomy, and xylem function.
The effect of xylem age on volume yield and sugar content of sugar maple sap
Carter B. Gibbs; Carter B. Gibbs
1969-01-01
At the Burlington, Vermont, research unit of the Northeastern Forest Experiment Station, a study was begun in 1966 in an effort to identify the portions of the xylem that produce the most sap and the sap with the highest sugar content. The study revealed that the greatest volume of sap comes from xylem that is about 35 years old, and that the sweetest sap comes from...
Compositions and methods for xylem-specific expression in plant cells
DOE Office of Scientific and Technical Information (OSTI.GOV)
Han, Kyung-Hwan; Ko, Jae-Heung
The invention provides promoter sequences that regulate specific expression of operably linked sequences in developing xylem cells and/or in developing xylem tissue. The developing xylem-specific sequences are exemplified by the DX5, DX8, DX11, and DX15 promoters, portions thereof, and homologs thereof. The invention further provides expression vectors, cells, tissues and plants that contain the invention's sequences. The compositions of the invention and methods of using them are useful in, for example, improving the quantity (biomass) and/or the quality (wood density, lignin content, sugar content etc.) of expressed biomass feedstock products that may be used for bioenergy, biorefinary, and generating woodmore » products such as pulp, paper, and solid wood.« less
Torres-Ruiz, José M; Cochard, Hervé; Mayr, Stefan; Beikircher, Barbara; Diaz-Espejo, Antonio; Rodriguez-Dominguez, Celia M; Badel, Eric; Fernández, José Enrique
2014-11-01
Different methods have been devised to analyze vulnerability to cavitation of plants. Although a good agreement between them is usually found, some discrepancies have been reported when measuring samples from long-vesseled species. The aim of this study was to evaluate possible artifacts derived from different methods and sample sizes. Current-year shoot segments of mature olive trees (Olea europaea), a long-vesseled species, were used to generate vulnerability curves (VCs) by bench dehydration, pressure collar and both static- and flow-centrifuge methods. For the latter, two different rotors were used to test possible effects of the rotor design on the curves. Indeed, high-resolution computed tomography (HRCT) images were used to evaluate the functional status of xylem at different water potentials. Measurements of native embolism were used to validate the methods used. The pressure collar and the two centrifugal methods showed greater vulnerability to cavitation than the dehydration method. The shift in vulnerability thresholds in centrifuge methods was more pronounced in shorter samples, supporting the open-vessel artifact hypothesis as a higher proportion of vessels were open in short samples. The two different rotor designs used for the flow-centrifuge method revealed similar vulnerability to cavitation. Only the bench dehydration or HRCT methods produced VCs that agreed with native levels of embolism and water potential values measured in the field. © 2014 Scandinavian Plant Physiology Society.
Genetic and hormonal regulation of cambial development.
Ursache, Robertas; Nieminen, Kaisa; Helariutta, Ykä
2013-01-01
The stems and roots of most dicot plants increase in diameter by radial growth, due to the activity of secondary meristems. Two types of meristems function in secondary plant body formation: the vascular cambium, which gives rise to secondary xylem and phloem, and the cork cambium, which produces a bark layer that replaces the epidermis and protects the plant stem from mechanical damage and pathogens. Cambial development, the initiation and activity of the vascular cambium, leads to an accumulation of wood, the secondary xylem tissue. The thick, cellulose-rich cell walls of wood provide a source of cellulose and have the potential to be used as a raw material for sustainable and renewable energy production. In this review, we will discuss what is known about the mechanisms regulating the cambium and secondary tissue development. Copyright © Physiologia Plantarum 2012.
Sevanto, Sanna; Holbrook, N. Michele; Ball, Marilyn C.
2012-06-06
Bubble formation in the conduits of woody plants sets a challenge for uninterrupted water transportation from the soil up to the canopy. Freezing and thawing of stems has been shown to increase the number of air-filled (embolized) conduits, especially in trees with large conduit diameters. Despite numerous experimental studies, the mechanisms leading to bubble formation during freezing have not been addressed theoretically. We used classical nucleation theory and fluid mechanics to show which mechanisms are most likely to be responsible for bubble formation during freezing and what parameters determine the likelihood of the process. Our results confirm the common assumptionmore » that bubble formation during freezing is most likely due to gas segregation by ice. If xylem conduit walls are not permeable to the salts expelled by ice during the freezing process, osmotic pressures high enough for air seeding could be created. The build-up rate of segregated solutes in front of the ice-water interface depends equally on conduit diameter and freezing velocity. Therefore, bubble formation probability depends on these variables. The dependence of bubble formation probability on freezing velocity means that the experimental results obtained for cavitation threshold conduit diameters during freeze/thaw cycles depend on the experimental setup; namely sample size and cooling rate. The velocity dependence also suggests that to avoid bubble formation during freezing trees should have narrow conduits where freezing is likely to be fast (e.g., branches or outermost layer of the xylem). Avoidance of bubble formation during freezing could thus be one piece of the explanation why xylem conduit size of temperate and boreal zone trees varies quite systematically.« less
Sevanto, Sanna; Holbrook, N Michele; Ball, Marilyn C
2012-01-01
Bubble formation in the conduits of woody plants sets a challenge for uninterrupted water transportation from the soil up to the canopy. Freezing and thawing of stems has been shown to increase the number of air-filled (embolized) conduits, especially in trees with large conduit diameters. Despite numerous experimental studies, the mechanisms leading to bubble formation during freezing have not been addressed theoretically. We used classical nucleation theory and fluid mechanics to show which mechanisms are most likely to be responsible for bubble formation during freezing and what parameters determine the likelihood of the process. Our results confirm the common assumption that bubble formation during freezing is most likely due to gas segregation by ice. If xylem conduit walls are not permeable to the salts expelled by ice during the freezing process, osmotic pressures high enough for air seeding could be created. The build-up rate of segregated solutes in front of the ice-water interface depends equally on conduit diameter and freezing velocity. Therefore, bubble formation probability depends on these variables. The dependence of bubble formation probability on freezing velocity means that the experimental results obtained for cavitation threshold conduit diameters during freeze/thaw cycles depend on the experimental setup; namely sample size and cooling rate. The velocity dependence also suggests that to avoid bubble formation during freezing trees should have narrow conduits where freezing is likely to be fast (e.g., branches or outermost layer of the xylem). Avoidance of bubble formation during freezing could thus be one piece of the explanation why xylem conduit size of temperate and boreal zone trees varies quite systematically.
Easy Come, Easy Go: Capillary Forces Enable Rapid Refilling of Embolized Primary Xylem Vessels.
Rolland, Vivien; Bergstrom, Dana M; Lenné, Thomas; Bryant, Gary; Chen, Hua; Wolfe, Joe; Holbrook, N Michele; Stanton, Daniel E; Ball, Marilyn C
2015-08-01
Protoxylem plays an important role in the hydraulic function of vascular systems of both herbaceous and woody plants, but relatively little is known about the processes underlying the maintenance of protoxylem function in long-lived tissues. In this study, embolism repair was investigated in relation to xylem structure in two cushion plant species, Azorella macquariensis and Colobanthus muscoides, in which vascular water transport depends on protoxylem. Their protoxylem vessels consisted of a primary wall with helical thickenings that effectively formed a pit channel, with the primary wall being the pit channel membrane. Stem protoxylem was organized such that the pit channel membranes connected vessels with paratracheal parenchyma or other protoxylem vessels and were not exposed directly to air spaces. Embolism was experimentally induced in excised vascular tissue and detached shoots by exposing them briefly to air. When water was resupplied, embolized vessels refilled within tens of seconds (excised tissue) to a few minutes (detached shoots) with water sourced from either adjacent parenchyma or water-filled vessels. Refilling occurred in two phases: (1) water refilled xylem pit channels, simplifying bubble shape to a rod with two menisci; and (2) the bubble contracted as the resorption front advanced, dissolving air along the way. Physical properties of the protoxylem vessels (namely pit channel membrane porosity, hydrophilic walls, vessel dimensions, and helical thickenings) promoted rapid refilling of embolized conduits independent of root pressure. These results have implications for the maintenance of vascular function in both herbaceous and woody species, because protoxylem plays a major role in the hydraulic systems of leaves, elongating stems, and roots. © 2015 American Society of Plant Biologists. All Rights Reserved.
Linking Tropical Forest Function to Hydraulic Traits in a Size-Structured and Trait-Based Model
NASA Astrophysics Data System (ADS)
Christoffersen, B. O.; Gloor, M.; Fauset, S.; Fyllas, N.; Galbraith, D.; Baker, T. R.; Rowland, L.; Fisher, R.; Binks, O.; Sevanto, S.; Xu, C.; Jansen, S.; Choat, B.; Mencuccini, M.; McDowell, N. G.; Meir, P.
2015-12-01
A major weakness of forest ecosystem models is their inability to capture the diversity of responses to changes in water availability, severely hampering efforts to predict the fate of tropical forests under climate change. Such models often prescribe moisture sensitivity using heuristic response functions that are uniform across all individuals and lack important knowledge about trade-offs in hydraulic traits. We address this weakness by implementing a process representation of plant hydraulics into an individual- and trait-based model (Trait Forest Simulator; TFS) intended for application at discrete sites where community-level distributions of stem and leaf trait spectra (wood density, leaf mass per area, leaf nitrogen and phosphorus content) are known. The model represents a trade-off in the safety and efficiency of water conduction in xylem tissue through hydraulic traits, while accounting for the counteracting effects of increasing hydraulic path length and xylem conduit taper on whole-plant hydraulic resistance with increasing tree size. Using existing trait databases and additional meta-analyses from the rich literature on tropical tree ecophysiology, we obtained all necessary hydraulic parameters associated with xylem conductivity, vulnerability curves, pressure-volume curves, and hydraulic architecture (e.g., leaf-to-sapwood area ratios) as a function of the aforementioned traits and tree size. Incorporating these relationships in the model greatly improved the diversity of tree response to seasonal changes in water availability as well as in response to drought, as determined by comparison with field observations and experiments. Importantly, this individual- and trait-based framework provides a testbed for identifying both critical processes and functional traits needed for inclusion in coarse-scale Dynamic Global Vegetation Models, which will lead to reduced uncertainty in the future state of tropical forests.
Gourieroux, Aude M; Holzapfel, Bruno P; Scollary, Geoffrey R; McCully, Margaret E; Canny, Martin J; Rogiers, Suzy Y
2016-08-01
Amino acids are essential to grape berry and seed development and they are transferred to the reproductive structures through the phloem and xylem from various locations within the plant. The diurnal and seasonal dynamics of xylem and phloem amino acid composition in the leaf petiole and bunch rachis of field-grown Cabernet Sauvignon are described to better understand the critical periods for amino acid import into the berry. Xylem sap was extracted by the centrifugation of excised leaf petioles and rachises, while phloem exudate was collected by immersing these structures in an ethylenediaminetetraacetic acid (EDTA) buffer. Glutamine and glutamic acid were the predominant amino acids in the xylem sap of both grapevine rachises and petioles, while arginine and glycine were the principal amino acids of the phloem exudate. The amino acid concentrations within the xylem sap and phloem exudate derived from these structures were greatest during anthesis and fruit set, and a second peak occurred within the rachis phloem at the onset of ripening. The concentrations of the amino acids within the phloem and xylem sap of the rachis were highest just prior to or after midnight while the flow of sugar through the rachis phloem was greatest during the early afternoon. Sugar exudation rates from the rachis was greater than that of the petiole phloem between anthesis and berry maturity. In summary, amino acid and sugar delivery through the vasculature to grape berries fluctuates over the course of the day as well as through the season and is not necessarily related to levels near the source. Copyright © 2016 Elsevier Masson SAS. All rights reserved.
Branch xylem density variations across the Amazon Basin
NASA Astrophysics Data System (ADS)
Patiño, S.; Lloyd, J.; Paiva, R.; Baker, T. R.; Quesada, C. A.; Mercado, L. M.; Schmerler, J.; Schwarz, M.; Santos, A. J. B.; Aguilar, A.; Czimczik, C. I.; Gallo, J.; Horna, V.; Hoyos, E. J.; Jimenez, E. M.; Palomino, W.; Peacock, J.; Peña-Cruz, A.; Sarmiento, C.; Sota, A.; Turriago, J. D.; Villanueva, B.; Vitzthum, P.; Alvarez, E.; Arroyo, L.; Baraloto, C.; Bonal, D.; Chave, J.; Costa, A. C. L.; Herrera, R.; Higuchi, N.; Killeen, T.; Leal, E.; Luizão, F.; Meir, P.; Monteagudo, A.; Neil, D.; Núñez-Vargas, P.; Peñuela, M. C.; Pitman, N.; Priante Filho, N.; Prieto, A.; Panfil, S. N.; Rudas, A.; Salomão, R.; Silva, N.; Silveira, M.; Soares Dealmeida, S.; Torres-Lezama, A.; Vásquez-Martínez, R.; Vieira, I.; Malhi, Y.; Phillips, O. L.
2009-04-01
Xylem density is a physical property of wood that varies between individuals, species and environments. It reflects the physiological strategies of trees that lead to growth, survival and reproduction. Measurements of branch xylem density, ρx, were made for 1653 trees representing 598 species, sampled from 87 sites across the Amazon basin. Measured values ranged from 218 kg m-3 for a Cordia sagotii (Boraginaceae) from Mountagne de Tortue, French Guiana to 1130 kg m-3 for an Aiouea sp. (Lauraceae) from Caxiuana, Central Pará, Brazil. Analysis of variance showed significant differences in average ρx across regions and sampled plots as well as significant differences between families, genera and species. A partitioning of the total variance in the dataset showed that species identity (family, genera and species) accounted for 33% with environment (geographic location and plot) accounting for an additional 26%; the remaining "residual" variance accounted for 41% of the total variance. Variations in plot means, were, however, not only accountable by differences in species composition because xylem density of the most widely distributed species in our dataset varied systematically from plot to plot. Thus, as well as having a genetic component, branch xylem density is a plastic trait that, for any given species, varies according to where the tree is growing in a predictable manner. Within the analysed taxa, exceptions to this general rule seem to be pioneer species belonging for example to the Urticaceae whose branch xylem density is more constrained than most species sampled in this study. These patterns of variation of branch xylem density across Amazonia suggest a large functional diversity amongst Amazonian trees which is not well understood.
Lignin Composition and Structure Differs between Xylem, Phloem and Phellem in Quercus suber L.
Lourenço, Ana; Rencoret, Jorge; Chemetova, Catarina; Gominho, Jorge; Gutiérrez, Ana; del Río, José C.; Pereira, Helena
2016-01-01
The composition and structure of lignin in different tissues—phellem (cork), phloem and xylem (wood)—of Quercus suber was studied. Whole cell walls and their respective isolated milled lignins were analyzed by pyrolysis coupled with gas chromatography/mass spectrometry (Py-GC/MS), two-dimensional nuclear magnetic resonance spectroscopy (2D-NMR) and derivatization followed by reductive cleavage (DFRC). Different tissues presented varied p-hydroxyphenyl:guaiacyl:syringyl (H:G:S) lignin compositions. Whereas lignin from cork has a G-rich lignin (H:G:S molar ratio 2:85:13), lignin from phloem presents more S-units (H:G:S molar ratio of 1:58:41) and lignin from xylem is slightly enriched in S-lignin (H:G:S molar ratio 1:45:55). These differences were reflected in the relative abundances of the different interunit linkages. Alkyl-aryl ethers (β–O–4′) were predominant, increasing from 68% in cork, to 71% in phloem and 77% in xylem, as consequence of the enrichment in S-lignin units. Cork lignin was enriched in condensed structures such as phenylcoumarans (β-5′, 20%), dibenzodioxocins (5–5′, 5%), as corresponds to a lignin enriched in G-units. In comparison, lignin from phloem and xylem presented lower levels of condensed linkages. The lignin from cork was highly acetylated at the γ-OH of the side-chain (48% lignin acetylation), predominantly over G-units; while the lignins from phloem and xylem were barely acetylated and this occurred mainly over S-units. These results are a first time overview of the lignin structure in xylem, phloem (generated by cambium), and in cork (generated by phellogen), in agreement with literature that reports that lignin biosynthesis is flexible and cell specific. PMID:27833631
Liu, X-P; Gong, C-M; Fan, Y-Y; Eiblmeier, M; Zhao, Z; Han, G; Rennenberg, H
2013-01-01
This study aimed to identify drought-mediated differences in amino nitrogen (N) composition and content of xylem and phloem in trees having different symbiotic N(2)-fixing bacteria. Under controlled water availability, 1-year-old seedlings of Robinia pseudoacacia (nodules with Rhizobium), Hippophae rhamnoides (symbiosis with Frankia) and Buddleja alternifolia (no such root symbiosis) were exposed to control, medium drought and severe drought, corresponding soil water content of 70-75%, 45-50% and 30-35% of field capacity, respectively. Composition and content of amino compounds in xylem sap and phloem exudates were analysed as a measure of N nutrition. Drought strongly reduced biomass accumulation in all species, but amino N content in xylem and phloem remained unaffected only in R. pseudoacacia. In H. rhamnoides and B. alternifolia, amino N in phloem remained constant, but increased in xylem of both species in response to drought. There were differences in composition of amino compounds in xylem and phloem of the three species in response to drought. Proline concentrations in long-distance transport pathways of all three species were very low, below the limit of detection in phloem of H. rhamnoides and in phloem and xylem of B. alternifolia. Apparently, drought-mediated changes in N composition were much more connected with species-specific changes in C:N ratios. Irrespective of soil water content, the two species with root symbioses did not show similar features for the different types of symbiosis, neither in N composition nor in N content. There was no immediate correlation between symbiotic N fixation and drought-mediated changes in amino N in the transport pathways. © 2012 German Botanical Society and The Royal Botanical Society of the Netherlands.
Reusche, Michael; Thole, Karin; Janz, Dennis; Truskina, Jekaterina; Rindfleisch, Sören; Drübert, Christine; Polle, Andrea; Lipka, Volker; Teichmann, Thomas
2012-01-01
The soilborne fungal plant pathogen Verticillium longisporum invades the roots of its Brassicaceae hosts and proliferates in the plant vascular system. Typical aboveground symptoms of Verticillium infection on Brassica napus and Arabidopsis thaliana are stunted growth, vein clearing, and leaf chloroses. Here, we provide evidence that vein clearing is caused by pathogen-induced transdifferentiation of chloroplast-containing bundle sheath cells to functional xylem elements. In addition, our findings suggest that reinitiation of cambial activity and transdifferentiation of xylem parenchyma cells results in xylem hyperplasia within the vasculature of Arabidopsis leaves, hypocotyls, and roots. The observed de novo xylem formation correlates with Verticillium-induced expression of the VASCULAR-RELATED NAC DOMAIN (VND) transcription factor gene VND7. Transgenic Arabidopsis plants expressing the chimeric repressor VND7-SRDX under control of a Verticillium infection-responsive promoter exhibit reduced de novo xylem formation. Interestingly, infected Arabidopsis wild-type plants show higher drought stress tolerance compared with noninfected plants, whereas this effect is attenuated by suppression of VND7 activity. Together, our results suggest that V. longisporum triggers a tissue-specific developmental plant program that compensates for compromised water transport and enhances the water storage capacity of infected Brassicaceae host plants. In conclusion, we provide evidence that this natural plant–fungus pathosystem has conditionally mutualistic features. PMID:23023171
A broad survey of hydraulic and mechanical safety in the xylem of conifers
Bouche, Pauline S.; Larter, Maximilien; Domec, Jean-Christophe; Burlett, Régis; Gasson, Peter; Jansen, Steven; Delzon, Sylvain
2014-01-01
Drought-induced forest dieback has been widely reported over the last decades, and the evidence for a direct causal link between survival and hydraulic failure (xylem cavitation) is now well known. Because vulnerability to cavitation is intimately linked to the anatomy of the xylem, the main objective of this study was to better understand the xylem anatomical properties associated with cavitation resistance. An extensive data set of cavitation resistance traits and xylem anatomical properties was developed for 115 conifer species, with special attention given to the micro-morphology of bordered pits. The ratio of torus to pit aperture diameter, so-called torus overlap, increased with increasing cavitation resistance, while the flexibility of the margo does not seem to play a role, suggesting that air-seeding is located at the seal between the aspirated torus and pit aperture. Moreover, punctured tori were reported in various Pinaceae species. Species resistant to cavitation had thicker tracheid walls, while their lumen diameter (conduit size) was only slightly reduced, minimizing the impact on hydraulic conductance. The results also demonstrated (i) the existence of an indirect trade-off between hydraulic safety and mechanical strength; and (ii) a consistency between species distribution and xylem anatomy: species with a wide torus overlap and high valve effects are found in arid environments such as the Mediterranean region. PMID:24916072
Burrows, G. E.; Meagher, P. F.; Heady, R. D.
2007-01-01
Background and Aims The branch-base xylem structure of the endangered Wollemia nobilis was anatomically investigated. Wollemia nobilis is probably the only extant tree species that produces only first-order branches and where all branches are cleanly abscised. An investigation was carried out to see if these unusual features might influence branch-base xylem structure and water supply to the foliage. Methods The xylem was sectioned at various distances along the branch bases of 6-year-old saplings. Huber values and relative theoretical hydraulic conductivities were calculated for various regions of the branch base. Key Results The most proximal branch base featured a pronounced xylem constriction. The constriction had only 14–31 % (average 21 %) of the cross-sectional area and 20–42 % (average 28 %) of the theoretical hydraulic conductivity of the more distal branch xylem. Wollemia nobilis had extremely low Huber values for a conifer. Conclusions The branch-base xylem constriction would appear to facilitate branch abscission, while the associated Huber values show that W. nobilis supplies a relatively large leaf area through a relatively small diameter ‘pipe’. It is tempting to suggest that the pronounced decline of W. nobilis in the Tertiary is related to its unusual branch-base structure but physiological studies of whole plant conductance are still needed. PMID:17272303
Puértolas, Jaime; Alcobendas, Rosalía; Alarcón, Juan J; Dodd, Ian C
2013-08-01
To determine how root-to-shoot abscisic acid (ABA) signalling is regulated by vertical soil moisture gradients, root ABA concentration ([ABA](root)), the fraction of root water uptake from, and root water potential of different parts of the root zone, along with bulk root water potential, were measured to test various predictive models of root xylem ABA concentration [RX-ABA](sap). Beans (Phaseolus vulgaris L. cv. Nassau) were grown in soil columns and received different irrigation treatments (top and basal watering, and withholding water for varying lengths of time) to induce different vertical soil moisture gradients. Root water uptake was measured at four positions within the column by continuously recording volumetric soil water content (θv). Average θv was inversely related to bulk root water potential (Ψ(root)). In turn, Ψ(root) was correlated with both average [ABA](root) and [RX-ABA](sap). Despite large gradients in θv, [ABA](root) and root water potential was homogenous within the root zone. Consequently, unlike some split-root studies, root water uptake fraction from layers with different soil moisture did not influence xylem sap (ABA). This suggests two different patterns of ABA signalling, depending on how soil moisture heterogeneity is distributed within the root zone, which might have implications for implementing water-saving irrigation techniques. © 2013 John Wiley & Sons Ltd.
Mistletoe ecophysiology: Host-parasite interactions
G. Glatzel; B. W. Geils
2009-01-01
Mistletoes are highly specialized perennial flowering plants adapted to parasitic life on aerial parts of their hosts. In our discussion on the physiological interactions between parasite and host, we focus on water relations, mineral nutrition, and the effect of host vigour. When host photosynthesis is greatest, the xylem water potential of the host is most negative....
Douglass F. Jacobs; Francis Salifu; Anthony Davis
2005-01-01
Transplant shock, implicated by depressed seedling physiological response associated with moisture or nutrient stress immediately following planting, limits early plantation establishment. We investigated the impacts of simulated drought and transplant root volume on predawn leaf xylem water potential, photosynthetic assimilation rates, stomatal conductance, and growth...
Nardini, Andrea; Dimasi, Federica; Klepsch, Matthias; Jansen, Steven
2012-12-01
The 'ionic effect', i.e., changes in xylem hydraulic conductivity (k(xyl)) due to variation of the ionic sap composition in vessels, was studied in four Acer species growing in contrasting environments differing in water availability. Hydraulic measurements of the ionic effect were performed together with measurements on the sap electrical conductivity, leaf water potential and vessel anatomy. The low ionic effect recorded in Acer pseudoplatanus L. and Acer campestre L. (15.8 and 14.7%, respectively), which represented two species from shady and humid habitats, was associated with a low vessel grouping index, high sap electrical conductivity and least negative leaf water potential. Opposite traits were found for Acer monspessulanum L. and Acer platanoides L., which showed an ionic effect of 23.6 and 23.1%, respectively, and represent species adapted to higher irradiance and/or lower water availability. These findings from closely related species provide additional support that the ionic effect could function as a compensation mechanism for embolism-induced loss of k(xyl), either as a result of high evaporative demand or increased risk of hydraulic failure.
Hydraulic Redistribution: A Modeling Perspective
NASA Astrophysics Data System (ADS)
Daly, E.; Verma, P.; Loheide, S. P., III
2014-12-01
Roots play a key role in the soil water balance. They extract and transport water for transpiration, which usually represents the most important soil water loss in vegetated areas, and can redistribute soil water, thereby increasing transpiration rates and enhancing root nutrient uptake. We present here a two-dimensional model capable of describing two key aspects of root water uptake: root water compensation and hydraulic redistribution. Root water compensation is the ability of root systems to respond to the reduction of water uptake from areas of the soil with low soil water potential by increasing the water uptake from the roots in soil parts with higher water potential. Hydraulic redistribution is a passive transfer of water through the root system from areas of the soil with greater water potential to areas with lower water potential. Both mechanisms are driven by gradients of water potential in the soil and the roots. The inclusion of root water compensation and hydraulic redistribution in models can be achieved by describing root water uptake as a function of the difference in water potential between soil and root xylem. We use a model comprising the Richards equation for the water flow in variably saturated soils and the Darcy's equation for the water flow in the xylem. The two equations are coupled via a sink term, which is assumed to be proportional to the difference between soil and xylem water potentials. The model is applied in two case studies to describe vertical and horizontal hydraulic redistribution and the interaction between vegetation with different root depths. In the case of horizontal redistribution, the model is used to reproduce the fluxes of water across the root system of a tree subjected to uneven irrigation. This example can be extended to situations when only part of the root system has access to water, such as vegetation near creeks, trees at the edge of forests, and street trees in urban areas. The second case is inspired by recent agro-ecosystems experiments that combined different vegetation species to increase crop yield. The presence of deep rooted plants (nursing species) near shallow rooted crops (nursed species) enhanced crop growth thanks to vertical and horizontal hydraulic redistribution. The model is able to reproduce the patterns of water redistribution observed in this scenario.
A mathematical model of water and nutrient transport in xylem vessels of a wheat plant.
Payvandi, S; Daly, K R; Jones, D L; Talboys, P; Zygalakis, K C; Roose, T
2014-03-01
At a time of increasing global demand for food, dwindling land and resources, and escalating pressures from climate change, the farming industry is undergoing financial strain, with a need to improve efficiency and crop yields. In order to improve efficiencies in farming, and in fertiliser usage in particular, understanding must be gained of the fertiliser-to-crop-yield pathway. We model one aspect of this pathway; the transport of nutrients within the vascular tissues of a crop plant from roots to leaves. We present a mathematical model of the transport of nutrients within the xylem vessels in response to the evapotranspiration of water. We determine seven different classes of flow, including positive unidirectional flow, which is optimal for nutrient transport from the roots to the leaves; and root multidirectional flow, which is similar to the hydraulic lift process observed in plants. We also investigate the effect of diffusion on nutrient transport and find that diffusion can be significant at the vessel termini especially if there is an axial efflux of nutrient, and at night when transpiration is minimal. Models such as these can then be coupled to whole-plant models to be used for optimisation of nutrient delivery scenarios.
Nanobubbles: a new paradigm for air-seeding in xylem.
Schenk, H Jochen; Steppe, Kathy; Jansen, Steven
2015-04-01
Long-distance water transport in plants relies on a system that typically operates under negative pressure and is prone to hydraulic failure due to gas bubble formation. One primary mechanism of bubble formation takes place at nanoporous pit membranes between neighboring conduits. We argue that this process is likely to snap off nanobubbles because the local increase in liquid pressure caused by entry of air-water menisci into the complex pit membrane pores would energetically favor nanobubble formation over instant cavitation. Nanobubbles would be stabilized by surfactants and by gas supersaturation of the sap, may dissolve, fragment into smaller bubbles, or create embolisms. The hypothesis that safe and stable nanobubbles occur in plants adds a new component supporting the cohesion-tension theory. Copyright © 2015 Elsevier Ltd. All rights reserved.
NASA Astrophysics Data System (ADS)
Christoffersen, Bradley O.; Gloor, Manuel; Fauset, Sophie; Fyllas, Nikolaos M.; Galbraith, David R.; Baker, Timothy R.; Kruijt, Bart; Rowland, Lucy; Fisher, Rosie A.; Binks, Oliver J.; Sevanto, Sanna; Xu, Chonggang; Jansen, Steven; Choat, Brendan; Mencuccini, Maurizio; McDowell, Nate G.; Meir, Patrick
2016-11-01
Forest ecosystem models based on heuristic water stress functions poorly predict tropical forest response to drought partly because they do not capture the diversity of hydraulic traits (including variation in tree size) observed in tropical forests. We developed a continuous porous media approach to modeling plant hydraulics in which all parameters of the constitutive equations are biologically interpretable and measurable plant hydraulic traits (e.g., turgor loss point πtlp, bulk elastic modulus ɛ, hydraulic capacitance Cft, xylem hydraulic conductivity ks,max, water potential at 50 % loss of conductivity for both xylem (P50,x) and stomata (P50,gs), and the leaf : sapwood area ratio Al : As). We embedded this plant hydraulics model within a trait forest simulator (TFS) that models light environments of individual trees and their upper boundary conditions (transpiration), as well as providing a means for parameterizing variation in hydraulic traits among individuals. We synthesized literature and existing databases to parameterize all hydraulic traits as a function of stem and leaf traits, including wood density (WD), leaf mass per area (LMA), and photosynthetic capacity (A
Linking xylem water storage with anatomical parameters in five temperate tree species.
Jupa, Radek; Plavcová, Lenka; Gloser, Vít; Jansen, Steven
2016-06-01
The release of water from storage compartments to the transpiration stream is an important functional mechanism that provides the buffering of sudden fluctuations in water potential. The ability of tissues to release water per change in water potential, referred to as hydraulic capacitance, is assumed to be associated with the anatomy of storage tissues. However, information about how specific anatomical parameters determine capacitance is limited. In this study, we measured sapwood capacitance (C) in terminal branches and roots of five temperate tree species (Fagus sylvatica L., Picea abies L., Quercus robur L., Robinia pseudoacacia L., Tilia cordata Mill.). Capacitance was calculated separately for water released mainly from capillary (CI; open vessels, tracheids, fibres, intercellular spaces and cracks) and elastic storage compartments (CII; living parenchyma cells), corresponding to two distinct phases of the moisture release curve. We found that C was generally higher in roots than branches, with CI being 3-11 times higher than CII Sapwood density and the ratio of dead to living xylem cells were most closely correlated with C In addition, the magnitude of CI was strongly correlated with fibre/tracheid lumen area, whereas CII was highly dependent on the thickness of axial parenchyma cell walls. Our results indicate that water released from capillary compartments predominates over water released from elastic storage in both branches and roots, suggesting the limited importance of parenchyma cells for water storage in juvenile xylem of temperate tree species. Contrary to intact organs, water released from open conduits in our small wood samples significantly increased CI at relatively high water potentials. Linking anatomical parameters with the hydraulic capacitance of a tissue contributes to a better understanding of water release mechanisms and their implications for plant hydraulics. © The Author 2016. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.
Temporal and spatial patterns of internal and external stem CO2 fluxes in a sub-Mediterranean oak.
Salomón, Roberto L; Valbuena-Carabaña, María; Gil, Luis; McGuire, Mary Anne; Teskey, Robert O; Aubrey, Doug P; González-Doncel, Inés; Rodríguez-Calcerrada, Jesús
2016-11-01
To accurately estimate stem respiration (R S ), measurements of both carbon dioxide (CO 2 ) efflux to the atmosphere (E A ) and internal CO 2 flux through xylem (F T ) are needed because xylem sap transports respired CO 2 upward. However, reports of seasonal dynamics of F T and E A are scarce and no studies exist in Mediterranean species under drought stress conditions. Internal and external CO 2 fluxes at three stem heights, together with radial stem growth, temperature, sap flow and shoot water potential, were measured in Quercus pyrenaica Willd. in four measurement campaigns during one growing season. Substantial daytime depressions in temperature-normalized E A were observed throughout the experiment, including prior to budburst, indicating that diel hysteresis between stem temperature and E A cannot be uniquely ascribed to diversion of CO 2 in the transpiration stream. Low internal [CO 2 ] (<0.5%) resulted in low contributions of F T to R S throughout the growing season, and R S was mainly explained by E A (>90%). Internal [CO 2 ] was found to vary vertically along the stems. Seasonality in resistance to radial CO 2 diffusion was related to shoot water potential. The low internal [CO 2 ] and F T observed in our study may result from the downregulation of xylem respiration in response to a legacy of coppicing as well as high radial diffusion of CO 2 through cambium, phloem and bark tissues, which was related to low water content of stems. Long-term studies analyzing temporal and spatial variation in internal and external CO 2 fluxes and their interactions are needed to mechanistically understand and model respiration of woody tissues. © The Author 2016. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.
Stem anatomy and relative growth rate in seedlings of a wide range of woody plant species and types.
Castro-Díez, P; Puyravaud, J P; Cornelissen, J H C; Villar-Salvador, P
1998-08-01
Stem traits were analysed in laboratory-grown seedlings of 80 European woody and semiwoody species of known potential relative growth rate (RGR) and of similar ontogenetic phase. The objectives were, firstly, to assess the relation between stem structure and plant growth potential and, secondly, to explore how stem structure varies among species differing in life form and leaf habit. Hydraulic conductance was represented by the mean diameter of the widest xylem conduits (Dmax), and structural strength by the percentage of xylem tissue occupied by cell wall material (CWx) or stem tissue density (SD). Across all species RGR showed a weak positive correlation with Dmax and weak negative ones with CWx and SD, with slow-growers showing great dispersion of stem trait values. In the RGR-Dmax relationship this dispersion disappeared when trees were removed from the analysis. None of the relationships were significant among tree species alone. It was suggested that fast-growers require a xylem with wide conduits (high Dmax) to achieve high hydraulic conductivity, and "cheaply" constructed stems (low CWx and SD) to maximise allocation to leaves. However, the possession of such traits does not guarantee fast growth, as other factors may constrain RGR elsewhere in the plant. Deciduous seedlings showed higher Dmax and lower CWx than evergreens. Higher Dmax could reflect an innate higher tolerance of conductivity loss by freeze-induced embolism in deciduous plants, which are not burdened by the maintenance of foliage in winter. In contrast, life forms were differentiated most clearly by SD. For instance, shrub seedlings had less dense stem tissues than tree seedlings, possibly because they need less investment in long-term strength and stature.
Gričar, Jožica; Prislan, Peter; de Luis, Martin; Gryc, Vladimír; Hacurová, Jana; Vavrčík, Hanuš; Čufar, Katarina
2015-01-01
There is limited information on intra-annual plasticity of secondary tissues of tree species growing under different environmental conditions. To increase the knowledge about the plasticity of secondary growth, which allows trees to adapt to specific local climatic regimes, we examined climate–radial growth relationships of Norway spruce [Picea abies (L.) H. Karst.] from three contrasting locations in the temperate climatic zone by analyzing tree-ring widths for the period 1932–2010, and cell characteristics in xylem and phloem increments formed in the years 2009–2011. Variation in the structure of xylem and phloem increments clearly shows that plasticity in seasonal dynamics of cambial cell production and cell differentiation exists on xylem and phloem sides. Anatomical characteristics of xylem and phloem cells are predominantly site-specific characteristics, because they varied among sites but were fairly uniform among years in trees from the same site. Xylem and phloem tissues formed in the first part of the growing season seemed to be more stable in structure, indicating their priority over latewood and late phloem for tree performance. Long-term climate and radial growth analyses revealed that growth was in general less dependent on precipitation than on temperature; however, growth sensitivity to local conditions differed among the sites. Only partial dependence of radial growth of spruce on climatic factors on the selected sites confirms its strategy to adapt the structure of wood and phloem increments to function optimally in local conditions. PMID:26442044
[Seasonal development of phloem in Scots pine stems].
Antonova, G F; Stasova, V V
2006-01-01
The formation of phloem was studied for two years in stems of 50 to 60 year old trees of Scots pine (Pinus sylvestris L.) growing in nature. The development of phloem of the current year begins 10 to 20 days before the xylem formation and is completed with the termination of shoot growth in the end of June. Observations over the seasonal activity of cambium producing sieve-like cells of phloem and duration of their differentiation as compared to the xylem derivatives of cambium have shown that the maxima of formation of phloem and xylem cells could coincide or not coincide by season, while the activities of their differentiation were always at antiphase. The sieve-like cells of early phloem were separated from those of late phloem by a layer of tannin-containing cells, which are formed simultaneously with the formation of late xylem cells by the cambium. Seasonal dynamics of accumulation of starch grain in structural elements of the phloem is related to the xylem development. The content of metabolites in differentiating and mature phloem elements, in the cambium zone, and in the xylem cells growing in the radial direction depended on cell specificity, stage of their development, and type of forming wood, early or late, which differ in the cell wall parameters and, hence, requirement of assimilates. Significant differences were described between the content of low molecular weigh carbohydrates, amino acids, organic acids, and phenol compounds using two methods of calculation: per dry weight and per cell.
Perrin, Magali; Rossi, Sergio; Isabel, Nathalie
2017-05-01
Bud and cambial phenology represent the adaptation of species to the local environment that allows the growing season to be maximized while minimizing the risk of frost for the developing tissues. The temporal relationship between the apical and radial meristems can help in the understanding of tree growth as a whole process. The aim of this study was to compare cambial phenology in black spruce (Picea mariana (Mill.) B.S.P.) provenances classified as early and late bud flushing. The different phases of cambial phenology were assessed on wood microcores sampled weekly from April to October in 2014 and 2015 from 61 trees growing in a provenance trial in Quebec, Canada. Trees showing an early bud flush also exhibited early reactivation of xylem differentiation, although an average difference of 12 days for buds corresponded to small although significant differences of 4 days for xylem. Provenances with early bud flush had an early bud set and completed xylem formation earlier than late bud flush provenances. No significant difference in the period of xylem formation and total growth was observed between the flushing classes. Our results demonstrate that the ecotype differentiation of black spruce provenances represented by the phenological adaptation of buds to the local climate corresponds to specific growth dynamics of the xylem. © The Author 2017. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.
Pérez-Donoso, Alonso G; Greve, L Carl; Walton, Jeffrey H; Shackel, Ken A; Labavitch, John M
2007-02-01
It is conventionally thought that multiplication of the xylem-limited bacterium Xylella fastidiosa (Xf) within xylem vessels is the sole factor responsible for the blockage of water movement in grapevines (Vitis vinifera) affected by Pierce's disease. However, results from our studies have provided substantial support for the idea that vessel obstructions, and likely other aspects of the Pierce's disease syndrome, result from the grapevine's active responses to the presence of Xf, rather than to the direct action of the bacterium. The use of magnetic resonance imaging (MRI) to observe the distribution of water within the xylem has allowed us to follow nondestructively the development of vascular system obstructions subsequent to inoculation of grapevines with Xf. Because we have hypothesized a role for ethylene produced in vines following infection, the impact of vine ethylene exposure on obstruction development was also followed using MRI. In both infected and ethylene-exposed plants, MRI shows that an important proportion of the xylem vessels become progressively air embolized after the treatments. The loss of xylem water-transporting function, assessed by MRI, has been also correlated with a decrease in stem-specific hydraulic conductivity (K(S)) and the presence of tyloses in the lumens of obstructed water conduits. We have observed that the ethylene production of leaves from infected grapevines is greater than that from healthy vines and, therefore, propose that ethylene may be involved in a series of cellular events that coordinates the vine's response to the pathogen.
De novo transcriptome assemblies of four xylem sap-feeding insects
Tassone, Erica E.; Cowden, Charles C.
2017-01-01
Abstract Background: Spittle bugs and sharpshooters are well-known xylem sap-feeding insects and vectors of the phytopathogenic bacterium Xylella fastidiosa (Wells), a causal agent of Pierce's disease of grapevines and other crop diseases. Specialized feeding on nutrient-deficient xylem sap is relatively rare among insect herbivores, and only limited genomic and transcriptomic information has been generated for xylem-sap feeders. To develop a more comprehensive understanding of biochemical adaptations and symbiotic relationships that support survival on a nutritionally austere dietary source, transcriptome assemblies for three sharpshooter species and one spittlebug species were produced. Findings: Trinity-based de novo transcriptome assemblies were generated for all four xylem-sap feeders using raw sequencing data originating from whole-insect preps. Total transcripts for each species ranged from 91 384 for Cuerna arida to 106 998 for Homalodisca liturata with transcript totals for Graphocephala atropunctata and the spittlebug Clastoptera arizonana falling in between. The percentage of transcripts comprising complete open reading frames ranged from 60% for H. liturata to 82% for C. arizonana. Bench-marking universal single-copy orthologs analyses for each dataset indicated quality assemblies and a high degree of completeness for all four species. Conclusions: These four transcriptomes represent a significant expansion of data for insect herbivores that feed exclusively on xylem sap, a nutritionally deficient dietary source relative to other plant tissues and fluids. Comparison of transcriptome data with insect herbivores that utilize other dietary sources may illuminate fundamental differences in the biochemistry of dietary specialization. PMID:28327966
A broad survey of hydraulic and mechanical safety in the xylem of conifers.
Bouche, Pauline S; Larter, Maximilien; Domec, Jean-Christophe; Burlett, Régis; Gasson, Peter; Jansen, Steven; Delzon, Sylvain
2014-08-01
Drought-induced forest dieback has been widely reported over the last decades, and the evidence for a direct causal link between survival and hydraulic failure (xylem cavitation) is now well known. Because vulnerability to cavitation is intimately linked to the anatomy of the xylem, the main objective of this study was to better understand the xylem anatomical properties associated with cavitation resistance. An extensive data set of cavitation resistance traits and xylem anatomical properties was developed for 115 conifer species, with special attention given to the micro-morphology of bordered pits. The ratio of torus to pit aperture diameter, so-called torus overlap, increased with increasing cavitation resistance, while the flexibility of the margo does not seem to play a role, suggesting that air-seeding is located at the seal between the aspirated torus and pit aperture. Moreover, punctured tori were reported in various Pinaceae species. Species resistant to cavitation had thicker tracheid walls, while their lumen diameter (conduit size) was only slightly reduced, minimizing the impact on hydraulic conductance. The results also demonstrated (i) the existence of an indirect trade-off between hydraulic safety and mechanical strength; and (ii) a consistency between species distribution and xylem anatomy: species with a wide torus overlap and high valve effects are found in arid environments such as the Mediterranean region. © The Author 2014. Published by Oxford University Press on behalf of the Society for Experimental Biology.
Castagneri, Daniele; Petit, Giai; Carrer, Marco
2015-12-01
Climate change can induce substantial modifications in xylem structure and water transport capacity of trees exposed to environmental constraints. To elucidate mechanisms of xylem plasticity in response to climate, we retrospectively analysed different cell anatomical parameters over tree-ring series in Norway spruce (Picea abies L. Karst.). We sampled 24 trees along an altitudinal gradient (1200, 1600 and 2100 m above sea level, a.s.l.) and processed 2335 ± 1809 cells per ring. Time series for median cell lumen area (MCA), cell number (CN), tree-ring width (RW) and tree-ring-specific hydraulic conductivity (Kr) were crossed with daily temperature and precipitation records (1926-2011) to identify climate influence on xylem anatomical traits. Higher Kr at the low elevation site was due to higher MCA and CN. These variables were related to different aspects of intra-seasonal climatic variability under different environmental conditions, with MCA being more sensitive to summer precipitation. Winter precipitation (snow) benefited most parameters in all the sites. Descending the gradient, sensitivity of xylem features to summer climate shifted mostly from temperature to precipitation. In the context of climate change, our results indicate that higher summer temperatures at high elevations will benefit cell production and xylem hydraulic efficiency, whereas reduced water availability at lower elevations could negatively affect tracheids enlargement and thus stem capacity to transport water. © The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.
Ishikawa, Tetsuya; Shabala, Sergey
2018-05-15
Control of xylem Na + loading has often been named as the essential components of salinity tolerance mechanism. However, it is less clear to what extent the difference in this trait may determine differential salinity tolerance between species. In this study barley (Hordeum vulgare L. cv. CM72) and rice (Oryza sativa L. cv Dongjin) plants were grown under two levels of salinity. Na + and K + concentrations in the xylem sap, and shoot and root tissues were measured at different time points after stress onset. Salt-exposed rice plants prevented xylem Na + loading for several days, but failed to control this process in the longer term, ultimately resulting in a massive Na + shoot loading. Barley plants quickly increased xylem Na + concentration and its delivery to the shoot (most likely for the purpose of osmotic adjustment) but were able to reduce this process later on, keeping most of accumulated Na + in the root, thus maintaining non-toxic shoot Na + level. Rice plants increased shoot K + concentration, while barley plants maintained higher root K + concentration. Control of xylem Na + loading is remarkably different between rice and barley; this difference may differentiate the extent of the salinity tolerance between species. This trait should be investigated in more details to be used in the breeding programs aimed to improve salinity tolerance in crops. This article is protected by copyright. All rights reserved. This article is protected by copyright. All rights reserved.
Karuppanapandian, T; Geilfus, C-M; Mühling, K-H; Novák, O; Gloser, V
2017-02-01
Changes in pH of the apoplast have recently been discussed as an important factor in adjusting transpiration and water relations under conditions of drought via modulatory effect on abscisic acid (ABA) concentration. Using Vicia faba L., we investigated whether changes in the root, shoot and leaf apoplastic pH correlated with (1) a drought-induced reduction in transpiration and with (2) changes in ABA concentration. Transpiration, leaf water potential and ABA in leaves were measured and correlated with root and shoot xylem pH, determined by a pH microelectrode, and pH of leaf apoplast quantified by microscopy-based in vivo ratiometric analysis. Results revealed that a reduction in transpiration rate in the early phase of soil drying could not be linked with changes in the apoplastic pH via effects on the stomata-regulating hormone ABA. Moreover, drought-induced increase in pH of xylem or leaf apoplast was not the remote effect of an acropetal transport of alkaline sap from root, because root xylem acidified during progressive soil drying, whereas the shoot apoplast alkalized. We reason that other, yet unknown signalling mechanism was responsible for reduction of transpiration rate in the early phase of soil drying. Copyright © 2016 Elsevier Ireland Ltd. All rights reserved.
Casting light on xylem vulnerability in an herbaceous species reveals a lack of segmentation.
Skelton, Robert P; Brodribb, Timothy J; Choat, Brendan
2017-04-01
Finding thresholds at which loss of plant functionality occurs during drought is critical for predicting future crop productivity and survival. Xylem resistance to embolism has been suggested as a key trait associated with water-stress tolerance. Although a substantial literature exists describing the vulnerability of woody stems to embolism, leaves and roots of herbaceous species remain under-represented. Also, little is known about vulnerability to embolism at a whole-plant scale or propagation of embolism within plants. New techniques to view the process of embolism formation provide opportunities to resolve long-standing questions. Here, we used multiple visual techniques, including X-ray micro-computed tomography and the optical vulnerability method, to investigate the spread of embolism within intact stems, leaves and roots of Solanum lycopersicum (common tomato). We found that roots, stems and leaves of tomato plants all exhibited similar vulnerability to embolism, suggesting that embolism rapidly propagates among tissues. Although we found scarce evidence for differentiation of xylem vulnerability among tissues at the scale of the whole plant, within a leaf the midrib embolized at higher water potentials than lower order veins. Substantial overlap between the onset of cavitation and incipient leaf damage suggests that cavitation represents a substantial damage to plants, but the point of lethal cavitation in this herbaceous species remains uncertain. © 2017 The Authors. New Phytologist © 2017 New Phytologist Trust.
Cell Wall Modifications in Arabidopsis Plants with Altered α-l-Arabinofuranosidase Activity[C][W
Chávez Montes, Ricardo A.; Ranocha, Philippe; Martinez, Yves; Minic, Zoran; Jouanin, Lise; Marquis, Mélanie; Saulnier, Luc; Fulton, Lynette M.; Cobbett, Christopher S.; Bitton, Frédérique; Renou, Jean-Pierre; Jauneau, Alain; Goffner, Deborah
2008-01-01
Although cell wall remodeling is an essential feature of plant growth and development, the underlying molecular mechanisms are poorly understood. This work describes the characterization of Arabidopsis (Arabidopsis thaliana) plants with altered expression of ARAF1, a bifunctional α-l-arabinofuranosidase/β-d-xylosidase (At3g10740) belonging to family 51 glycosyl-hydrolases. ARAF1 was localized in several cell types in the vascular system of roots and stems, including xylem vessels and parenchyma cells surrounding the vessels, the cambium, and the phloem. araf1 T-DNA insertional mutants showed no visible phenotype, whereas transgenic plants that overexpressed ARAF1 exhibited a delay in inflorescence emergence and altered stem architecture. Although global monosaccharide analysis indicated only slight differences in cell wall composition in both mutant and overexpressing lines, immunolocalization experiments using anti-arabinan (LM6) and anti-xylan (LM10) antibodies indicated cell type-specific alterations in cell wall structure. In araf1 mutants, an increase in LM6 signal intensity was observed in the phloem, cambium, and xylem parenchyma in stems and roots, largely coinciding with ARAF1 expression sites. The ectopic overexpression of ARAF1 resulted in an increase in LM10 labeling in the secondary walls of interfascicular fibers and xylem vessels. The combined ARAF1 gene expression and immunolocalization studies suggest that arabinan-containing pectins are potential in vivo substrates of ARAF1 in Arabidopsis. PMID:18344421
Hudson, P J; Limousin, J M; Krofcheck, D J; Boutz, A L; Pangle, R E; Gehres, N; McDowell, N G; Pockman, W T
2018-02-01
Hydraulic architecture imposes a fundamental control on water transport, underpinning plant productivity, and survival. The extent to which hydraulic architecture of mature trees acclimates to chronic drought is poorly understood, limiting accuracy in predictions of forest responses to future droughts. We measured seasonal shoot hydraulic performance for multiple years to assess xylem acclimation in mature piñon (Pinus edulis) and juniper (Juniperus monosperma) after 3+ years of precipitation manipulation. Our treatments consisted of water addition (+20% ambient precipitation), partial precipitation-exclusion (-45% ambient precipitation), and exclusion-structure control. Supplemental watering elevated leaf water potential, sapwood-area specific hydraulic conductivity, and leaf-area specific hydraulic conductivity relative to precipitation exclusion. Shifts in allocation of leaf area to sapwood area enhanced differences between irrigated and droughted K L in piñon but not juniper. Piñon and juniper achieved similar K L under ambient conditions, but juniper matched or outperformed piñon in all physiological measurements under both increased and decreased precipitation treatments. Embolism vulnerability and xylem anatomy were unaffected by treatments in either species. Absence of significant acclimation combined with inferior performance for both hydraulic transport and safety suggests piñon has greater risk of local extirpation if aridity increases as predicted in the southwestern USA. © 2017 John Wiley & Sons Ltd.
Separation and characterization of needle and xylem maritime pine proteins.
Costa, P; Pionneau, C; Bauw, G; Dubos, C; Bahrmann, N; Kremer, A; Frigerio, J M; Plomion, C
1999-01-01
Two-dimensional gel electrophoresis (2-DE) and image analysis are currently used for proteome analysis in maritime pine (Pinus pinaster Ait.). This study presents a database of expressed proteins extracted from needles and xylem, two important tissues for growth and wood formation. Electrophoresis was carried out by isoelectric focusing (IEF) in the first dimension and sodium dodecyl sulfate-polyacrylamide gel electrophoresis (SDS-PAGE) in the second. Silver staining made it possible to detect an average of 900 and 600 spots on 2-DE gels from needles and xylem, respectively. A total of 28 xylem and 35 needle proteins were characterized by internal peptide microsequencing. Out of these 63 proteins, 57 (90%) could be identified based on amino acid similarity with known proteins, of which 24 (42%) have already been described in conifers. Overall comparison of both tissues indicated that 29% and 36% of the spots were specific to xylem and needles, respectively, while the other spots were of identical molecular weight and isoelectric point. The homology of spot location in 2-DE patterns was further validated by sequence analysis of proteins present in both tissues. A proteomic database of maritime pine is accessible on the internet (http://www.pierroton.inra.fr/genetics/2D/).
de Silva, Nayana Dilini Gardiyehewa; Cholewa, Ewa; Ryser, Peter
2012-10-01
The effects of heavy metal stress, drought stress, and their combination on xylem structure in red maple (Acer rubrum) seedlings were investigated in an outdoor pot experiment. As metal-contaminated substrate, a mixture of 1.5% slag with sand was used, with Ni, Cu, Co, and Cr as the main contaminants. Plants grown on contaminated substrate had increased leaf metal concentrations. The two stresses reduced plant growth in an additive manner. The effects of metal and drought stresses on xylem characteristics were similar to each other, with a reduced proportion of xylem tissue, reduced conduit density in stems, and reduced conduit size in the roots. This resulted, in both stems and roots, in reductions in hydraulic conductance, xylem-specific conductivity, and leaf-specific conductivity. The similarity of the responses to the two stresses suggests that the plants' response to metals was actually a drought response, probably due to the reduced water uptake capacity of the metal-exposed roots. The only plant responses specific to metal stress were decreasing trends of stomatal density and chlorophyll content. In conclusion, the exposure to metals aggravates water stress in an additive manner, making the plants more vulnerable to drought.
NASA Astrophysics Data System (ADS)
Bittner, S.; Priesack, E.
2012-04-01
We apply a functional-structural model of tree water flow to single old-growth trees in a temperate broad-leaved forest stand. Roots, stems and branches are represented by connected porous cylinder elements further divided into the inner heartwood cylinders surrounded by xylem and phloem. Xylem water flow is simulated by applying a non-linear Darcy flow in porous media driven by the water potential gradient according to the cohesion-tension theory. The flow model is based on physiological input parameters such as the hydraulic conductivity, stomatal response to leaf water potential and root water uptake capability and, thus, can reflect the different properties of tree species. The actual root water uptake is calculated using also a non-linear Darcy law based on the gradient between root xylem water potential and rhizosphere soil water potential and by the simulation of soil water flow applying Richards equation. A leaf stomatal conductance model is combined with the hydrological tree and soil water flow model and a spatially explicit three-dimensional canopy light model. The structure of the canopy and the tree architectures are derived by applying an automatic tree skeleton extraction algorithm from point clouds obtained by use of a terrestrial laser scanner allowing an explicit representation of the water flow path in the stem and branches. The high spatial resolution of the root and branch geometry and their connectivity makes the detailed modelling of the water use of single trees possible and allows for the analysis of the interaction between single trees and the influence of the canopy light regime (including different fractions of direct sunlight and diffuse skylight) on the simulated sap flow and transpiration. The model can be applied at various sites and to different tree species, enabling the up-scaling of the water usage of single trees to the total transpiration of mixed stands. Examples are given to reveal differences between diffuse- and ring-porous tree species and to simulate the diurnal dynamics of transpiration, stem sap flux, and root water uptake observed during the vegetation period in the year 2009.
Large and small plants of creosotebush and mesquite were subjected to drought during summer 1993. arge and small plants responded to drought with lowered xylem water potential and lowered photosynthetic gas exchange. arge plants appear to maintain a reduced but constant photosynt...
NASA Astrophysics Data System (ADS)
Smith, A.; Tetzlaff, D.; Soulsby, C.
2017-12-01
Identifying the sources of water which sustain plant water uptake is an essential prerequisite to understanding the interactions of vegetation and water within the critical zone. Estimating the sources of root-water uptake is complicated by ecohydrological separation, or the notion of "two-water worlds" which distinguishes more mobile and immobile water sources which respectively sustain streamflow and evapotranspiration. Water mobility within the soil determines both the transit time/residence time of water through/in soils and the subsequent age of root-uptake and xylem water. We used time-variant StorAge Selection (SAS) functions to conceptualise the transit/residence times in the critical zone using a dual-storage soil column differentiating gravity (mobile) and tension dependent (immobile) water, calibrated to measured stable isotope signatures of soil water. Storage-discharge relationships [Brutsaert and Nieber, 1977] were used to identify gravity and tension dependent storages. A temporally variable distribution for root water uptake was identified using simulated stable isotopes in xylem and soil water. Composition of δ2H and δ18O was measured in soil water at 4 depths (5, 10, 15, and 20 cm) on 10 occasions, and 5 times for xylem water within the dominant heather (Calluna sp. and Erica sp.) vegetation in a Scottish Highland catchment over a two-year period. Within a 50 cm soil column, we found that more than 53% of the total stored water was water that was present before the start of the simulation. Mean residence times of the mobile water in the upper 20 cm of the soil were 16, 25, 36, and 44 days, respectively. Mean evaporation transit time varied between 9 and 40 days, driven by seasonal changes and precipitation events. Lastly, mean transit times of xylem water ranged between 95-205 days, driven by changes in soil moisture. During low soil moisture (i.e. lower than mean soil moisture), root-uptake was from lower depths, while higher than mean soil moisture showed preferential uptake of near surface water. In our humid, low energy environment, we found that xylem water is comprised of both mobile and immobile water. The division of soil storage into two storages, gravity and tension dependent, has shown potential to identify the sources of plant water and vegetation and soil water interactions.
Backus, Elaine A; Andrews, Kim B; Shugart, Holly J; Carl Greve, L; Labavitch, John M; Alhaddad, Hasan
2012-07-01
A few phytophagous hemipteran species such as the glassy-winged sharpshooter, Homalodisca vitripennis, (Germar), subsist entirely on xylem fluid. Although poorly understood, aspects of the insect's salivary physiology may facilitate both xylem-feeding and transmission of plant pathogens. Xylella fastidiosa is a xylem-limited bacterium that causes Pierce's disease of grape and other scorch diseases in many important crops. X. fastidiosa colonizes the anterior foregut (precibarium and cibarium) of H. vitripennis and other xylem-feeding vectors. Bacteria form a dense biofilm anchored in part by an exopolysaccharide (EPS) matrix that is reported to have a β-1,4-glucan backbone. Recently published evidence supports the following, salivation-egestion hypothesis for the inoculation of X. fastidiosa during vector feeding. The insect secretes saliva into the plant and then rapidly takes up a mixture of saliva and plant constituents. During turbulent fluid movements in the precibarium, the bacteria may become mechanically and enzymatically dislodged; the mixture is then egested back out through the stylets into plant cells, possibly including xylem vessels. The present study found that proteins extracted from dissected H. vitripennis salivary glands contain several enzyme activities capable of hydrolyzing glycosidic linkages in polysaccharides such as those found in EPS and plant cell walls, based on current information about the structures of those polysaccharides. One of these enzymes, a β-1,4-endoglucanase (EGase) was enriched in the salivary gland protein extract by subjecting the extract to a few, simple purification steps. The EGase-enriched extract was then used to generate a polyclonal antiserum that was used for immunohistochemical imaging of enzymes in sharpshooter salivary sheaths in grape. Results showed that enzyme-containing gelling saliva is injected into xylem vessels during sharpshooter feeding, in one case being carried by the transpiration stream away from the injection site. Thus, the present study provides support for the salivation-egestion hypothesis. Published by Elsevier Ltd.
Plant GSK3 proteins regulate xylem cell differentiation downstream of TDIF-TDR signalling
NASA Astrophysics Data System (ADS)
Kondo, Yuki; Ito, Tasuku; Nakagami, Hirofumi; Hirakawa, Yuki; Saito, Masato; Tamaki, Takayuki; Shirasu, Ken; Fukuda, Hiroo
2014-03-01
During plant radial growth typically seen in trees, procambial and cambial cells act as meristematic cells in the vascular system to self-proliferate and differentiate into xylem cells. These two processes are regulated by a signalling pathway composed of a peptide ligand and its receptor; tracheary element differentiation inhibitory factor (TDIF) and TDIF RECEPTOR (TDR). Here we show that glycogen synthase kinase 3 proteins (GSK3s) are crucial downstream components of the TDIF signalling pathway suppressing xylem differentiation from procambial cells. TDR interacts with GSK3s at the plasma membrane and activates GSK3s in a TDIF-dependent fashion. Consistently, a specific inhibitor of plant GSK3s strongly induces xylem cell differentiation through BRI1-EMS SUPPRESSOR 1 (BES1), a well-known target transcription factor of GSK3s. Our findings provide insight into the regulation of cell fate determination in meristem maintenance.
Palaeo-adaptive properties of the xylem of Metasequoia: mechanical/hydraulic compromises.
Jagels, Richard; Visscher, George E; Lucas, John; Goodell, Barry
2003-07-01
The xylem of Metasequoia glyptostroboides Hu et Cheng is characterized by very low density (average specific gravity = 0.27) and tracheids with relatively large dimensions (length and diameter). The microfibril angle in the S2 layer of tracheid walls is large, even in outer rings, suggesting a cambial response to compressive rather than tensile stresses. In some cases, this compressive stress is converted to irreversible strain (plastic deformation), as evidenced by cell wall corrugations. The heartwood is moderately decay resistant, helping to prevent Brazier buckling. These xylem properties are referenced to the measured bending properties of modulus of rupture and modulus of elasticity, and compared with other low-to-moderate density conifers. The design strategy for Metasequoia is to produce a mechanically weak but hydraulically efficient xylem that permits rapid height growth and crown development to capture and dominate a wet site environment. The adaptability of these features to a high-latitude Eocene palaeoenvironment is discussed.
Halis, Youcef; Mayouf, Rabah; Benhaddya, Mohamed Lamine; Belhamra, Mohamed
2013-03-01
The main objective of this study was to evaluate the role of intervessel contacts in determining the patterns of hydraulic integration both within and between xylem sectors. The degree of intervessel contacts and the lateral exchange capability within and between sectors were examined and correlated in different xeric shrubs. A dye injection method was used to detect the connections between vessels; an apoplastic dye was sucked through a known number of vessels and its distribution in the xylem network was followed. Hydraulic techniques were used to measure axial and tangential conductivity both within and between xylem sectors. The intra- and inter-sector integration indexes were then determined as the ratio of tangential to axial conductance. Species differed significantly in the degree of intervessel contacts, intra- and inter-sector integration index. In all cases, hydraulic integration was observed to be higher within sector than between sectors. From the correlation analyses, the intervessel contacts showed a very weak relationship with inter-sector integration index and a strong positive relationship with intra-sector integration index. Results suggested that (1) the factors affecting patterns of lateral flow within xylem sectors might be relatively different from those between sectors. (2) The degree of intervessel contacts was a major determinant of hydraulic integration within the same xylem sector. (3) Intervessel connectivity alone was a poor predictor of hydraulic integration between different sectors, implying a significant contribution of other anatomical, physiological and environmental factors in determining the patterns of integrated-sectored transport within woody stems.
Uptake, transport, distribution and Bio-effects of SiO2 nanoparticles in Bt-transgenic cotton.
Le, Van Nhan; Rui, Yukui; Gui, Xin; Li, Xuguang; Liu, Shutong; Han, Yaning
2014-12-05
SiO2 nanoparticle is one of the most popular nanomaterial which has been used in various fields, such as wastewater treatment, environmental remediation, food processing, industrial and household applications, biomedicine, disease labeling, and biosensor, etc. In agriculture, the use of SiO2 nanoparticles as insecticide, carriers in drug delivery, or in uptake and translocation of nutrient elements, etc., has been given attention. However, the effects of nanoparticles on plants have been seldom studied. In this work, the toxicity of SiO2 nanoparticles and their uptake, transport, distribution and bio-effects have been investigated in Bt-transgenic cotton. The phytotoxic effects of SiO2 nanoparticles were exhibited in Bt-transgenic cotton with different SiO2 concentrations of 0, 10, 100, 500 and 2000 mg.L(-1) for 3 weeks through dry biomasses, nutrient elements, xylem sap, enzymes activities, and hormone concentrations. The uptake and distribution of nanoparticles by the plants were confirmed using transmission electron microscopy (TEM). The SiO2 nanoparticles decreased significantly the plant height, shoot and root biomasses; the SiO2 nanoparticles also affected the contents of Cu, Mg in shoots and Na in roots of transgenic cotton; and SOD activity and IAA concentration were significantly influenced by SiO2 nanoparticles. In addition, SiO2 nanoparticles were present in the xylem sap and roots as examined by TEM showing that the SiO2 nanoparticles were transported from roots to shoots via xylem sap. This is the first report of the transportation of SiO2 nanoparticles via xylem sap within Bt-transgenic cotton. This study provides direct evidence for the bioaccumulation of SiO2 nanoparticles in plants, which shows the potential risks of SiO2 nanoparticles impact on food crops and human health.
López, Rosana; López de Heredia, Unai; Collada, Carmen; Cano, Francisco Javier; Emerson, Brent C; Cochard, Hervé; Gil, Luis
2013-06-01
It is widely accepted that hydraulic failure due to xylem embolism is a key factor contributing to drought-induced mortality in trees. In the present study, an attempt is made to disentangle phenotypic plasticity from genetic variation in hydraulic traits across the entire distribution area of a tree species to detect adaptation to local environments. A series of traits related to hydraulics (vulnerability to cavitation and hydraulic conductivity in branches), growth performance and leaf mass per area were assessed in eight Pinus canariensis populations growing in two common gardens under contrasting environments. In addition, the neutral genetic variability (FST) and the genetic differentiation of phenotypic variation (QST) were compared in order to identify the evolutionary forces acting on these traits. The variability for hydraulic traits was largely due to phenotypic plasticity. Nevertheless, the vulnerability to cavitation displayed a significant genetic variability (approx. 5 % of the explained variation), and a significant genetic × environment interaction (between 5 and 19 % of the explained variation). The strong correlation between vulnerability to cavitation and survival in the xeric common garden (r = -0·81; P < 0·05) suggests a role for the former in the adaptation to xeric environments. Populations from drier sites and higher temperature seasonality were less vulnerable to cavitation than those growing at mesic sites. No trade-off between xylem safety and efficiency was detected. QST of parameters of the vulnerability curve (0·365 for P50 and the slope of the vulnerability curve and 0·452 for P88) differed substantially from FST (0·091), indicating divergent selection. In contrast, genetic drift alone was found to be sufficient to explain patterns of differentiation for xylem efficiency and growth. The ability of P. canariensis to inhabit a wide range of ecosystems seemed to be associated with high phenotypic plasticity and some degree of local adaptations of xylem and leaf traits. Resistance to cavitation conferred adaptive potential for this species to adapt successfully to xeric conditions.
Does short-term potassium fertilization improve recovery from drought stress in laurel?
Oddo, Elisabetta; Inzerillo, Simone; Grisafi, Francesca; Sajeva, Maurizio; Salleo, Sebastiano; Nardini, Andrea
2014-08-01
Xylem hydraulic conductance varies in response to changes in sap solute content, and in particular of potassium (K(+)) ion concentration. This phenomenon, known as the 'ionic effect', is enhanced in embolized stems, where it can compensate for cavitation-induced loss of hydraulic conductance. Previous studies have shown that in well-watered laurel plants (Laurus nobilis L.), potassium concentration of the xylem sap and plant hydraulic conductance increased 24 h after fertilization with KCl. The aim of this work was to test whether water-stressed laurel plants, grown under low potassium availability, could recover earlier from stress when irrigated with a KCl solution instead of potassium-free water. Two-year-old potted laurel seedlings were subjected to water stress by suspending irrigation until leaf conductance to water vapour (g(L)) dropped to ∼30% of its initial value and leaf water potential (ψ(L)) reached the turgor loss point (ψ(TLP)). Plants were then irrigated either with water or with 25 mM KCl and monitored for water status, gas exchange and plant hydraulics recovery at 3, 6 and 24 h after irrigation. No significant differences were found between the two experimental groups in terms of ψ(L), g(L), plant transpiration, plant hydraulic conductance or leaf-specific shoot hydraulic conductivity. Analysis of xylem sap potassium concentration showed that there were no significant differences between treatments, and potassium levels were similar to those of potassium-starved but well-watered plants. In conclusion, potassium uptake from the soil solution and/or potassium release to the xylem appeared to be impaired in water-stressed plants, at least up to 24 h after relief from water stress, so that fertilization after the onset of stress did not result in any short-term advantage for recovery from drought. © The Author 2014. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.
Iakimova, Elena T; Woltering, Ernst J
2017-04-01
Physiological and molecular studies support the view that xylogenesis can largely be determined as a specific form of vacuolar programmed cell death (PCD). The studies in xylogenic zinnia cell culture have led to many breakthroughs in xylogenesis research and provided a background for investigations in other experimental models in vitro and in planta . This review discusses the most essential earlier and recent findings on the regulation of xylem elements differentiation and PCD in zinnia and other xylogenic systems. Xylogenesis (the formation of water conducting vascular tissue) is a paradigm of plant developmental PCD. The xylem vessels are composed of fused tracheary elements (TEs)-dead, hollow cells with patterned lignified secondary cell walls. They result from the differentiation of the procambium and cambium cells and undergo cell death to become functional post-mortem. The TE differentiation proceeds through a well-coordinated sequence of events in which differentiation and the programmed cellular demise are intimately connected. For years a classical experimental model for studies on xylogenesis was the xylogenic zinnia (Zinnia elegans) cell culture derived from leaf mesophyll cells that, upon induction by cytokinin and auxin, transdifferentiate into TEs. This cell system has been proven very efficient for investigations on the regulatory components of xylem differentiation which has led to many discoveries on the mechanisms of xylogenesis. The knowledge gained from this system has potentiated studies in other xylogenic cultures in vitro and in planta. The present review summarises the previous and latest findings on the hormonal and biochemical signalling, metabolic pathways and molecular and gene determinants underlying the regulation of xylem vessels differentiation in zinnia cell culture. Highlighted are breakthroughs achieved through the use of xylogenic systems from other species and newly introduced tools and analytical approaches to study the processes. The mutual dependence between PCD signalling and the differentiation cascade in the program of TE development is discussed.
Scoffoni, Christine; Vuong, Christine; Diep, Steven; Cochard, Hervé; Sack, Lawren
2014-01-01
Leaf shrinkage with dehydration has attracted attention for over 100 years, especially as it becomes visibly extreme during drought. However, little has been known of its correlation with physiology. Computer simulations of the leaf hydraulic system showed that a reduction of hydraulic conductance of the mesophyll pathways outside the xylem would cause a strong decline of leaf hydraulic conductance (Kleaf). For 14 diverse species, we tested the hypothesis that shrinkage during dehydration (i.e. in whole leaf, cell and airspace thickness, and leaf area) is associated with reduction in Kleaf at declining leaf water potential (Ψleaf). We tested hypotheses for the linkage of leaf shrinkage with structural and physiological water relations parameters, including modulus of elasticity, osmotic pressure at full turgor, turgor loss point (TLP), and cuticular conductance. Species originating from moist habitats showed substantial shrinkage during dehydration before reaching TLP, in contrast with species originating from dry habitats. Across species, the decline of Kleaf with mild dehydration (i.e. the initial slope of the Kleaf versus Ψleaf curve) correlated with the decline of leaf thickness (the slope of the leaf thickness versus Ψleaf curve), as expected based on predictions from computer simulations. Leaf thickness shrinkage before TLP correlated across species with lower modulus of elasticity and with less negative osmotic pressure at full turgor, as did leaf area shrinkage between full turgor and oven desiccation. These findings point to a role for leaf shrinkage in hydraulic decline during mild dehydration, with potential impacts on drought adaptation for cells and leaves, influencing plant ecological distributions. PMID:24306532
Leaf shrinkage with dehydration: coordination with hydraulic vulnerability and drought tolerance.
Scoffoni, Christine; Vuong, Christine; Diep, Steven; Cochard, Hervé; Sack, Lawren
2014-04-01
Leaf shrinkage with dehydration has attracted attention for over 100 years, especially as it becomes visibly extreme during drought. However, little has been known of its correlation with physiology. Computer simulations of the leaf hydraulic system showed that a reduction of hydraulic conductance of the mesophyll pathways outside the xylem would cause a strong decline of leaf hydraulic conductance (K(leaf)). For 14 diverse species, we tested the hypothesis that shrinkage during dehydration (i.e. in whole leaf, cell and airspace thickness, and leaf area) is associated with reduction in K(leaf) at declining leaf water potential (Ψ(leaf)). We tested hypotheses for the linkage of leaf shrinkage with structural and physiological water relations parameters, including modulus of elasticity, osmotic pressure at full turgor, turgor loss point (TLP), and cuticular conductance. Species originating from moist habitats showed substantial shrinkage during dehydration before reaching TLP, in contrast with species originating from dry habitats. Across species, the decline of K(leaf) with mild dehydration (i.e. the initial slope of the K(leaf) versus Ψ(leaf) curve) correlated with the decline of leaf thickness (the slope of the leaf thickness versus Ψ(leaf) curve), as expected based on predictions from computer simulations. Leaf thickness shrinkage before TLP correlated across species with lower modulus of elasticity and with less negative osmotic pressure at full turgor, as did leaf area shrinkage between full turgor and oven desiccation. These findings point to a role for leaf shrinkage in hydraulic decline during mild dehydration, with potential impacts on drought adaptation for cells and leaves, influencing plant ecological distributions.
An annual pattern of native embolism in upper branches of four tall conifer species.
McCulloh, Katherine A; Johnson, Daniel M; Meinzer, Frederick C; Lachenbruch, Barbara
2011-06-01
The Pacific Northwest of North America experiences relatively mild winters and dry summers. For the tall coniferous trees that grow in this region, we predicted that loss in the hydraulic conductivity of uppermost branches would be avoided because of difficulty reversing accumulated emboli in xylem that is always under negative pressure. To test this hypothesis, we measured native percent loss in hydraulic conductivity (PLC; the decrease of in situ hydraulic conductivity relative to the maximum) monthly throughout 2009 in branches at the tops (∼50 m) of four species in an old growth forest in southern Washington. Contrary to our prediction, freeze-thaw cycles resulted in considerable native PLC. Branches showed hydraulic recovery in the spring and after a moderate increase in native embolism that was observed after an unusually hot period in August. The September recovery occurred despite decreases in the leaf and stem water potentials compared to August values. Recoveries in branches of these trees could not have occurred by raising the water potential enough to dissolve bubbles simply by transporting water from roots and must have occurred either through water absorption through needles and/or refilling under negative pressure. Excluding the August value, native embolism values correlated strongly with air temperature of the preceding 10 d. For three species, we found that branches with lower wood density had higher specific conductivity, but not greater native PLC than branches with higher wood density, which calls into question whether there is any hydraulic benefit to higher wood density in small branches in those species.
Ceballos-Laita, Laura; Gutierrez-Carbonell, Elain; Takahashi, Daisuke; Abadía, Anunciación; Uemura, Matsuo; Abadía, Javier; López-Millán, Ana Flor
2018-01-06
The aim of this work was to study the effects of Fe and Mn deficiencies on the xylem sap proteome of tomato using a shotgun proteomic approach, with the final goal of elucidating plant response mechanisms to these stresses. This approach yielded 643 proteins reliably identified and quantified with 70% of them predicted as secretory. Iron and Mn deficiencies caused statistically significant and biologically relevant abundance changes in 119 and 118 xylem sap proteins, respectively. In both deficiencies, metabolic pathways most affected were protein metabolism, stress/oxidoreductases and cell wall modifications. First, results suggest that Fe deficiency elicited more stress responses than Mn deficiency, based on the changes in oxidative and proteolytic enzymes. Second, both nutrient deficiencies affect the secondary cell wall metabolism, with changes in Fe deficiency occurring via peroxidase activity, and in Mn deficiency involving peroxidase, Cu-oxidase and fasciclin-like arabinogalactan proteins. Third, the primary cell wall metabolism was affected by both nutrient deficiencies, with changes following opposite directions as judged from the abundances of several glycoside-hydrolases with endo-glycolytic activities and pectin esterases. Fourth, signaling pathways via xylem involving CLE and/or lipids as well as changes in phosphorylation and N-glycosylation also play a role in the responses to these stresses. Biological significance In spite of being essential for the delivery of nutrients to the shoots, our knowledge of xylem responses to nutrient deficiencies is very limited. The present work applies a shotgun proteomic approach to unravel the effects of Fe and Mn deficiencies on the xylem sap proteome. Overall, Fe deficiency seems to elicit more stress in the xylem sap proteome than Mn deficiency, based on the changes measured in proteolytic and oxido-reductase proteins, whereas both nutrients exert modifications in the composition of the primary and secondary cell wall. Cell wall modifications could affect the mechanical and permeability properties of the xylem sap vessels, and therefore ultimately affect solute transport and distribution to the leaves. Results also suggest that signaling cascades involving lipid and peptides might play a role in nutrient stress signaling and pinpoint interesting candidates for future studies. Finally, both nutrient deficiencies seem to affect phosphorylation and glycosylation processes, again following an opposite pattern. Copyright © 2017 Elsevier B.V. All rights reserved.
How do sharpshooter leafhoppers feed and survive on nutritionally depauperate xylem fluid?
USDA-ARS?s Scientific Manuscript database
Sharpshooters (Cicadellidae: Cicadellinae) are large, tropical and semi-tropical leafhoppers that are unique among all non-sessile hemipterans in ingesting primarily from xylem vessels. This presentation will summarize research on behavioral and physiological adaptations that permit sharpshooters ...
Use of gold nanoparticles to detect water uptake in vascular plants.
Hwang, Bae Geun; Ahn, Sungsook; Lee, Sang Joon
2014-01-01
Direct visualization of water-conducting pathways and sap flows in xylem vessels is important for understanding the physiology of vascular plants and their sap ascent. Gold nanoparticles (AuNPs) combined with synchrotron X-ray imaging technique is a new promising tool for investigating plant hydraulics in opaque xylem vessels of vascular plants. However, in practical applications of AuNPs for real-time quantitative visualization of sap flows, their interaction with a vascular network needs to be verified in advance. In this study, the effect of AuNPs on the water-refilling function of xylem vessels is experimentally investigated with three monocot species. Discrepancy in the water uptakes starts to appear at about 20 min to 40 min after the supply of AuNP solution to the test plant by the possible gradual accumulation of AuNPs on the internal structures of vasculature. However conclusively, it is observed that the water-refilling speeds in individual xylem vessels are virtually unaffected by hydrophilically surface-modified AuNPs (diameter ∼20 nm). Therefore, the AuNPs can be effectively used as flow tracers in the xylem vessels in the first 20∼30 min without any physiological barrier. As a result, AuNPs are found to be useful for visualizing various fluid dynamic phenomena occurring in vascular plants.
Metabolic Responses of Poplar to Apripona germari (Hope) as Revealed by Metabolite Profiling
Wang, Lijuan; Qu, Liangjian; Zhang, Liwei; Hu, Jianjun; Tang, Fang; Lu, Mengzhu
2016-01-01
Plants have developed biochemical responses to adapt to biotic stress. To characterize the resistance mechanisms in poplar tree against Apripona germari, comprehensive metabolomic changes of poplar bark and xylem in response to A. germari infection were examined by gas chromatography time-of-flight mass spectrometry (GC–TOF/MS). It was found that, four days after feeding (stage I), A. germari infection brought about changes in various metabolites, such as phenolics, amino acids and sugars in both bark and xylem. Quinic acid, epicatechin, epigallocatechin and salicin might play a role in resistance response in bark, while coniferyl alcohol, ferulic acid and salicin contribute resistance in xylem. At feeding stages II when the larvae fed for more than one month, fewer defensive metabolites were induced, but levels of many intermediates of glycolysis and the tricarboxylic acid (TCA) cycle were reduced, especially in xylem. These results suggested that the defense strategies against A. germari might depend mainly on the early defense responses in poplar. In addition, it was found that bark and xylem in infected trees accumulated higher levels of salicylic acid and 4-aminobutyric acid, respectively, these tissues displaying a direct and systemic reaction against A. germari. However, the actual role of the two metabolites in A. germari-induced defense in poplar requires further investigation. PMID:27331808
Cellulase and cell differentiation in Acer pseudoplatanus.
Sheldrake, A R
1970-06-01
Homogenates of differentiating xylem and phloem tissue have higher cellulase activities than cambial samples; the highest activity is always found in phloem. Callus tissue, in which no vascular differentiation occurs, contains only low cellulase activity. The results suggest that cellulase is involved in vascular differentiation. Different pH optima of cellulase activity were found: in cambium, xylem and phloem tissue, cellulase activity with an optimum at about pH 5.9 is predominantly membrane-bound; it is sedimentable at 100,000 g and releasable by Triton X-100. The same may be true of activity with an optimum at pH 5.3. Phloem tissue also contains a soluble, cytoplasmic cellulase of high activity at pH 7.1, and xylem tissue contains cytoplasmic cellulase with an optimum at pH 6.5. Low cellulase activity with a pH optimum similar to that of xylem homogenates was found in xylem sap. Cellulase activity in abscission zones increases greatly just before leaf abscission. Abscission zone cellulase has two pH optima, et 5.3 and 5.9; both activities are increased by Triton treatment of homogenates. The possible existence of several different cellulases forming part of a cellulase complex, and the rôle of the enzymes in hydrolysing wall material during cell differentiation are discussed.
Air method measurements of apple vessel length distributions with improved apparatus and theory
Shabtal Cohen; John Bennink; Mel Tyree
2003-01-01
Studies showing that rootstock dwarfing potential is related to plant hydraulic conductance led to the hypothesis that xylem properties are also related. Vessel length distribution and other properties of apple wood from a series of varieties were measured using the 'air method' in order to test this hypothesis. Apparatus was built to measure and monitor...
USDA-ARS?s Scientific Manuscript database
Low water potentials in xylem can result in damaging levels of cavitation, yet little is understood about which hydraulic traits have most influence in delaying the onset of hydraulic dysfunction during periods of drought. We examined three traits contributing to longer desiccation times in excised ...
New star on the stage: amount of ray parenchyma in tree rings shows a link to climate.
Olano, José Miguel; Arzac, Alberto; García-Cervigón, Ana I; von Arx, Georg; Rozas, Vicente
2013-04-01
Tree-ring anatomy reflects the year-by-year impact of environmental factors on tree growth. Up to now, research in this field has mainly focused on the hydraulic architecture, with ray parenchyma neglected despite the growing recognition of its relevance for xylem function. Our aim was to address this gap by exploring the potential of the annual patterns of xylem parenchyma as a climate proxy. We constructed ring-width and ray-parenchyma chronologies from 1965 to 2004 for 20 Juniperus thurifera trees growing in a Mediterranean continental climate. Chronologies were related to climate records by means of correlation, multiple regression and partial correlation analyses. Ray parenchyma responded to climatic conditions at critical stages during the xylogenetic process; namely, at the end of the previous year's xylogenesis (October) and at the onset of earlywood (May) and latewood formation (August). Ray parenchyma-based chronologies have potential to complement ring-width chronologies as a tool for climate reconstructions. Furthermore, medium- and low-frequency signals in the variation of ray parenchyma may improve our understanding of how trees respond to environmental fluctuations and to global change. © 2013 The Authors. New Phytologist © 2013 New Phytologist Trust.
Pavlik, B M
2001-08-01
Measurements of xylem water potential, leaf conductance, and leaf pressure-volume characteristics on the geothermal endemic Dichanthelium lanuginosum var. thermale (DILA) were used to delineate operational ranges during wet and dry years and among several microsites at Little Geysers, Sonoma County, California, USA. Plants seldom experienced water potentials more negative that -1.5 MPa. Other nongeothermal, widespread species experienced the lower water potentials typical of chaparral and woodland plants. DILA was able to effectively utilize geothermal water while the widespread species could not and was able to keep stomata open during most of the year. There was evidence to suggest that DILA had some ability to acclimate with significant shifts in Pio and psio during the dry 1994 summer, especially in the upland microhabitat. Nevertheless, minimum leaf turgor values in the upland came very close to, or dropped below, the 0.2-0.3 MPa threshold thought necessary to maintain stomatal opening and photosynthesis. DILA thus depends upon the unique water status of fumarole soils in the vicinity of the Little Geysers to persist in an otherwise lethal regional mosaic of climate, soil, and vegetation. The physiological data were used to derive reference ranges for subsequent monitoring of DILA at Little Geysers. Such ranges are required to determine the future impact, if any, of geothermal development on the persistence of this rare grass and its complex ecosystem.
Holmlund, Helen I; Lekson, Victoria M; Gillespie, Breahna M; Nakamatsu, Nicole A; Burns, Amanda M; Sauer, Kaitlyn E; Pittermann, Jarmila; Davis, Stephen D
2016-09-01
California experienced severe drought between 2012 and 2016. During this period, we compared seasonal changes in tissue-water relations among eight fern species in the Santa Monica Mountains of southern California to elucidate differential mechanisms of drought survival and physiological performance during extreme water deficits. We monitored seasonal changes in water potential (Ψmd) and dark-adapted chlorophyll fluorescence (Fv/Fm), assessed tissue-water relations including osmotic potential at saturation and the turgor loss point (Ψπ, sat and Ψπ, tlp), and measured, for two evergreen species, xylem-specific and leaf-specific hydraulic conductivity (Ks and Kl) and vulnerability of stem xylem to water stress-induced embolism (water potential at 50% loss hydraulic conductivity, Ψ50). Species grew in either riparian or chaparral understory. The five chaparral species had a wider range of seasonal water potentials, root depths, and frond phenological traits, including one evergreen, two summer-deciduous, and two desiccation-tolerant (resurrection) species. Evergreen species were especially diverse, with an evergreen riparian species maintaining seasonal water potentials above -1.3 MPa, while an evergreen chaparral species had seasonal water potentials below -8 MPa. In those two species the Ψ50 values were -2.5 MPa and -4.3 MPa, respectively. Observed differences in physiological performance among eight fern species reflected niche partitioning in water utilization and habitat preference associated with distinct phenological traits. We predict differential survival among fern species as future drought events in California intensify, with desiccation-tolerant resurrection ferns being the most resistant. © 2016 Botanical Society of America.
Hacke, Uwe G; Venturas, Martin D; MacKinnon, Evan D; Jacobsen, Anna L; Sperry, John S; Pratt, R Brandon
2015-01-01
The standard centrifuge method has been frequently used to measure vulnerability to xylem cavitation. This method has recently been questioned. It was hypothesized that open vessels lead to exponential vulnerability curves, which were thought to be indicative of measurement artifact. We tested this hypothesis in stems of olive (Olea europea) because its long vessels were recently claimed to produce a centrifuge artifact. We evaluated three predictions that followed from the open vessel artifact hypothesis: shorter stems, with more open vessels, would be more vulnerable than longer stems; standard centrifuge-based curves would be more vulnerable than dehydration-based curves; and open vessels would cause an exponential shape of centrifuge-based curves. Experimental evidence did not support these predictions. Centrifuge curves did not vary when the proportion of open vessels was altered. Centrifuge and dehydration curves were similar. At highly negative xylem pressure, centrifuge-based curves slightly overestimated vulnerability compared to the dehydration curve. This divergence was eliminated by centrifuging each stem only once. The standard centrifuge method produced accurate curves of samples containing open vessels, supporting the validity of this technique and confirming its utility in understanding plant hydraulics. Seven recommendations for avoiding artefacts and standardizing vulnerability curve methodology are provided. © 2014 The Authors. New Phytologist © 2014 New Phytologist Trust.
Mayr, Stefan; Sperry, John S
2010-03-01
*The 'thaw-expansion hypothesis' postulates that xylem embolism is caused by the formation of gas bubbles on freezing and their expansion on thawing. We evaluated the hypothesis using centrifuge experiments and ultrasonic emission monitoring in Pinus contorta. *Stem samples were exposed to freeze-thaw cycles at varying xylem pressure (P) in a centrifuge before the percentage loss of hydraulic conductivity (PLC) was measured. Ultrasonic acoustic emissions were registered on samples exposed to freeze-thaw cycles in a temperature chamber. *Freeze-thaw exposure of samples spun at -3 MPa induced a PLC of 32% (one frost cycle) and 50% (two cycles). An increase in P to -0.5 MPa during freezing had no PLC effect, whereas increased P during thaw lowered PLC to 7%. Ultrasonic acoustic emissions were observed during freezing and thawing at -3 MPa, but not in air-dried or water-saturated samples. A decrease in minimum temperature caused additional ultrasonic acoustic emissions, but had no effect on PLC. *The centrifuge experiments indicate that the 'thaw-expansion hypothesis' correctly describes the embolization process. Possible explanations for the increase in PLC on repeated frost cycles and for the ultrasonic acoustic emissions observed during freezing and with decreasing ice temperature are discussed.
The dynamics of gas bubbles in conduits of vascular plants and implications for embolism repair.
Konrad, W; Roth-Nebelsick, A
2003-09-07
Pressure-induced tensions in the xylem, the water conducting tissue of vascular plants, can lead to embolism in the water-conducting cells. The details and mechanisms of embolism repair in vascular plants are still not well understood. In particular, experimental results which indicate that embolism repair may occur during xylem tension cause great problems with respect to current paradigms of plant water transport. The present paper deals with a theoretical analysis of interfacial effects at the pits (pores in the conduit walls), because it was suggested that gas-water interfaces at the pit pores may be involved in the repair process by hydraulically isolating the embolized conduit. The temporal behaviour of bubbles at the pit pores was especially studied since the question of whether these pit bubbles are able to persist is of crucial importance for the suggested mechanism to work. The results indicate that (1) the physical preconditions which are necessary for the suggested mechanism appear to be satisfied, (2) pit bubbles can achieve temporal stability and therefore persist and (3) dissolving of bubbles in the conduit lumen may lead to the final breakdown of the hydraulic isolation. The whole process is, however, complex and strongly dependent on the detailed anatomy of the pit and the contact angle.
DELZON, SYLVAIN; DOUTHE, CYRIL; SALA, ANNA; COCHARD, HERVE
2010-01-01
Resistance to water-stress induced cavitation is an important indicator of drought tolerance in woody species and is known to be intimately linked to the anatomy of the xylem. However, the actual mechanical properties of the pit membrane are not well known and the exact mode of air-seeding by which cavitation occurs is still uncertain. We examined the relationship between cavitation resistance and bordered pit structure and function in 40 coniferous species. Xylem pressure inducing 50% loss of hydraulic conductance (P50, a proxy for cavitation resistance) varied widely among species, from −2.9 to −11.3 MPa. The valve effect of the pit membrane, measured as a function of margo flexibility and torus overlap, explained more variation in cavitation-resistance than simple anatomical traits such as pit membrane, pit aperture or torus size. Highly cavitation resistant species exhibited both a high flexibility of the margo and a large overlap between the torus and the pit aperture, allowing the torus to tightly seal the pit aperture. Our results support the hypothesis of seal capillary-seeding as the most likely mode of air-seeding, and suggest that the adhesion of the torus to the pit border may be the main determinant of cavitation resistance in conifers. PMID:20636490
Delzon, Sylvain; Douthe, Cyril; Sala, Anna; Cochard, Herve
2010-12-01
Resistance to water-stress induced cavitation is an important indicator of drought tolerance in woody species and is known to be intimately linked to the anatomy of the xylem. However, the actual mechanical properties of the pit membrane are not well known and the exact mode of air-seeding by which cavitation occurs is still uncertain. We examined the relationship between cavitation resistance and bordered pit structure and function in 40 coniferous species. Xylem pressure inducing 50% loss of hydraulic conductance (P(50), a proxy for cavitation resistance) varied widely among species, from -2.9 to -11.3 MPa. The valve effect of the pit membrane, measured as a function of margo flexibility and torus overlap, explained more variation in cavitation-resistance than simple anatomical traits such as pit membrane, pit aperture or torus size. Highly cavitation resistant species exhibited both a high flexibility of the margo and a large overlap between the torus and the pit aperture, allowing the torus to tightly seal the pit aperture. Our results support the hypothesis of seal capillary-seeding as the most likely mode of air-seeding, and suggest that the adhesion of the torus to the pit border may be the main determinant of cavitation resistance in conifers. © 2010 Blackwell Publishing Ltd.
Frost fatigue and spring recovery of xylem vessels in three diffuse-porous trees in situ.
Christensen-Dalsgaard, Karen K; Tyree, Melvin T
2014-05-01
Frost has been shown to cause frost fatigue (reduced cavitation resistance) in branch segments in the lab. Here, we studied the change in cavitation resistance and percent loss of conductivity (PLC) from fall to spring over 2 consecutive years in three diffuse-porous species in situ. We used the cavitron technique to measure P25 , P50 and P90 (the xylem pressure causing a 25, 50 and 90% conductivity loss) and PLC and stained functioning vessels. Cavitation resistance was reduced by 64-87% (in terms of P50 ), depending on the species and year. P25 was impacted the most and P90 the least, changing the vulnerability curves from s- to r-shaped over the winter in all three species. The branches suffered an almost complete loss of conductivity, but frost fatigue did not necessarily occur concurrently with increases in PLC. In two species, there was a trade-off between conduit size and vulnerability. Spring recovery occurred by growth of new vessels, and in two species by partial refilling of embolized conduits. Although newly grown and functioning conduits appeared more vulnerable to cavitation than year-old vessels, cavitation resistance generally improved in spring, suggesting other mechanisms for partial frost fatigue repair. © 2013 John Wiley & Sons Ltd.
Pita, Pilar; Rodríguez-Calcerrada, Jesús; Medel, David; Gil, Luis
2018-02-01
Dutch elm disease (DED) is a vascular disease that has killed over 1 billion elm trees. The pathogen spreads throughout the xylem network triggering vessel blockage, which results in water stress, tissue dehydration and extensive leaf wilting in susceptible genotypes. We investigated the differences between four Ulmus minor Mill. clones of contrasting susceptibility to Ophiostoma novo-ulmi Brasier regarding morphological, anatomical and physiological traits affecting water transport, in order to gain a better understanding of the mechanisms underlying DED susceptibility. We analyzed the differential response to water shortage and increased air vapor pressure deficit (VPD) to investigate whether resistance to water stress might be related to DED tolerance. Sixteen plants per clone, aged 2 years, were grown inside a greenhouse under differential watering. Stomatal conductance was measured under ambient and increased VPD. Growth, bark water content and stem hydraulic and anatomical parameters were measured 22 days after starting differential watering. Vessel lumen area, lumen fraction and hydraulic conductance were highest in susceptible clones. Stomatal conductance was lowest under low VPD and decreased faster under increased VPD in resistant clones. We found a negative relationship between the decrease in stomatal conductance at increased VPD and specific hydraulic conductance, revealing a narrower hydraulic margin for sustaining transpiration in resistant clones. The effect of water shortage was greater on radial stem growth than on leaf area, which could be explained through an extensive use of capacitance water to buffer xylem water potential. Water shortage reduced stomatal conductance and vessel lumen area. Bark water content under conditions of water shortage only decreased in susceptible clones. Higher hydraulic constraints to sap flow in resistant clones may determine higher stomatal sensitivity to VPD and so contribute to DED resistance by limiting pathogen expansion and reducing water loss and metabolic impairment in cells involved in fighting against infection. © The Author 2017. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.
Volaire, Florence; Lens, Frederic; Cochard, Hervé; Xu, Hueng; Chacon-Doria, Larissa; Bristiel, Pauline; Balachowski, Jennifer; Rowe, Nick; Violle, Cyrille; Picon-Cochard, Catherine
2018-05-17
More intense droughts under climate change threaten species resilience. Hydraulic strategies determine drought survival in woody plants but have been hardly studied in herbaceous species. We explored the intraspecific variability of hydraulic and morphological traits as indicators of dehydration tolerance in a perennial grass, cocksfoot (Dactylis glomerata), which has a large biogeographical distribution in Europe. Twelve populations of cocksfoot originating from Mediterranean, Temperate and Northern European areas were grown in a controlled environment in pots. Dehydration tolerance, leaf and stem anatomical traits and xylem pressure associated with 88 or 50 % loss of xylem conductance (P88, P50) were measured. Across the 12 populations of cocksfoot, P50 ranged from -3.06 to - 6.36 MPa, while P88 ranged from -5.06 to -11.6 MPa. This large intraspecific variability of embolism thresholds corresponded with the biogeographical distribution and some key traits of the populations. In particular, P88 was correlated with dehydration tolerance (r = -0.79). The dehydration-sensitive Temperate populations exhibited the highest P88 (-6.1 MPa). The most dehydration-tolerant Mediterranean populations had the greatest leaf dry matter content and leaf fracture toughness, and the lowest P88 (-10.4 MPa). The Northern populations displayed intermediate trait values, potentially attributable to frost resistance. The thickness of metaxylem vessel walls in stems was highly correlated with P50 (r = -0.92), but no trade-off with stem lignification was observed. The relevance of the linkage between hydraulic and stomatal traits is discussed for drought survival in perennial grasses. Compared with woody species, the large intraspecific variability in dehydration tolerance and embolism resistance within cocksfoot has consequences for its sensitivity to climate change. To better understand adaptive strategies of herbaceous species to increasing drought and frost requires further exploration of the role of hydraulic and mechanical traits using a larger inter- and intraspecific range of species.
Nilsson, Robert; Bernfur, Katja; Gustavsson, Niklas; Bygdell, Joakim; Wingsle, Gunnar; Larsson, Christer
2010-02-01
By exploiting the abundant tissues available from Populus trees, 3-4 m high, we have been able to isolate plasma membranes of high purity from leaves, xylem, and cambium/phloem at a time (4 weeks after bud break) when photosynthesis in the leaves and wood formation in the xylem should have reached a steady state. More than 40% of the 956 proteins identified were found in the plasma membranes of all three tissues and may be classified as "housekeeping" proteins, a typical example being P-type H(+)-ATPases. Among the 213 proteins predicted to be integral membrane proteins, transporters constitute the largest class (41%) followed by receptors (14%) and proteins involved in cell wall and carbohydrate metabolism (8%) and membrane trafficking (8%). ATP-binding cassette transporters (all members of subfamilies B, C, and G) and receptor-like kinases (four subfamilies) were two of the largest protein families found, and the members of these two families showed pronounced tissue distribution. Leaf plasma membranes were characterized by a very high proportion of transporters, constituting almost half of the integral proteins. Proteins involved in cell wall synthesis (such as cellulose and sucrose synthases) and membrane trafficking were most abundant in xylem plasma membranes in agreement with the role of the xylem in wood formation. Twenty-five integral proteins and 83 soluble proteins were exclusively found in xylem plasma membranes, which identifies new candidates associated with cell wall synthesis and wood formation. Among the proteins uniquely found in xylem plasma membranes were most of the enzymes involved in lignin biosynthesis, which suggests that they may exist as a complex linked to the plasma membrane.
Quantitative Wood Anatomy-Practical Guidelines.
von Arx, Georg; Crivellaro, Alan; Prendin, Angela L; Čufar, Katarina; Carrer, Marco
2016-01-01
Quantitative wood anatomy analyzes the variability of xylem anatomical features in trees, shrubs, and herbaceous species to address research questions related to plant functioning, growth, and environment. Among the more frequently considered anatomical features are lumen dimensions and wall thickness of conducting cells, fibers, and several ray properties. The structural properties of each xylem anatomical feature are mostly fixed once they are formed, and define to a large extent its functionality, including transport and storage of water, nutrients, sugars, and hormones, and providing mechanical support. The anatomical features can often be localized within an annual growth ring, which allows to establish intra-annual past and present structure-function relationships and its sensitivity to environmental variability. However, there are many methodological challenges to handle when aiming at producing (large) data sets of xylem anatomical data. Here we describe the different steps from wood sample collection to xylem anatomical data, provide guidance and identify pitfalls, and present different image-analysis tools for the quantification of anatomical features, in particular conducting cells. We show that each data production step from sample collection in the field, microslide preparation in the lab, image capturing through an optical microscope and image analysis with specific tools can readily introduce measurement errors between 5 and 30% and more, whereby the magnitude usually increases the smaller the anatomical features. Such measurement errors-if not avoided or corrected-may make it impossible to extract meaningful xylem anatomical data in light of the rather small range of variability in many anatomical features as observed, for example, within time series of individual plants. Following a rigid protocol and quality control as proposed in this paper is thus mandatory to use quantitative data of xylem anatomical features as a powerful source for many research topics.
Ren, Ping; Rossi, Sergio; Gricar, Jozica; Liang, Eryuan; Cufar, Katarina
2015-01-01
Background and Aims A series of studies have shown that temperature triggers the onset of xylogenesis of trees after winter dormancy. However, little is known about whether and how moisture availability influences xylogenesis in spring in drought-prone areas. Methods Xylogenesis was monitored in five mature Qilian junipers (Juniperus przewalskii) by microcore sampling from 2009 to 2011 in a semi-arid area of the north-eastern Tibetan Plateau. A simple physical model of xylem cell production was developed and its sensitivity was analysed. The relationship between climate and growth was then evaluated, using weekly wood production data and climatic data from the study site. Key Results Delayed onset of xylogenesis in 2010 corresponded to a negative standardized precipitation evapotranspiration index (SPEI) value and a continuous period without rainfall in early May. The main period of wood formation was in June and July, and drier conditions from May to July led to a smaller number of xylem cells. Dry conditions in July could cause early cessation of xylem differentiation. The final number of xylem cells was mainly determined by the average production rate rather than the duration of new cell production. Xylem growth showed a positive and significant response to precipitation, but not to temperature. Conclusions Precipitation in late spring and summer can play a critical role in the onset of xylogenesis and xylem cell production. The delay in the initiation of xylogenesis under extremely dry conditions seems to be a stress-avoidance strategy against hydraulic failure. These findings could thus demonstrate an evolutionary adaptation of Qilian juniper to the extremely dry conditions of the north-eastern Tibetan Plateau. PMID:25725006
Turtola, S; Manninen, A M; Holopainen, J K; Levula, T; Raitio, H; Kainulainen, P
2002-01-01
Secondary compounds are known to be associated with the resistance of conifer xylem against insects and fungi. The effects of long-term forest fertilization with nitrogen (N) or with N, calcium (Ca), and phosphorus (P) on secondary compounds in the xylem of 50-yr-old Scots pine (Pinus sylvestris L.) trees were examined. Xylem samples were collected from trees growing in three locations in southern Finland: Vilppula, Padasjoki, and Punkaharju. Forests were fertilized every fifth (Vilppula and Padasjoki) or tenth (Punkaharju) year since the 1950s. We compared concentrations of individual and total monoterpenes and resin acids in the heartwood and sapwood of Scots pine. Terpene emissions were analyzed from the sapwood and total phenolics from the heartwood. Fertilization did not have any significant effect on the concentrations and emissions of xylem monoterpenes. Concentrations of several individual terpenes in sapwood were positively correlated with the corresponding terpene emission. The concentrations of individual resin acids (i.e., abietic and dehydroabietic) decreased significantly in Punkaharju, but increased in the sapwood of N-fertilized trees compared with control ones at Padasjoki and Vilppula. The concentrations of resin acids in the heartwood were not significantly affected by fertilization. Both fertilization treatments decreased the total phenolic concentrations in the heartwood of trees growing in Padasjoki. There was a significant positive correlation between the total phenolics and total resin acid concentration. Overall, resin acids and phenolics seemed be more responsive than monoterpenes to N treatment. These results suggest that forest fertilization might cause slight changes in secondary compound concentrations of xylem, and thus might have significance in the decay resistance of wood.
Quantitative Wood Anatomy—Practical Guidelines
von Arx, Georg; Crivellaro, Alan; Prendin, Angela L.; Čufar, Katarina; Carrer, Marco
2016-01-01
Quantitative wood anatomy analyzes the variability of xylem anatomical features in trees, shrubs, and herbaceous species to address research questions related to plant functioning, growth, and environment. Among the more frequently considered anatomical features are lumen dimensions and wall thickness of conducting cells, fibers, and several ray properties. The structural properties of each xylem anatomical feature are mostly fixed once they are formed, and define to a large extent its functionality, including transport and storage of water, nutrients, sugars, and hormones, and providing mechanical support. The anatomical features can often be localized within an annual growth ring, which allows to establish intra-annual past and present structure-function relationships and its sensitivity to environmental variability. However, there are many methodological challenges to handle when aiming at producing (large) data sets of xylem anatomical data. Here we describe the different steps from wood sample collection to xylem anatomical data, provide guidance and identify pitfalls, and present different image-analysis tools for the quantification of anatomical features, in particular conducting cells. We show that each data production step from sample collection in the field, microslide preparation in the lab, image capturing through an optical microscope and image analysis with specific tools can readily introduce measurement errors between 5 and 30% and more, whereby the magnitude usually increases the smaller the anatomical features. Such measurement errors—if not avoided or corrected—may make it impossible to extract meaningful xylem anatomical data in light of the rather small range of variability in many anatomical features as observed, for example, within time series of individual plants. Following a rigid protocol and quality control as proposed in this paper is thus mandatory to use quantitative data of xylem anatomical features as a powerful source for many research topics. PMID:27375641
De novo transcriptome assemblies of four xylem sap-feeding insects.
Tassone, Erica E; Cowden, Charles C; Castle, S J
2017-03-01
Spittle bugs and sharpshooters are well-known xylem sap-feeding insects and vectors of the phytopathogenic bacterium Xylella fastidiosa (Wells), a causal agent of Pierce's disease of grapevines and other crop diseases. Specialized feeding on nutrient-deficient xylem sap is relatively rare among insect herbivores, and only limited genomic and transcriptomic information has been generated for xylem-sap feeders. To develop a more comprehensive understanding of biochemical adaptations and symbiotic relationships that support survival on a nutritionally austere dietary source, transcriptome assemblies for three sharpshooter species and one spittlebug species were produced. Trinity-based de novo transcriptome assemblies were generated for all four xylem-sap feeders using raw sequencing data originating from whole-insect preps. Total transcripts for each species ranged from 91 384 for Cuerna arida to 106 998 for Homalodisca liturata with transcript totals for Graphocephala atropunctata and the spittlebug Clastoptera arizonana falling in between. The percentage of transcripts comprising complete open reading frames ranged from 60% for H. liturata to 82% for C. arizonana. Bench-marking universal single-copy orthologs analyses for each dataset indicated quality assemblies and a high degree of completeness for all four species. These four transcriptomes represent a significant expansion of data for insect herbivores that feed exclusively on xylem sap, a nutritionally deficient dietary source relative to other plant tissues and fluids. Comparison of transcriptome data with insect herbivores that utilize other dietary sources may illuminate fundamental differences in the biochemistry of dietary specialization. Published by Oxford University Press on behalf of GIGSCI 2017. This work is written by (a) US Government employee(s) and is in the public domain in the US.
Chuche, Julien; Sauvion, Nicolas; Thiéry, Denis
2017-10-01
In phytophagous piercing-sucking insects, salivary sheath-feeding species are often described as xylem- or phloem-sap feeding specialists. Because these two food sources have very different characteristics, two feeding tactics are often associated with this supposed specialization. Studying the feeding behavior of insects provides substantial information on their biology, ecology, and evolution. Furthermore, study of feeding behavior is of primary importance to elucidate the transmission ability of insects that act as vectors of plant pathogens. In this study, we compared the durations of ingestion performed in xylem versus phloem by a leafhopper species, Scaphoideus titanus Ball, 1932. This was done by characterizing and statistically analyzing electrical signals recorded using the electropenetrography technique, derived from the feeding behaviors of males and females. We identified three groups of S. titanus based on their feeding behavior: 1) a group that reached the phloem quickly and probed for a longer time in phloem tissue than the other groups, 2) a group that reached the xylem quickly and probed for a longer time in xylem tissue than the other groups, and 3) a group where individuals did not ingest much sap. In addition, the numbers and durations of waveforms representing ingestion of xylem and phloem saps differed significantly depending on the sex of the leafhopper, indicating that the two sexes exhibit different feeding behaviors. Males had longer phloem ingestion events than did females, which indicates that males are greater phloem feeders than females. These differences are discussed, specifically in relation to hypotheses about evolution of sap feeding and phytoplasma transmission from plant to plant. Copyright © 2017 Elsevier Ltd. All rights reserved.
Are flowers vulnerable to xylem cavitation during drought?
Zhang, Feng-Ping; Brodribb, Timothy J
2017-05-17
Water stress is known to cause xylem cavitation in the leaves, roots and stems of plants, but little is known about the vulnerability of flowers to xylem damage during drought. This is an important gap in our understanding of how and when plants become damaged by water stress. Here we address fundamental questions about if and when flowers suffer cavitation damage, using a new technique of cavitation imaging to resolve the timing of cavitation in water-stressed flower petals compared with neighbouring leaves. Leaves and flowers from a sample of two herbaceous and two woody eudicots were exposed to a severe water stress while the spatial and temporal propagation of embolism through veins was recorded. Although in most cases water potentials inducing 50% embolism of herbaceous flower veins were more negative than neighbouring leaves, there was no significant difference between the average vulnerability of leaves and petals of herbaceous species. In both woody species, petals were more vulnerable to cavitation than leaves, in one case by more than 3 MPa. Early cavitation and subsequent damage of flowers in the two woody species would thus be expected to precede leaf damage during drought. Similar cavitation thresholds of flowers and leaves in the herb sample suggest that cavitation during water shortage in these species will occur simultaneously among aerial tissues. Species-specific differences in the cavitation thresholds of petals provide a new axis of variation that may explain contrasting flowering ecology among plant species. © 2017 The Author(s).
de Freitas, Sergio Tonetto; Shackel, Kenneth A; Mitcham, Elizabeth J
2011-05-01
Calcium (Ca) uptake into fruit and leaves is dependent on xylemic water movement, and hence presumably driven by transpiration and growth. High leaf transpiration is thought to restrict Ca movement to low-transpiring tomato fruit, which may increase fruit susceptibility to the Ca-deficiency disorder, blossom end rot (BER). The objective of this study was to analyse the effect of reduced leaf transpiration in abscisic acid (ABA)-treated plants on fruit and leaf Ca uptake and BER development. Tomato cultivars Ace 55 (Vf) and AB2 were grown in a greenhouse environment under Ca-deficit conditions and plants were treated weekly after pollination with water (control) or 500 mg l(-1) ABA. BER incidence was completely prevented in the ABA-treated plants and reached values of 30-45% in the water-treated controls. ABA-treated plants had higher stem water potential, lower leaf stomatal conductance, and lower whole-plant water loss than water-treated plants. ABA treatment increased total tissue and apoplastic water-soluble Ca concentrations in the fruit, and decreased Ca concentrations in leaves. In ABA-treated plants, fruit had a higher number of Safranin-O-stained xylem vessels at early stages of growth and development. ABA treatment reduced the phloem/xylem ratio of fruit sap uptake. The results indicate that ABA prevents BER development by increasing fruit Ca uptake, possibly by a combination of whole-plant and fruit-specific mechanisms.
The hydraulic architecture of Juniperus communis L. ssp. communis: shrubs and trees compared.
Beikircher, Barbara; Mayr, Stefan
2008-11-01
Juniperus communis ssp. communis can grow like a shrub or it can develop a tree-like habit. In this study, the hydraulic architecture of these contrasting growth forms was compared. We analysed the hydraulic efficiency (leaf-specific conductivity, k(l); specific conductivity, k(s); Huber value, HV) and the vulnerability to cavitation (the water potential corresponding to a 50% loss of conductivity, Psi(50)), as well as anatomical parameters [mean tracheid diameter, d; mean hydraulic diameter, d(h); cell wall reinforcement (t/b)(h)(2)] of shrub shoots, tree stems and tree branches. Shrub shoots were similar to tree branches (especially to lower branches) in growth form and conductivity (k(l) = 1.93 +/- 0.11 m(2) s(-1) MPa(-1) 10(-7), k(s) = 5.71 +/- 0.19 m(2) s(-1) MPa(-1) 10(-4)), but were similar to tree stems in their vulnerability to cavitation (Psi(50) = -5.81 +/- 0.08 MPa). Tree stems showed extraordinarily high k(l) and k(s) values, and HV increased from the base up. Stem xylem was more vulnerable to cavitation than branch xylem, where Psi(50) increased from lower (Psi(50) = -6.44 +/- 0.19 MPa) to upper branches (Psi(50) = -5.98 +/- 0.13 MPa). Conduit diameters were correlated with k(l) and k(s). Data indicate that differences in hydraulic architecture correspond to changes in growth form. In some aspects, the xylem hydraulics of tree-like Juniperus communis differs from that of other coniferous tree species.
Maximum height in a conifer is associated with conflicting requirements for xylem design.
Domec, Jean-Christophe; Lachenbruch, Barbara; Meinzer, Frederick C; Woodruff, David R; Warren, Jeffrey M; McCulloh, Katherine A
2008-08-19
Despite renewed interest in the nature of limitations on maximum tree height, the mechanisms governing ultimate and species-specific height limits are not yet understood, but they likely involve water transport dynamics. Tall trees experience increased risk of xylem embolism from air-seeding because tension in their water column increases with height because of path-length resistance and gravity. We used morphological measurements to estimate the hydraulic properties of the bordered pits between tracheids in Douglas-fir trees along a height gradient of 85 m. With increasing height, the xylem structural modifications that satisfied hydraulic requirements for avoidance of runaway embolism imposed increasing constraints on water transport efficiency. In the branches and trunks, the pit aperture diameter of tracheids decreases steadily with height, whereas torus diameter remains relatively constant. The resulting increase in the ratio of torus to pit aperture diameter allows the pits to withstand higher tensions before air-seeding but at the cost of reduced pit aperture conductance. Extrapolations of vertical trends for trunks and branches show that water transport across pits will approach zero at a heights of 109 m and 138 m, respectively, which is consistent with historic height records of 100-127 m for this species. Likewise, the twig water potential corresponding to the threshold for runaway embolism would be attained at a height of approximately 107 m. Our results suggest that the maximum height of Douglas-fir trees may be limited in part by the conflicting requirements for water transport and water column safety.
Soper, Fiona M; Boutton, Thomas W; Sparks, Jed P
2015-01-01
Biological nitrogen fixation (BNF) in woody plants is often investigated using foliar measurements of δ(15) N and is of particular interest in ecosystems experiencing increases in BNF due to woody plant encroachment. We sampled δ(15) N along the entire N uptake pathway including soil solution, xylem sap and foliage to (1) test assumptions inherent to the use of foliar δ(15) N as a proxy for BNF; (2) determine whether seasonal divergences occur between δ(15) Nxylem sap and δ(15) Nsoil inorganic N that could be used to infer variation in BNF; and (3) assess patterns of δ(15) N with tree age as indicators of shifting BNF or N cycling. Measurements of woody N-fixing Prosopis glandulosa and paired reference non-fixing Zanthoxylum fagara at three seasonal time points showed that δ(15) Nsoil inorganic N varied temporally and spatially between species. Fractionation between xylem and foliar δ(15) N was consistently opposite in direction between species and varied on average by 2.4‰. Accounting for these sources of variation caused percent nitrogen derived from fixation values for Prosopis to vary by up to ∼70%. Soil-xylem δ(15) N separation varied temporally and increased with Prosopis age, suggesting seasonal variation in N cycling and BNF and potential long-term increases in BNF not apparent through foliar sampling alone. © 2014 John Wiley & Sons Ltd.
Ceratocystis smalleyi colonization of bitternut hickkory and host responses in the xylem
J.-H. Park; J. Juzwik
2014-01-01
Colonization of Carya cordiformis sapwood by Ceratocystis smalleyi and subsequent host defence responses following artificial inoculation were investigated using anatomical and histological techniques. Hyphae of C. smalleyi were observed in all sapwood xylem features confirming the ability of the pathogen to...
Relating xylem cavitation to gas exchange in cotton
USDA-ARS?s Scientific Manuscript database
Acoustic emissions (AEs) from xylem cavitation events are characteristic of transpiration processes. Though a body of work using AE exists with a large number of species, cotton and other agronomically important crops have either not been investigated, or limited information exists. The objective of...
Relating xylem cavitation to transpiration in cotton
USDA-ARS?s Scientific Manuscript database
Acoustic emmisions (AEs) from xylem cavitation events are characteristic of transpiration processes. Even though a body of work employing AE exists with a large number of species, cotton and other agronomically important crops have either not been investigated, or limited information exists. A few s...
X-ray CT and histological imaging of xylem vessels organization in Mimosa pudica.
Lee, Sang Joon; Song, Kahye; Kim, Hae Koo; Park, Joonghyuk
2013-11-01
Mimosa pudica has three distinct specialized organs, namely, pulvinus, secondary pulvinus, and pulvinule, which are respectively controlling the movements of petioles, leaflets, and pinna in response to external stimuli. Water flow is a key factor for such movements, but detailed studies on the organization of the vascular system for water transport in these organs have not been published yet. In this study, organizations of the xylem vessels and morphological features of the pulvinus, the secondary pulvinus, and the pulvinule were experimentally investigated by X-ray computed tomography and histological technique. Results showed that the xylem vessels were circularly distributed in the specialized motile organs and reorganized into distinct vascular bundles at the extremities. The number and the total cross-sectional area of the xylem vessels were increased inside the specialized motile organs. Morphological characteristics obtained in this study provided new insight to understand the functions of the vascular networks in the dynamic movements of M. pudica. Copyright © 2013 Wiley Periodicals, Inc.
NASA Astrophysics Data System (ADS)
Park, Keunhwan; Tixier, Aude; Christensen, Anneline; Arnbjerg-Nielsen, Sif; Zwieniecki, Maciej; Jensen, Kaare
2017-11-01
Water and minerals flow from plant roots to leaves in the xylem, an interconnected network of vascular conduits that spans the full length of the organism. When a plant is subjected to drought stress, air pockets can spread inside the xylem, threatening the survival of the plant. Many plants prevent propagation of air by using hydrophobic nano-membranes in the ``pit'' pores that link adjacent xylem cells. This adds considerable resistance to flow. Interestingly, torus-margo pit pores in conifers are open and offer less resistance. To prevent propagation of air, conifers use a soft gating mechanism, which relies on hydrodynamic interactions between the xylem liquid and the elastic pit. However, it is unknown exactly how it is able to combine the seemingly antagonist functions of high permeability and resistance to propagation of air. We conduct experiments on biomimetic pores to elucidate the flow regulation mechanism. The design of plant valves is compared to other natural systems and optimal strategies are discussed. This work was supported by a research Grant (13166) from VILLUM FONDEN.
Ciraulo, Maristela Boaceff; Santos, Daiene Souza; Rodrigues, Ana Claudia de Freitas Oliveira; de Oliveira, Marcus Vinícius; Rodrigues, Tiago; de Oliveira, Regina Costa; Nunes, Luiz R
2010-01-01
Xylella fastidiosa is a xylem-limited bacterium responsible for important plant diseases, like citrus-variegated chlorosis (CVC) and grapevine Pierce's disease (PD). Interestingly, in vitro growth of X. fastidiosa in chemically defined media that resemble xylem fluid has been achieved, allowing studies of metabolic processes used by xylem-dwelling bacteria to thrive in such nutrient-poor conditions. Thus, we performed microarray hybridizations to compare transcriptomes of X. fastidiosa cells grown in 3G10-R, a medium that resembles grape sap, and in Periwinkle Wilt (PW), the complex medium traditionally used to cultivate X. fastidiosa. We identified 299 transcripts modulated in response to growth in these media. Some 3G10R-overexpressed genes have been shown to be upregulated in cells directly isolated from infected plants and may be involved in plant colonization, virulence and environmental competition. In contrast, cells cultivated in PW show a metabolic switch associated with increased aerobic respiration and enhanced bacterial growth rates.
Ceballos-Laita, Laura; Gutierrez-Carbonell, Elain; Takahashi, Daisuke; Abadía, Anunciación; Uemura, Matsuo; Abadía, Javier; López-Millán, Ana Flor
2018-04-01
This article contains consolidated proteomic data obtained from xylem sap collected from tomato plants grown in Fe- and Mn-sufficient control, as well as Fe-deficient and Mn-deficient conditions. Data presented here cover proteins identified and quantified by shotgun proteomics and Progenesis LC-MS analyses: proteins identified with at least two peptides and showing changes statistically significant (ANOVA; p ≤ 0.05) and above a biologically relevant selected threshold (fold ≥ 2) between treatments are listed. The comparison between Fe-deficient, Mn-deficient and control xylem sap samples using a multivariate statistical data analysis (Principal Component Analysis, PCA) is also included. Data included in this article are discussed in depth in the research article entitled "Effects of Fe and Mn deficiencies on the protein profiles of tomato ( Solanum lycopersicum) xylem sap as revealed by shotgun analyses" [1]. This dataset is made available to support the cited study as well to extend analyses at a later stage.
Ciraulo, Maristela Boaceff; Santos, Daiene Souza; Rodrigues, Ana Claudia de Freitas Oliveira; de Oliveira, Marcus Vinícius; Rodrigues, Tiago; de Oliveira, Regina Costa; Nunes, Luiz R.
2010-01-01
Xylella fastidiosa is a xylem-limited bacterium responsible for important plant diseases, like citrus-variegated chlorosis (CVC) and grapevine Pierce's disease (PD). Interestingly, in vitro growth of X. fastidiosa in chemically defined media that resemble xylem fluid has been achieved, allowing studies of metabolic processes used by xylem-dwelling bacteria to thrive in such nutrient-poor conditions. Thus, we performed microarray hybridizations to compare transcriptomes of X. fastidiosa cells grown in 3G10-R, a medium that resembles grape sap, and in Periwinkle Wilt (PW), the complex medium traditionally used to cultivate X. fastidiosa. We identified 299 transcripts modulated in response to growth in these media. Some 3G10R-overexpressed genes have been shown to be upregulated in cells directly isolated from infected plants and may be involved in plant colonization, virulence and environmental competition. In contrast, cells cultivated in PW show a metabolic switch associated with increased aerobic respiration and enhanced bacterial growth rates. PMID:20625415
Two Complementary Mechanisms Underpin Cell Wall Patterning during Xylem Vessel Development[OPEN
Tang, Lu; Barkwill, Sarah; Lathe, Rahul; McFarlane, Heather E.
2017-01-01
The evolution of the plant vasculature was essential for the emergence of terrestrial life. Xylem vessels are solute-transporting elements in the vasculature that possess secondary wall thickenings deposited in intricate patterns. Evenly dispersed microtubule (MT) bands support the formation of these wall thickenings, but how the MTs direct cell wall synthesis during this process remains largely unknown. Cellulose is the major secondary wall constituent and is synthesized by plasma membrane-localized cellulose synthases (CesAs) whose catalytic activity propels them through the membrane. We show that the protein CELLULOSE SYNTHASE INTERACTING1 (CSI1)/POM2 is necessary to align the secondary wall CesAs and MTs during the initial phase of xylem vessel development in Arabidopsis thaliana and rice (Oryza sativa). Surprisingly, these MT-driven patterns successively become imprinted and sufficient to sustain the continued progression of wall thickening in the absence of MTs and CSI1/POM2 function. Hence, two complementary principles underpin wall patterning during xylem vessel development. PMID:28947492
Palaeo‐adaptive Properties of the Xylem of Metasequoia: Mechanical/Hydraulic Compromises
JAGELS, RICHARD; VISSCHER, GEORGE E.; LUCAS, JOHN; GOODELL, BARRY
2003-01-01
The xylem of Metasequoia glyptostroboides Hu et Cheng is characterized by very low density (average specific gravity = 0·27) and tracheids with relatively large dimensions (length and diameter). The microfibril angle in the S2 layer of tracheid walls is large, even in outer rings, suggesting a cambial response to compressive rather than tensile stresses. In some cases, this compressive stress is converted to irreversible strain (plastic deformation), as evidenced by cell wall corrugations. The heartwood is moderately decay resistant, helping to prevent Brazier buckling. These xylem properties are referenced to the measured bending properties of modulus of rupture and modulus of elasticity, and compared with other low‐to‐moderate density conifers. The design strategy for Metasequoia is to produce a mechanically weak but hydraulically efficient xylem that permits rapid height growth and crown development to capture and dominate a wet site environment. The adaptability of these features to a high‐latitude Eocene palaeoenvironment is discussed. PMID:12763758
Ito, Akiko; Sugiura, Toshihiko; Sakamoto, Daisuke; Moriguchi, Takaya
2013-04-01
In order to elucidate which physiological event(s) are involved in the seasonal changes of carbohydrate dynamics during winter, we examined the effects of different low temperatures on the carbohydrate concentrations of Japanese pear (Pyrus pyrifolia (Burm.) Nakai). For four winter seasons, large increases in the sorbitol concentration of shoot xylem sap occurred during mid- to late December, possibly due to the endodormancy completion and low-temperature responses. When trees were kept at 15 °C from 3 November to 3 December in order to postpone the initiation and completion of chilling accumulation that would break endodormancy, sorbitol accumulation in xylem sap was always higher from trees with sufficient chilling accumulation than from trees that received insufficient chilling. However, an additional increase in xylem sap sorbitol occurred around late December in trees regardless of whether their chilling accumulation naturally progressed or was postponed. To examine different temperature effects more closely, we compared the carbohydrate concentrations of trees subjected to either 6 or 0 °C treatment. The sorbitol concentration in xylem sap tremendously increased at 0 °C treatment compared with 6 °C treatment. However, an additional increase in xylem sap sorbitol occurred at both the temperatures when sufficient chilling accumulated with a peak coinciding with the peak expression in shoots of the sorbitol transporter gene (PpSOT2). Interestingly, the total carbohydrate concentration of shoots tremendously increased with exposure to 0 °C compared with exposure to 6 °C, but was not affected by the amount of accumulated chilling. Instead, as chilling accumulated the ratio of sorbitol to total soluble sugars in shoots increased. We presumed that carbohydrates in the shoot tissues may be converted to sorbitol and loaded into the xylem sap so that the sorbitol accumulation patterns were synchronized with the progression of dormancy, whereas the total carbohydrate transported into shoots from other storage organs may be related to freezing tolerance acquisition independent of dormancy progression. We thus propose that there are different effects of dormancy progression and low-temperature responses on carbohydrate dynamics in Japanese pear.
2011-01-01
Background Lignin, after cellulose, is the second most abundant biopolymer accounting for approximately 15-35% of the dry weight of wood. As an important component during wood formation, lignin is indispensable for plant structure and defense. However, it is an undesirable component in the pulp and paper industry. Removal of lignin from cellulose is costly and environmentally hazardous process. Tremendous efforts have been devoted to understand the role of enzymes and genes in controlling the amount and composition of lignin to be deposited in the cell wall. However, studies on the impact of downregulation and overexpression of monolignol biosynthesis genes in model species on lignin content, plant fitness and viability have been inconsistent. Recently, non-coding RNAs have been discovered to play an important role in regulating the entire monolignol biosynthesis pathway. As small RNAs have critical functions in various biological process during wood formation, small RNA profiling is an important tool for the identification of complete set of differentially expressed small RNAs between low lignin and high lignin secondary xylem. Results In line with this, we have generated two small RNAs libraries from samples with contrasting lignin content using Illumina GAII sequencer. About 10 million sequence reads were obtained in secondary xylem of Am48 with high lignin content (41%) and a corresponding 14 million sequence reads were obtained in secondary xylem of Am54 with low lignin content (21%). Our results suggested that A. mangium small RNAs are composed of a set of 12 highly conserved miRNAs families found in plant miRNAs database, 82 novel miRNAs and a large proportion of non-conserved small RNAs with low expression levels. The predicted target genes of those differentially expressed conserved and non-conserved miRNAs include transcription factors associated with regulation of the lignin biosynthetic pathway genes. Some of these small RNAs play an important role in epigenetic silencing. Differential expression of the small RNAs between secondary xylem tissues with contrasting lignin content suggests that a cascade of miRNAs play an interconnected role in regulating the lignin biosynthetic pathway in Acacia species. Conclusions Our study critically demonstrated the roles of small RNAs during secondary wall formation. Comparison of the expression pattern of small RNAs between secondary xylem tissues with contrasting lignin content strongly indicated that small RNAs play a key regulatory role during lignin biosynthesis. Our analyses suggest an evolutionary mechanism for miRNA targets on the basis of the length of their 5’ and 3’ UTRs and their cellular roles. The results obtained can be used to better understand the roles of small RNAs during lignin biosynthesis and for the development of gene constructs for silencing of specific genes involved in monolignol biosynthesis with minimal effect on plant fitness and viability. For the first time, small RNAs were proven to play an important regulatory role during lignin biosynthesis in A. mangium. PMID:22369296
Ong, Seong Siang; Wickneswari, Ratnam
2011-11-30
Lignin, after cellulose, is the second most abundant biopolymer accounting for approximately 15-35% of the dry weight of wood. As an important component during wood formation, lignin is indispensable for plant structure and defense. However, it is an undesirable component in the pulp and paper industry. Removal of lignin from cellulose is costly and environmentally hazardous process. Tremendous efforts have been devoted to understand the role of enzymes and genes in controlling the amount and composition of lignin to be deposited in the cell wall. However, studies on the impact of downregulation and overexpression of monolignol biosynthesis genes in model species on lignin content, plant fitness and viability have been inconsistent. Recently, non-coding RNAs have been discovered to play an important role in regulating the entire monolignol biosynthesis pathway. As small RNAs have critical functions in various biological process during wood formation, small RNA profiling is an important tool for the identification of complete set of differentially expressed small RNAs between low lignin and high lignin secondary xylem. In line with this, we have generated two small RNAs libraries from samples with contrasting lignin content using Illumina GAII sequencer. About 10 million sequence reads were obtained in secondary xylem of Am48 with high lignin content (41%) and a corresponding 14 million sequence reads were obtained in secondary xylem of Am54 with low lignin content (21%). Our results suggested that A. mangium small RNAs are composed of a set of 12 highly conserved miRNAs families found in plant miRNAs database, 82 novel miRNAs and a large proportion of non-conserved small RNAs with low expression levels. The predicted target genes of those differentially expressed conserved and non-conserved miRNAs include transcription factors associated with regulation of the lignin biosynthetic pathway genes. Some of these small RNAs play an important role in epigenetic silencing. Differential expression of the small RNAs between secondary xylem tissues with contrasting lignin content suggests that a cascade of miRNAs play an interconnected role in regulating the lignin biosynthetic pathway in Acacia species. Our study critically demonstrated the roles of small RNAs during secondary wall formation. Comparison of the expression pattern of small RNAs between secondary xylem tissues with contrasting lignin content strongly indicated that small RNAs play a key regulatory role during lignin biosynthesis. Our analyses suggest an evolutionary mechanism for miRNA targets on the basis of the length of their 5' and 3' UTRs and their cellular roles. The results obtained can be used to better understand the roles of small RNAs during lignin biosynthesis and for the development of gene constructs for silencing of specific genes involved in monolignol biosynthesis with minimal effect on plant fitness and viability. For the first time, small RNAs were proven to play an important regulatory role during lignin biosynthesis in A. mangium.
Leaf vein xylem conduit diameter influences susceptibility to embolism and hydraulic decline
USDA-ARS?s Scientific Manuscript database
Ecosystems worldwide are facing increasingly severe and prolonged droughts during which hydraulic failure from drought-induced embolism can lead to organ or whole plant death. Understanding the determinants of xylem failure across species is critical especially in leaves, the engine of plant growth....
Seasonal development of cambial activity in relation to xylem formation in Chinese fir.
Wu, Hongyang; Xu, Huimin; Li, Hanyin; Wei, Dongmei; Lin, Jinxing; Li, Xiaojuan
2016-05-20
The vascular cambium is a lateral meristem which can differentiate into secondary phloem and xylem. The secondary growth of woody plants resulting from vascular cambium activity has been a focus of considerable attention, but the quantitative relationships between cambial activity and secondary xylem formation have been little studied. Our analysis of cytological changes in the cambium of Chinese fir (Cunninghamia lanceolata), revealed a significant positive correlation between vascular cambium cell numbers and cambium zone width through the seasonal cycle. Cambium cell numbers and the cambium cell radial diameter were closely related to xylem formation. Immuno-labeling showed that de-esterified homogalacturonan and (1-4)-β-d-galactan epitopes were highly abundant in cell walls of dormant-stage cambium, whereas high methylesterified homogalacturonan was strongly labeled in the active stage. Raman spectroscopy detected significant changes in the chemical composition of cell walls during the active-dormant stage transition. More pectin and less monolignols occurred in radial cell walls than in tangential walls during the dormant stage, but no significant changes were found in other stages, indicating that pectin accumulation facilitates cell wall expansion, with cambium activity transition. Our quantitative analysis of the relationship between cambial activity and xylem formation, as well as the cell wall modification during the active stage provides useful information about cambial characteristics and xylogenesis. Copyright © 2016. Published by Elsevier GmbH.
Burbank, Lindsey P; Stenger, Drake C
2017-04-01
Xylella fastidiosa, the causal agent of Pierce's disease of grapes, is a slow-growing, xylem-limited, bacterial pathogen. Disease progression is characterized by systemic spread of the bacterium through xylem vessel networks, causing leaf-scorching symptoms, senescence, and vine decline. It appears to be advantageous to this pathogen to avoid excessive blockage of xylem vessels, because living bacterial cells are generally found in plant tissue with low bacterial cell density and minimal scorching symptoms. The DinJ/RelE toxin-antitoxin system is characterized here for a role in controlling bacterial proliferation and population size during plant colonization. The DinJ/RelE locus is transcribed from two separate promoters, allowing for coexpression of antitoxin DinJ with endoribonuclease toxin RelE, in addition to independent expression of RelE. The ratio of antitoxin/toxin expressed is dependent on bacterial growth conditions, with lower amounts of antitoxin present under conditions designed to mimic grapevine xylem sap. A knockout mutant of DinJ/RelE exhibits a hypervirulent phenotype, with higher bacterial populations and increased symptom development and plant decline. It is likely that DinJ/RelE acts to prevent excessive population growth, contributing to the ability of the pathogen to spread systemically without completely blocking the xylem vessels and increasing probability of acquisition by the insect vector.
Uraguchi, Shimpei; Mori, Shinsuke; Kuramata, Masato; Kawasaki, Akira; Arao, Tomohito; Ishikawa, Satoru
2009-01-01
Physiological properties involved in divergent cadmium (Cd) accumulation among rice genotypes were characterized using the indica cultivar ‘Habataki’ (high Cd in grains) and the japonica cultivar ‘Sasanishiki’ (low Cd in grains). Time-dependence and concentration-dependence of symplastic Cd absorption in roots were revealed not to be responsible for the different Cd accumulation between the two cultivars because root Cd uptake was not greater in the Cd-accumulating cultivar ‘Habataki’ compared with ‘Sasanishiki’. On the other hand, rapid and greater root-to-shoot Cd translocation was observed in ‘Habataki’, which could be mediated by higher abilities in xylem loading of Cd and transpiration rate as a driving force. To verify whether different abilities in xylem-mediated shoot-to-root translocation generally account for the genotypic variation in shoot Cd accumulation in rice, the world rice core collection, consisting of 69 accessions which covers the genetic diversity of almost 32 000 accessions of cultivated rice, was used. The results showed strong correlation between Cd levels in xylem sap and shoots and grains among the 69 rice accessions. Overall, the results presented in this study revealed that the root-to-shoot Cd translocation via the xylem is the major and common physiological process determining the Cd accumulation level in shoots and grains of rice plants. PMID:19401409
Wolkerstorfer, Silviya V; Rosner, Sabine; Hietz, Peter
2012-10-01
The vulnerability of the xylem to cavitation is an important trait in plant drought resistance and has been quantified by several methods. We present a modified method for the simultaneous measurement of cavitations, recorded as ultrasound acoustic emissions (UAEs), and the water potential, measured with a thermocouple psychrometer, in small samples of conifer wood. Analyzing the amplitude of the individual signals showed that a first phase, during which the mean amplitude increased, was followed by a second phase with distinctly lower signal amplitudes. We provide a method to separate the two groups of signals and show that for many samples plausible vulnerability curves require rejecting late low-energy UAEs. These very likely do not result from cavitations. This method was used to analyze the differences between juvenile wood, and early and late mature wood in Picea abies (L.) Karst. Juvenile earlywood was more resistant to cavitation than mature earlywood or latewood, which we relate to the tracheid anatomy of the samples. Copyright © Physiologia Plantarum 2012.
DOE Office of Scientific and Technical Information (OSTI.GOV)
Christoffersen, Bradley O.; Gloor, Manuel; Fauset, Sophie
Forest ecosystem models based on heuristic water stress functions poorly predict tropical forest response to drought partly because they do not capture the diversity of hydraulic traits (including variation in tree size) observed in tropical forests. We developed a continuous porous media approach to modeling plant hydraulics in which all parameters of the constitutive equations are biologically interpretable and measurable plant hydraulic traits (e.g., turgor loss point π tlp, bulk elastic modulus ε, hydraulic capacitance C ft, xylem hydraulic conductivity k s,max, water potential at 50 % loss of conductivity for both xylem ( P 50,x) and stomata ( Pmore » 50,gs), and the leaf : sapwood area ratio A l: A s). We embedded this plant hydraulics model within a trait forest simulator (TFS) that models light environments of individual trees and their upper boundary conditions (transpiration), as well as providing a means for parameterizing variation in hydraulic traits among individuals. We synthesized literature and existing databases to parameterize all hydraulic traits as a function of stem and leaf traits, including wood density (WD), leaf mass per area (LMA), and photosynthetic capacity ( A max ), and evaluated the coupled model (called TFS v.1-Hydro) predictions, against observed diurnal and seasonal variability in stem and leaf water potential as well as stand-scaled sap flux. Our hydraulic trait synthesis revealed coordination among leaf and xylem hydraulic traits and statistically significant relationships of most hydraulic traits with more easily measured plant traits. Using the most informative empirical trait–trait relationships derived from this synthesis, TFS v.1-Hydro successfully captured individual variation in leaf and stem water potential due to increasing tree size and light environment, with model representation of hydraulic architecture and plant traits exerting primary and secondary controls, respectively, on the fidelity of model predictions. The plant hydraulics model made substantial improvements to simulations of total ecosystem transpiration. As a result, remaining uncertainties and limitations of the trait paradigm for plant hydraulics modeling are highlighted.« less
Christoffersen, Bradley O.; Gloor, Manuel; Fauset, Sophie; ...
2016-11-24
Forest ecosystem models based on heuristic water stress functions poorly predict tropical forest response to drought partly because they do not capture the diversity of hydraulic traits (including variation in tree size) observed in tropical forests. We developed a continuous porous media approach to modeling plant hydraulics in which all parameters of the constitutive equations are biologically interpretable and measurable plant hydraulic traits (e.g., turgor loss point π tlp, bulk elastic modulus ε, hydraulic capacitance C ft, xylem hydraulic conductivity k s,max, water potential at 50 % loss of conductivity for both xylem ( P 50,x) and stomata ( Pmore » 50,gs), and the leaf : sapwood area ratio A l: A s). We embedded this plant hydraulics model within a trait forest simulator (TFS) that models light environments of individual trees and their upper boundary conditions (transpiration), as well as providing a means for parameterizing variation in hydraulic traits among individuals. We synthesized literature and existing databases to parameterize all hydraulic traits as a function of stem and leaf traits, including wood density (WD), leaf mass per area (LMA), and photosynthetic capacity ( A max ), and evaluated the coupled model (called TFS v.1-Hydro) predictions, against observed diurnal and seasonal variability in stem and leaf water potential as well as stand-scaled sap flux. Our hydraulic trait synthesis revealed coordination among leaf and xylem hydraulic traits and statistically significant relationships of most hydraulic traits with more easily measured plant traits. Using the most informative empirical trait–trait relationships derived from this synthesis, TFS v.1-Hydro successfully captured individual variation in leaf and stem water potential due to increasing tree size and light environment, with model representation of hydraulic architecture and plant traits exerting primary and secondary controls, respectively, on the fidelity of model predictions. The plant hydraulics model made substantial improvements to simulations of total ecosystem transpiration. As a result, remaining uncertainties and limitations of the trait paradigm for plant hydraulics modeling are highlighted.« less
What HR-CT imaging can teach us about xylem structure and function
USDA-ARS?s Scientific Manuscript database
It is well established that plant xylem is composed of a complex and interconnected system of vascular elements, but little is known about how the three-dimensional (3D) organization of this network influences properties such as plant hydraulics (Tyree & Zimmermann, 2002), and few studies have measu...
In vivo visualizations of drought-induced embol 35 ism spread in Vitis vinifera
USDA-ARS?s Scientific Manuscript database
Long distance water transport through plant xylem is vulnerable to hydraulic dysfunction during periods of increased tension on the xylem sap, often coinciding with drought. While the effects of local and systemic embolism on plant water transport and physiology are well documented, the spatial patt...
Metabolic Profiling of Xylem Sap from Pierce’s Disease Resistant and Susceptible Grapevines
USDA-ARS?s Scientific Manuscript database
Pierce’s Disease (PD) of grapevines is caused by a gram-negative, xylem-limited bacterium Xylella fastidiosa (Xf). All Vitis vinifera-based cultivars are highly susceptible to Xf infection. However, some grape species from the southern United States such as V. arizonica, V. Shuttleworthii, and Musca...
Invasion of xylem of mature tree stems by Phytophthora ramorum and P. kernoviae
Anna Brown; Clive Brasier
2008-01-01
The aetiology and frequency of Phytophthoras in discoloured xylem tissue beneath phloem lesions was investigated in a range of broadleaved trees infected with P. ramorum, P. kernoviae and several other Phytophthoras. Isolation was attempted from the inner surface of 81, 6 x 4 cm sterilised...
Ecophysiological responses of three evergreen woody Mediterranean species to water stress
NASA Astrophysics Data System (ADS)
Abril, Mireia; Hanano, Ralph
1998-08-01
The ecophysiological response to drought in three different evergreen Mediterranean species were compared. For a better interpretation of the mechanisms regulating physiological processes, the choice of species was based on evident differences in morphological and structural features (leaf size, leaf specific weight, water-conducting system). Seedlings of Ceanothus thyrsiflorus, Quercus agrifolia and Buxus microphylla grown in pots were subjected to natural stressing conditions during late spring in Southern California. Gas exchange, xylem water potential and abscisic acid concentration in xylem sap were measured in control (irrigated) and water-stressed plants, from predawn to sunset. Environmental, hydraulic and hormonal effects on water control and limitations to photosynthesis were analyzed. Q. agrifolia had the highest maximums of net photosynthesis, stomatal conductance and transpiration, which were significantly different from C. thyrsiflorus and B. microphylla. B. microphylla had the lowest values. Stressed individuals of C. thyrsiflorus and B. microphylla reached absolute minimum water potentials during the day and at predawn. Q. agrifolia plants had a water conservative behaviour and did not show these low values. Control plants from Q. agrifolia had the lowest values of hydraulic resistance with high maximum stomatal conductance, while B. microphylla control plants had the lowest maximum stomatal conductance due to higher hydraulic resistance. Changes in plant hydraulic resistance during soil drying were found, which differed among the species. In general, water-use efficiency was reduced during the day by water stress but increased as seasonal drought proceeded. On a long-term basis, Q. agrifolia was the most efficient species in water use. The results support the hypothesis that information on abscisic acid concentration in xylem sap may be one of the most important physiological keys when modelling stomatal conductance and canopy gas exchange over seasons as drought develops on Mediterranean vegetation. Nevertheless, we believe it is necessary to integrate all known factors that control stomatal behaviour in order to construct general models of the vegetation response to environmental changes.
Impact of warming and drought on carbon balance related to wood formation in black spruce.
Deslauriers, Annie; Beaulieu, Marilène; Balducci, Lorena; Giovannelli, Alessio; Gagnon, Michel J; Rossi, Sergio
2014-08-01
Wood formation in trees represents a carbon sink that can be modified in the case of stress. The way carbon metabolism constrains growth during stress periods (high temperature and water deficit) is now under debate. In this study, the amounts of non-structural carbohydrates (NSCs) for xylogenesis in black spruce, Picea mariana, saplings were assessed under high temperature and drought in order to determine the role of sugar mobilization for osmotic purposes and its consequences for secondary growth. Four-year-old saplings of black spruce in a greenhouse were subjected to different thermal conditions with respect to the outside air temperature (T0) in 2010 (2 and 5 °C higher than T0) and 2011 (6 °C warmer than T0 during the day or night) with a dry period of about 1 month in June of each year. Wood formation together with starch, NSCs and leaf parameters (water potential and photosynthesis) were monitored from May to September. With the exception of raffinose, the amounts of soluble sugars were not modified in the cambium even if gas exchange and photosynthesis were greatly reduced during drought. Raffinose increased more than pinitol under a pre-dawn water potential of less than -1 Mpa, presumably because this compound is better suited than polyol for replacing water and capturing free radicals, and its degradation into simple sugar is easier. Warming decreased the starch storage in the xylem as well the available hexose pool in the cambium and the xylem, probably because of an increase in respiration. Radial stem growth was reduced during drought due to the mobilization of NSCs for osmotic purposes and due to the lack of cell turgor. Thus plant water status during wood formation can influence the NSCs available for growth in the cambium and xylem. © The Author 2014. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please email: journals.permissions@oup.com.
Nickel affects xylem Sap RNase a and converts RNase A to a urease
2013-01-01
Background Nickel (Ni) is an essential micronutrient; however, its metabolic or physiological functions in plants and animals are largely uncharacterized. The ribonucleases (RNase, e.g., RNase A) are a large family of hydrolases found in one form or many forms facilitating nitrogen (N) cycling. It is currently unknown how either a deficiency or excess of Ni influences the functionality of ribonucleases, like RNase A. This is especially true for perennial crops possessing relatively high Ni requirements. Results We report that the 'rising’ xylem sap of pecan [Carya illinoinensis (Wangenh.) K. Koch, a long-lived tree] at bud break contains a 14 kDa RNase A (aka, RNase 1), which amount has a 33% greater in Ni-deficient as in Ni-sufficient trees when exposed to Ni ions exhibits ureolytic activity. The homologous 13.4 kDa bovine pancreatic RNase A likewise exhibits ureolytic activity upon exposure to Ni ions. Ni therefore affects enzymatic function of a typically non-metalloenzyme, such as it transforms to an enzyme capable of hydrolyzing a linear amide; thus, converting an endonuclease esterase into a urease. Conclusions We conclude that Ni potentially affects the level and activity of RNase A present in the spring xylem sap of pecan trees, and probably in other crops, it has the same influence. The catalytic property of RNase A appears to shift from a nuclease to a urease relying on Ni exposure. This is suggestive that RNase A might possess novel metabolic functionality regarding N-metabolism in perennial plants. The ability of Ni to convert the activity of plant and animal RNase A from that of a ribonuclease to a urease indicates a possible unrecognized beneficial metabolic function of Ni in organisms, while also identifying a potential detrimental effect of excessive Ni on N related metabolic activity if there is sufficient disruption of Ni homeostasis. PMID:24320827
NASA Astrophysics Data System (ADS)
Koppan, A.; Fenyvesi, A.; Szarka, L.; Wesztergom, V.
2002-05-01
Electrical potential differences (EPD) in the trunk of a Turkey oak tree (measured by using non-polarising electrodes deepened in the sap wood) have been continuously recorded in the Geophysical Observatory "Istv n Széchenyi" of the Hungarian Academy of Sciences since 1997. Besides of various geophysical observations, meteorological and direct sap-flow measurements have also been carried out in the observatory. As it was found (Kopp n A., Szarka L., Wesztergom V., 2000: Annual fluctuation in amplitudes of daily variations of electrical signals measured in the trunk of a standing tree. C.R. Acad. Sci. Paris, Life Sciences 323, 559-563), the measured electric potential difference data have a characteristic sinusoidal daily fluctuation, and the intensity of the diurnal variations has a double-peak annual characteristics, which coincides with the life activity maximums of the tree. We have found a remarkable inter-correlation between trunk EPD, water potential of air (derived from meteorological data), and direct sap flow velocity data from a neighboring tree. All these results clearly demonstrate that the sap streaming due to the transpiration and root pressure generates the largest part of measured potential differences. The ratio of the flow velocity of a diluted solution forced through stems and the potential differences was found to be constant (Gindl, W., L”ppert, H.-G., Wimmer, R., 1999: Relationship between streaming potential and sap velocity in Salix alba L. Phyton, 39, 217-224.). On the contrary in our in-vivo experiments the relationship between the measured sap flow velocity and EPD is non-linear, which means that the conductivity (i.e. ion concentration) of the xylem sap itself also has a daily fluctuation.
R.W. Hemingway; G.W. McGraw; S.J. Barras
1977-01-01
Since Ceratocystis minor is central to the death of pines infested by southern pine beetles, changes in polyphenols of infected loblolly pine were examined with regard to accumulation of fungal metabolites and changes in concentrations of fungitoxic and fungistatic phloem and xylem constitutents. C. minor grown in liquid culture...
Polyphenols in Ceratocystis minor infected Pinus taeda: fungal metabolites, phloem and xylem phenols
Richard W. Hemingway; Gerald W. McGraw; Stanley J. Barras
1977-01-01
Since Ceratocystis minor is central to the death of pines infested by southern pine beetles, changes in polyphenols of infected loblolly pine were examined with regard to accumulation of fungal metabolites and changes in concentrations of fungitoxic and fungistatic phloem and xylem constitutents. C. minor grown in liquid culture...
USDA-ARS?s Scientific Manuscript database
The formation of emboli in xylem conduits can dramatically reduce hydraulic capacity and represents one of the principal mechanisms of drought induced mortality in woody plants. Some angiosperm species possess a mechanism to rapidly repair embolism by dissolving gas back into solution. However, it i...
Direct micro-CT observation confirms the induction of embolism upon xylem cutting under tension
USDA-ARS?s Scientific Manuscript database
We used two different Synchrotron-based micro-CT facilities (SLS: Swiss Light Source, Villigen, Switzerland, and ALS: Advanced Light Source, Berkeley, CA USA) to test the excision artifact described by Wheeler et al. (2013). Specifically, we examined the impact of cutting xylem under tension and und...
Outside-xylem pathways, not xylem embolism, drive leaf hydraulic decline with dehydration
USDA-ARS?s Scientific Manuscript database
Leaf hydraulic supply is crucial to enable the maintenance of open stomata for CO2 capture and plant growth. During drought-induced leaf dehydration, the capacity for water flow through the leaf (Kleaf) declines, a phenomenon surprisingly attributed for the past fifty years solely to the formation o...
USDA-ARS?s Scientific Manuscript database
During drought, xylem conduits are susceptible to hydraulic dysfunction caused by cavitation and gas embolism. Embolism formation and spread within xylem is dependent on conduit structure and network connectivity, but detailed spatial analysis has been limited due to a lack of non-destructive method...
Xylem Resin in the Resistance of the Pinaceae to Bark Beetles
Richard H. Smith
1972-01-01
Xylem resin of Pinaceae is closely linked with their resistance and suseptibility to tree-killing bark beetles. This review of the literature on attacking adults suggests that all three resistance mechanisms proposed by Painter -- preference, antibiosis, and tolerance -- are active in this relationship: preference by attraction, repellency, and synergism; antibiosis...
Plant fluid proteomics: Delving into the xylem sap, phloem sap and apoplastic fluid proteomes
USDA-ARS?s Scientific Manuscript database
The phloem sap, xylem sap and apoplastic fluid play key roles in long and short distance transport of signals and nutrients, and act as a barrier against local and systemic pathogen infection. Among other components, these plant fluids contain proteins which are likely to be important players in the...
USDA-ARS?s Scientific Manuscript database
Xylella fastidiosa, the causal agent of Pierce’s disease of grapes, is a slow-growing, xylem-limited, bacterial pathogen. Disease progression is characterized by systemic spread of the bacterium through xylem vessel networks, causing leaf scorching symptoms, senescence, and vine decline. It appears ...
BRANCH JUNCTIONS AND THE FLOW OF WATER THROUGH XYLEM IN DOUGLAS-FIR AND PONDEROSA PINE STEMS
Water flowing through the xylem of most plants from the roots to the leaves must pass through junctions where branches have developed from the main stem. These junctions have been studied as both flow constrictions and components of a hydraulic segmentation mechanism to protect ...
Plant fluid proteomics: Delving into the xylem sap, phloem sap and apoplastic fluid proteomes
USDA-ARS?s Scientific Manuscript database
The phloem sap, xylem sap and apoplastic fluid play key roles in long and short distance transport of signals and nutrients, and act as a barrier against local and systemic pathogen infection. Among other components, these plant fluids contain proteins, which are likely to be important players in th...
A Plumber's-Eye View of Xylem Water Transport in Woody Plants
ERIC Educational Resources Information Center
Martinez-Vilalta, Jordi; Pinol, Josep
2004-01-01
We present a practical for university-level students aimed at measuring and comparing xylem hydraulic properties of co-existing plant species. After sampling branches of several woody species in the field, their main hydraulic properties were measured using a simple set-up. Hydraulic conductivity ("K[subscript h]") was calculated as the ratio…
A Proteomic Study of Clavibacter Michiganensis Subsp. Michiganensis Culture Supernatants
Hiery, Eva; Poetsch, Ansgar; Moosbauer, Tanja; Amin, Bushra; Hofmann, Jörg; Burkovski, Andreas
2015-01-01
Clavibacter michiganensis, subsp. michiganensis is a Gram-positive plant pathogen infecting tomato (Solanum lycopersicum). Despite a considerable economic importance due to significant losses of infected plants and fruits, knowledge about virulence factors of C. michiganensis subsp. michiganensis and host-pathogen interactions on a molecular level are rather limited. In the study presented here, the proteome of culture supernatants from C. michiganensis subsp. michiganensis NCPPB382 was analyzed. In total, 1872 proteins were identified in M9 and 1766 proteins in xylem mimicking medium. Filtration of supernatants before protein precipitation reduced these to 1276 proteins in M9 and 976 proteins in the xylem mimicking medium culture filtrate. The results obtained indicate that C. michiganensis subsp. michiganensis reacts to a sucrose- and glucose-depleted medium similar to the xylem sap by utilizing amino acids and host cell polymers as well as their degradation products, mainly peptides, amino acids and various C5 and C6 sugars. Interestingly, the bacterium expresses the previously described virulence factors Pat-1 and CelA not exclusively after host cell contact in planta but already in M9 minimal and xylem mimicking medium. PMID:28248277
Cabañero, Francisco J; Carvajal, Micaela
2007-10-01
In order to study the effect of nutrient stress on water uptake in pepper plants (Capsicum annuum L.), the excess or deficiency of the main cations involved in plant nutrition (K(+), Mg(2+), Ca(2+)) and two different degrees of salinity were related to the activity of plasma membrane H(+)-ATPase, the pH of the xylem sap, nutrient flux into the xylem (J(s)) and to a number of parameters related to water relations, such as root hydraulic conductance (L(0)), stomatal conductance (g(s)) and aquaporin activity. Excess of K(+), Ca(+) and NaCl produced a toxic effect on L(0) while Mg(2+) starvation produced a positive effect, which was in agreement with aquaporin functionality, but not with ATPase activity. The xylem pH was altered only by Ca treatments. The results obtained with each treatment could suggest that detection of the quality of the nutrient supply being received by roots can be related to aquaporins functionality, but also that each cation stress triggers specific responses that have to be assessed individually.
Two Complementary Mechanisms Underpin Cell Wall Patterning during Xylem Vessel Development.
Schneider, Rene; Tang, Lu; Lampugnani, Edwin R; Barkwill, Sarah; Lathe, Rahul; Zhang, Yi; McFarlane, Heather E; Pesquet, Edouard; Niittyla, Totte; Mansfield, Shawn D; Zhou, Yihua; Persson, Staffan
2017-10-01
The evolution of the plant vasculature was essential for the emergence of terrestrial life. Xylem vessels are solute-transporting elements in the vasculature that possess secondary wall thickenings deposited in intricate patterns. Evenly dispersed microtubule (MT) bands support the formation of these wall thickenings, but how the MTs direct cell wall synthesis during this process remains largely unknown. Cellulose is the major secondary wall constituent and is synthesized by plasma membrane-localized cellulose synthases (CesAs) whose catalytic activity propels them through the membrane. We show that the protein CELLULOSE SYNTHASE INTERACTING1 (CSI1)/POM2 is necessary to align the secondary wall CesAs and MTs during the initial phase of xylem vessel development in Arabidopsis thaliana and rice ( Oryza sativa ). Surprisingly, these MT-driven patterns successively become imprinted and sufficient to sustain the continued progression of wall thickening in the absence of MTs and CSI1/POM2 function. Hence, two complementary principles underpin wall patterning during xylem vessel development. © 2017 American Society of Plant Biologists. All rights reserved.
Hydrolase treatments help unravel the function of intervessel pits in xylem hydraulics.
Dusotoit-Coucaud, Anaïs; Brunel, Nicole; Tixier, Aude; Cochard, Hervé; Herbette, Stéphane
2014-03-01
Intervessel pits are structures that play a key role in the efficiency and safety functions of xylem hydraulics. However, little is known about the components of the pit membrane (PM) and their role in hydraulic functions, especially in resistance to cavitation. We tested the effect of commercial chemicals including a cellulase, a hemicellulase, a pectolyase, a proteinase and DTT on xylem hydraulic properties: vulnerability to cavitation (VC) and conductance. The effects were tested on branch segments from Fagus sylvatica (where the effects on pit structure were analyzed using TEM) and Populus tremula. Cellulose hydrolysis resulted in a sharp increase in VC and a significant increase in conductance, related to complete breakdown of the PM. Pectin hydrolysis also induced a sharp increase in VC but with no effect on conductance or pit structure observable by TEM. The other treatments with hemicellulase, proteinase or DTT showed no effect. This study brings evidence that cellulose and pectins are critical components underpinning VC, and that PM components may play distinct roles in the xylem hydraulic safety and efficiency. © 2013 Scandinavian Plant Physiology Society.
Koddenberg, Tim; Militz, Holger
2018-05-05
The popularity of X-ray based imaging methods has continued to increase in research domains. In wood research, X-ray micro-computed tomography (XμCT) is useful for structural studies examining the three-dimensional and complex xylem tissue of trees qualitatively and quantitatively. In this study, XμCT made it possible to visualize and quantify the spatial xylem organization of the angiosperm species Fraxinus excelsior L. on the microscopic level. Through image analysis, it was possible to determine morphological characteristics of the cellular axial tissue (vessel elements, fibers, and axial parenchyma cells) three-dimensionally. X-ray imaging at high resolutions provides very distinct visual insight into the xylem structure. Numerical analyses performed through semi-automatic procedures made it possible to quickly quantify cell characteristics (length, diameter, and volume of cells). Use of various spatial resolutions (0.87-5 μm) revealed boundaries users should be aware of. Nevertheless, our findings, both qualitative and quantitative, demonstrate XμCT to be a valuable tool for studying the spatial cell morphology of F. excelsior. Copyright © 2018. Published by Elsevier Ltd.
A Miniature Stem Thermocouple Hygrometer 1
Michel, Burlyn E.
1977-01-01
An unprotected chromel-constantan thermocouple was mounted in a cavity (4 × 2 × 1 mm) with rounded corners in a chrome-plated brass block (10 × 6 × 4 mm). When installed against a soybean (Glycine max [L.] Merr.) xylem face, sealed with caulking gum, and insulated with polyurethane foam and aluminum foil, even rapidly changing stem water potentials could be followed accurately. Temperature gradients can be a problem. PMID:16660155
Competitive Effects of Various Grasses and Forbs on Ponderosa Pine Seedlings
Katherine J. Elliott; Alan S. White
1987-01-01
Competition between ponderosa pine seedlings and various grasses and forbs was studied on :I site in northern Arizona burned in 1982 by a wildfire. Two-year-old pine seedlings were planted in 3.05 x 3.05 m plots in April 1983, followed by the sowing of grass and forb seeds on the same plots in July 1983 after summer rains had begun. Predawn xylem water potential...
Cornu, Jean-Yves; Deinlein, Ulrich; Höreth, Stephan; Braun, Manuel; Schmidt, Holger; Weber, Michael; Persson, Daniel P; Husted, Søren; Schjoerring, Jan K; Clemens, Stephan
2015-04-01
Elevated nicotianamine synthesis in roots of Arabidopsis halleri has been established as a zinc (Zn) hyperaccumulation factor. The main objective of this study was to elucidate the mechanism of nicotianamine-dependent root-to-shoot translocation of metals. Metal tolerance and accumulation in wild-type (WT) and AhNAS2-RNA interference (RNAi) plants were analysed. Xylem exudates were subjected to speciation analysis and metabolite profiling. Suppression of root nicotianamine synthesis had no effect on Zn and cadmium (Cd) tolerance but rendered plants nickel (Ni)-hypersensitive. It also led to a reduction of Zn root-to-shoot translocation, yet had the opposite effect on Ni mobility, even though both metals form coordination complexes of similar stability with nicotianamine. Xylem Zn concentrations were positively, yet nonstoichiometrically, correlated with nicotianamine concentrations. Two fractions containing Zn coordination complexes were detected in WT xylem. One of them was strongly reduced in AhNAS2-suppressed plants and coeluted with (67) Zn-labelled organic acid complexes. Organic acid concentrations were not responsive to nicotianamine concentrations and sufficiently high to account for complexing the coordinated Zn. We propose a key role for nicotianamine in controlling the efficiency of Zn xylem loading and thereby the formation of Zn coordination complexes with organic acids, which are the main Zn ligands in the xylem but are not rate-limiting for Zn translocation. © 2014 The Authors. New Phytologist © 2014 New Phytologist Trust.
Pieretti, Isabelle; Royer, Monique; Barbe, Valérie; Carrere, Sébastien; Koebnik, Ralf; Couloux, Arnaud; Darrasse, Armelle; Gouzy, Jérôme; Jacques, Marie-Agnès; Lauber, Emmanuelle; Manceau, Charles; Mangenot, Sophie; Poussier, Stéphane; Segurens, Béatrice; Szurek, Boris; Verdier, Valérie; Arlat, Matthieu; Gabriel, Dean W; Rott, Philippe; Cociancich, Stéphane
2012-11-21
Xanthomonas albilineans causes leaf scald, a lethal disease of sugarcane. X. albilineans exhibits distinctive pathogenic mechanisms, ecology and taxonomy compared to other species of Xanthomonas. For example, this species produces a potent DNA gyrase inhibitor called albicidin that is largely responsible for inducing disease symptoms; its habitat is limited to xylem; and the species exhibits large variability. A first manuscript on the complete genome sequence of the highly pathogenic X. albilineans strain GPE PC73 focused exclusively on distinctive genomic features shared with Xylella fastidiosa-another xylem-limited Xanthomonadaceae. The present manuscript on the same genome sequence aims to describe all other pathogenicity-related genomic features of X. albilineans, and to compare, using suppression subtractive hybridization (SSH), genomic features of two strains differing in pathogenicity. Comparative genomic analyses showed that most of the known pathogenicity factors from other Xanthomonas species are conserved in X. albilineans, with the notable absence of two major determinants of the "artillery" of other plant pathogenic species of Xanthomonas: the xanthan gum biosynthesis gene cluster, and the type III secretion system Hrp (hypersensitive response and pathogenicity). Genomic features specific to X. albilineans that may contribute to specific adaptation of this pathogen to sugarcane xylem vessels were also revealed. SSH experiments led to the identification of 20 genes common to three highly pathogenic strains but missing in a less pathogenic strain. These 20 genes, which include four ABC transporter genes, a methyl-accepting chemotaxis protein gene and an oxidoreductase gene, could play a key role in pathogenicity. With the exception of hypothetical proteins revealed by our comparative genomic analyses and SSH experiments, no genes potentially involved in any offensive or counter-defensive mechanism specific to X. albilineans were identified, supposing that X. albilineans has a reduced artillery compared to other pathogenic Xanthomonas species. Particular attention has therefore been given to genomic features specific to X. albilineans making it more capable of evading sugarcane surveillance systems or resisting sugarcane defense systems. This study confirms that X. albilineans is a highly distinctive species within the genus Xanthomonas, and opens new perpectives towards a greater understanding of the pathogenicity of this destructive sugarcane pathogen.
López, Rosana; López de Heredia, Unai; Collada, Carmen; Cano, Francisco Javier; Emerson, Brent C.; Cochard, Hervé; Gil, Luis
2013-01-01
Background and Aims It is widely accepted that hydraulic failure due to xylem embolism is a key factor contributing to drought-induced mortality in trees. In the present study, an attempt is made to disentangle phenotypic plasticity from genetic variation in hydraulic traits across the entire distribution area of a tree species to detect adaptation to local environments. Methods A series of traits related to hydraulics (vulnerability to cavitation and hydraulic conductivity in branches), growth performance and leaf mass per area were assessed in eight Pinus canariensis populations growing in two common gardens under contrasting environments. In addition, the neutral genetic variability (FST) and the genetic differentiation of phenotypic variation (QST) were compared in order to identify the evolutionary forces acting on these traits. Key Results The variability for hydraulic traits was largely due to phenotypic plasticity. Nevertheless, the vulnerability to cavitation displayed a significant genetic variability (approx. 5 % of the explained variation), and a significant genetic × environment interaction (between 5 and 19 % of the explained variation). The strong correlation between vulnerability to cavitation and survival in the xeric common garden (r = –0·81; P < 0·05) suggests a role for the former in the adaptation to xeric environments. Populations from drier sites and higher temperature seasonality were less vulnerable to cavitation than those growing at mesic sites. No trade-off between xylem safety and efficiency was detected. QST of parameters of the vulnerability curve (0·365 for P50 and the slope of the vulnerability curve and 0·452 for P88) differed substantially from FST (0·091), indicating divergent selection. In contrast, genetic drift alone was found to be sufficient to explain patterns of differentiation for xylem efficiency and growth. Conclusions The ability of P. canariensis to inhabit a wide range of ecosystems seemed to be associated with high phenotypic plasticity and some degree of local adaptations of xylem and leaf traits. Resistance to cavitation conferred adaptive potential for this species to adapt successfully to xeric conditions. PMID:23644361
Protein S-Nitrosylation Regulates Xylem Vessel Cell Differentiation in Arabidopsis.
Kawabe, Harunori; Ohtani, Misato; Kurata, Tetsuya; Sakamoto, Tomoaki; Demura, Taku
2018-01-01
Post-translational modifications of proteins have important roles in the regulation of protein activity. One such modification, S-nitrosylation, involves the covalent binding of nitric oxide (NO)-related species to a cysteine residue. Recent work showed that protein S-nitrosylation has crucial functions in plant development and environmental responses. In the present study, we investigated the importance of protein S-nitrosylation for xylem vessel cell differentiation using a forward genetics approach. We performed ethyl methanesulfonate mutagenesis of a transgenic Arabidopsis 35S::VND7-VP16-GR line in which the activity of VASCULAR-RELATED NAC-DOMAIN7 (VND7), a key transcription factor involved in xylem vessel cell differentiation, can be induced post-translationally by glucocorticoid treatment, with the goal of obtaining suppressor mutants that failed to differentiate ectopic xylem vessel cells; we named these mutants suppressor of ectopic vessel cell differentiation induced by VND7 (seiv) mutants. We found the seiv1 mutant to be a recessive mutant in which ectopic xylem cell differentiation was inhibited, especially in aboveground organs. In seiv1 mutants, a single nucleic acid substitution (G to A) leading to an amino acid substitution (E36K) was present in the gene encoding S-NITROSOGLUTATHIONE REDUCTASE 1 (GSNOR1), which regulates the turnover of the natural NO donor, S-nitrosoglutathione. An in vitro S-nitrosylation assay revealed that VND7 can be S-nitrosylated at Cys264 and Cys320 located near the transactivation activity-related domains, which were shown to be important for transactivation activity of VND7 by transient reporter assay. Our results suggest crucial roles for GSNOR1-regulated protein S-nitrosylation in xylem vessel cell differentiation, partly through the post-translational modification of VND7. © The Author 2017. Published by Oxford University Press on behalf of Japanese Society of Plant Physiologists. All rights reserved. For permissions, please email: journals.permissions@oup.com.
In vivo Visualization of the Water-refilling Process in Xylem Vessels Using X-ray Micro-imaging
Lee, Sang-Joon; Kim, Yangmin
2008-01-01
Background and Aims Xylem vessels containing gases (embolized) must be refilled with water if they are to resume transport of water through the plant, so refilling is of great importance for the maintenance of water balance in plants. However, the refilling process is poorly understood because of inadequate examination methods. Simultaneous measurements of plant anatomy and vessel refilling are essential to elucidate the mechanisms involved. In the present work, a new technique based on phase-contrast X-ray imaging is presented that visualizes, in vivo and in real time, both xylem anatomy and refilling of embolized vessels. Methods With the synchrotron X-ray micro-imaging technique, the refilling of xylem vessels of leaves and a stem of Phyllostachys bambusoides with water is demonstrated under different conditions. The technique employs phase contrast imaging of X-ray beams, which are transformed into visible light and are photographed by a charge coupled device camera. X-ray images were captured consecutively at every 0·5 s with an exposure time of 10 ms. Key Results The interface (meniscus) between the water and gas phases in refilling the xylem vessels is displayed. During refilling, the rising menisci in embolized vessels showed repetitive flow, i.e. they temporarily stopped at the end walls of the vessel elements while gas bubbles were removed. The meniscus then passed through the end wall at a faster rate than the speed of flow in the main vessels. In the light, the speed of refilling in a specific vessel was slower than that in the dark, but this rate increased again after repeated periods in darkness. Conclusions Real-time, non-destructive X-ray micro-imaging is an important, useful and novel technique to study the relationship between xylem structure and the refilling of embolized vessels in intact plants. It provides new insight into understanding the mechanisms of water transport and the refilling of embolized vessels, which are not understood well. PMID:18077466
Lavrič, Martina; Eler, Klemen; Ferlan, Mitja; Vodnik, Dominik; Gričar, Jožica
2017-01-01
Intra-annual variations in leaf development, radial growth, including the phloem part, and sap flow have rarely been studied in deciduous trees from drought-prone environments. In order to understand better the chronological order and temporal course of these processes, we monitored leaf phenology, xylem and phloem formation and sap flow in Quercus pubescens from abandoned karst grasslands in Slovenia during the growing season of 2014. We found that the initial earlywood vessel formation started before bud opening at the beginning of April. Buds started to open in the second half of April and full leaf unfolding occurred by the end of May. LAI values increased correspondingly with leaf development. About 28% of xylem and 22% of phloem annual increment were formed by the time of bud break. Initial earlywood vessels were fully lignified and ready for water transport, indicating that they are essential to provide hydraulic conductivity for axial water flow during leaf development. Sap flow became active and increasing contemporarily with leaf development and LAI values. Similar early spring patterns of xylem sap flow and LAI denoted that water transport in oaks broadly followed canopy leaf area development. In the initial 3 weeks of radial growth, phloem growth preceded that of xylem, indicating its priority over xylem at the beginning of the growing season. This may be related to the fact that after bud break, the developing foliage is a very large sink for carbohydrates but, at the same time, represents a small transpirational area. Whether the interdependence of the chronological sequence of the studied processes is fixed in Q. pubescens needs to be confirmed with more data and several years of analyses, although the ‘correct sequence’ of processes is essential for synchronized plant performance and response to environmental stress. PMID:28321232
Rita, Angelo; Borghetti, Marco; Todaro, Luigi; Saracino, Antonio
2016-01-01
In the Mediterranean region, the widely predicted rise in temperature, change in the precipitation pattern, and increase in the frequency of extreme climatic events are expected to alter the shape of ecological communities and to affect plant physiological processes that regulate ecosystem functioning. Although change in the mean values are important, there is increasing evidence that plant distribution, survival, and productivity respond to extremes rather than to the average climatic condition. The present study aims to assess the effects of both mean and extreme climatic conditions on radial growth and functional anatomical traits using long-term tree-ring time series of two co-existing Quercus spp. from a drought-prone site in Southern Italy. In particular, this is the first attempt to apply the Generalized Additive Model for Location, Scale, and Shape (GAMLSS) technique and Bayesian modeling procedures to xylem traits data set, with the aim of (i) detecting non-linear long-term responses to climate and (ii) exploring relationships between climate extreme and xylem traits variability in terms of probability of occurrence. This study demonstrates the usefulness of long-term xylem trait chronologies as records of environmental conditions at annual resolution. Statistical analyses revealed that most of the variability in tree-ring width and specific hydraulic conductivity might be explained by cambial age. Additionally, results highlighted appreciable relationships between xylem traits and climate variability more than tree-ring width, supporting also the evidence that the plant hydraulic traits are closely linked to local climate extremes rather than average climatic conditions. We reported that the probability of extreme departure in specific hydraulic conductivity (Ks) rises at extreme values of Standardized Precipitation Index (SPI). Therefore, changing frequency or intensity of extreme events might overcome the adaptive limits of vascular transport, resulting in substantial reduction of hydraulic functionality and, hence increased incidence of xylem dysfunctions.
Rita, Angelo; Borghetti, Marco; Todaro, Luigi; Saracino, Antonio
2016-01-01
In the Mediterranean region, the widely predicted rise in temperature, change in the precipitation pattern, and increase in the frequency of extreme climatic events are expected to alter the shape of ecological communities and to affect plant physiological processes that regulate ecosystem functioning. Although change in the mean values are important, there is increasing evidence that plant distribution, survival, and productivity respond to extremes rather than to the average climatic condition. The present study aims to assess the effects of both mean and extreme climatic conditions on radial growth and functional anatomical traits using long-term tree-ring time series of two co-existing Quercus spp. from a drought-prone site in Southern Italy. In particular, this is the first attempt to apply the Generalized Additive Model for Location, Scale, and Shape (GAMLSS) technique and Bayesian modeling procedures to xylem traits data set, with the aim of (i) detecting non-linear long-term responses to climate and (ii) exploring relationships between climate extreme and xylem traits variability in terms of probability of occurrence. This study demonstrates the usefulness of long-term xylem trait chronologies as records of environmental conditions at annual resolution. Statistical analyses revealed that most of the variability in tree-ring width and specific hydraulic conductivity might be explained by cambial age. Additionally, results highlighted appreciable relationships between xylem traits and climate variability more than tree-ring width, supporting also the evidence that the plant hydraulic traits are closely linked to local climate extremes rather than average climatic conditions. We reported that the probability of extreme departure in specific hydraulic conductivity (Ks) rises at extreme values of Standardized Precipitation Index (SPI). Therefore, changing frequency or intensity of extreme events might overcome the adaptive limits of vascular transport, resulting in substantial reduction of hydraulic functionality and, hence increased incidence of xylem dysfunctions. PMID:27532008
Soper, Fiona M; Richards, Anna E; Siddique, Ilyas; Aidar, Marcos P M; Cook, Garry D; Hutley, Lindsay B; Robinson, Nicole; Schmidt, Susanne
2015-05-01
Water and nitrogen (N) interact to influence soil N cycling and plant N acquisition. We studied indices of soil N availability and acquisition by woody plant taxa with distinct nutritional specialisations along a north Australian rainfall gradient from monsoonal savanna (1,600-1,300 mm annual rainfall) to semi-arid woodland (600-250 mm). Aridity resulted in increased 'openness' of N cycling, indicated by increasing δ(15)N(soil) and nitrate:ammonium ratios, as plant communities transitioned from N to water limitation. In this context, we tested the hypothesis that δ(15)N(root) xylem sap provides a more direct measure of plant N acquisition than δ(15)N(foliage). We found highly variable offsets between δ(15)N(foliage) and δ(15)N(root) xylem sap, both between taxa at a single site (1.3-3.4 ‰) and within taxa across sites (0.8-3.4 ‰). As a result, δ(15)N(foliage) overlapped between N-fixing Acacia and non-fixing Eucalyptus/Corymbia and could not be used to reliably identify biological N fixation (BNF). However, Acacia δ(15)N(root) xylem sap indicated a decline in BNF with aridity corroborated by absence of root nodules and increasing xylem sap nitrate concentrations and consistent with shifting resource limitation. Acacia dominance at arid sites may be attributed to flexibility in N acquisition rather than BNF capacity. δ(15)N(root) xylem sap showed no evidence of shifting N acquisition in non-mycorrhizal Hakea/Grevillea and indicated only minor shifts in Eucalyptus/Corymbia consistent with enrichment of δ(15)N(soil) and/or decreasing mycorrhizal colonisation with aridity. We propose that δ(15)N(root) xylem sap is a more direct indicator of N source than δ(15)N(foliage), with calibration required before it could be applied to quantify BNF.
MacMillan, Colleen P; Birke, Hannah; Chuah, Aaron; Brill, Elizabeth; Tsuji, Yukiko; Ralph, John; Dennis, Elizabeth S; Llewellyn, Danny; Pettolino, Filomena A
2017-07-18
Knowledge of plant secondary cell wall (SCW) regulation and deposition is mainly based on the Arabidopsis model of a 'typical' lignocellulosic SCW. However, SCWs in other plants can vary from this. The SCW of mature cotton seed fibres is highly cellulosic and lacks lignification whereas xylem SCWs are lignocellulosic. We used cotton as a model to study different SCWs and the expression of the genes involved in their formation via RNA deep sequencing and chemical analysis of stem and seed fibre. Transcriptome comparisons from cotton xylem and pith as well as from a developmental series of seed fibres revealed tissue-specific and developmentally regulated expression of several NAC transcription factors some of which are likely to be important as top tier regulators of SCW formation in xylem and/or seed fibre. A so far undescribed hierarchy was identified between the top tier NAC transcription factors SND1-like and NST1/2 in cotton. Key SCW MYB transcription factors, homologs of Arabidopsis MYB46/83, were practically absent in cotton stem xylem. Lack of expression of other lignin-specific MYBs in seed fibre relative to xylem could account for the lack of lignin deposition in seed fibre. Expression of a MYB103 homolog correlated with temporal expression of SCW CesAs and cellulose synthesis in seed fibres. FLAs were highly expressed and may be important structural components of seed fibre SCWs. Finally, we made the unexpected observation that cell walls in the pith of cotton stems contained lignin and had a higher S:G ratio than in xylem, despite that tissue's lacking many of the gene transcripts normally associated with lignin biosynthesis. Our study in cotton confirmed some features of the currently accepted gene regulatory cascade for 'typical' plant SCWs, but also revealed substantial differences, especially with key downstream NACs and MYBs. The lignocellulosic SCW of cotton xylem appears to be achieved differently from that in Arabidopsis. Pith cell walls in cotton stems are compositionally very different from that reported for other plant species, including Arabidopsis. The current definition of a 'typical' primary or secondary cell wall might not be applicable to all cell types in all plant species.
Obudulu, Ogonna; Bygdell, Joakim; Sundberg, Björn; Moritz, Thomas; Hvidsten, Torgeir R; Trygg, Johan; Wingsle, Gunnar
2016-02-18
Wood development is of outstanding interest both to basic research and industry due to the associated cellulose and lignin biomass production. Efforts to elucidate wood formation (which is essential for numerous aspects of both pure and applied plant science) have been made using transcriptomic analyses and/or low-resolution sampling. However, transcriptomic data do not correlate perfectly with levels of expressed proteins due to effects of post-translational modifications and variations in turnover rates. In addition, high-resolution analysis is needed to characterize key transitions. In order to identify protein profiles across the developmental region of wood formation, an in-depth and tissue specific sampling was performed. We examined protein profiles, using an ultra-performance liquid chromatography/quadrupole time of flight mass spectrometry system, in high-resolution tangential sections spanning all wood development zones in Populus tremula from undifferentiated cambium to mature phloem and xylem, including cell expansion and cell death zones. In total, we analyzed 482 sections, 20-160 μm thick, from four 47-year-old trees growing wild in Sweden. We obtained high quality expression profiles for 3,082 proteins exhibiting consistency across the replicates, considering that the trees were growing in an uncontrolled environment. A combination of Principal Component Analysis (PCA), Orthogonal Projections to Latent Structures (OPLS) modeling and an enhanced stepwise linear modeling approach identified several major transitions in global protein expression profiles, pinpointing (for example) locations of the cambial division leading to phloem and xylem cells, and secondary cell wall formation zones. We also identified key proteins and associated pathways underlying these developmental landmarks. For example, many of the lignocellulosic related proteins were upregulated in the expansion to the early developmental xylem zone, and for laccases with a rapid decrease in early xylem zones. We observed upregulation of two forms of xylem cysteine protease (Potri.002G005700.1 and Potri.005G256000.2; Pt-XCP2.1) in early xylem and their downregulation in late maturing xylem. Our data also show that Pt-KOR1.3 (Potri.003G151700.2) exhibits an expression pattern that supports the hypothesis put forward in previous studies that this is a key xyloglucanase involved in cellulose biosynthesis in primary cell walls and reduction of cellulose crystallinity in secondary walls. Our novel multivariate approach highlights important processes and provides confirmatory insights into the molecular foundations of wood development.
Bennett, Gordon M.; Moran, Nancy A.
2013-01-01
Many insects rely on bacterial symbionts with tiny genomes specialized for provisioning nutrients lacking in host diets. Xylem sap and phloem sap are both deficient as insect diets, but differ dramatically in nutrient content, potentially affecting symbiont genome evolution. For sap-feeding insects, sequenced symbiont genomes are available only for phloem-feeding examples from the suborder Sternorrhyncha and xylem-feeding examples from the suborder Auchenorrhyncha, confounding comparisons. We sequenced genomes of the obligate symbionts, Sulcia muelleri and Nasuia deltocephalinicola, of the phloem-feeding pest insect, Macrosteles quadrilineatus (Auchenorrhyncha: Cicadellidae). Our results reveal that Nasuia-ALF has the smallest bacterial genome yet sequenced (112 kb), and that the Sulcia-ALF genome (190 kb) is smaller than that of Sulcia in other insect lineages. Together, these symbionts retain the capability to synthesize the 10 essential amino acids, as observed for several symbiont pairs from xylem-feeding Auchenorrhyncha. Nasuia retains genes enabling synthesis of two amino acids, DNA replication, transcription, and translation. Both symbionts have lost genes underlying ATP synthesis through oxidative phosphorylation, possibly as a consequence of the enriched sugar content of phloem. Shared genomic features, including reassignment of the UGA codon from Stop to tryptophan, and phylogenetic results suggest that Nasuia-ALF is most closely related to Zinderia, the betaproteobacterial symbiont of spittlebugs. Thus, Nasuia/Zinderia and Sulcia likely represent ancient associates that have co-resided in hosts since the divergence of leafhoppers and spittlebugs >200 Ma, and possibly since the origin of the Auchenorrhyncha, >260 Ma. PMID:23918810
Differential expansion and expression of alpha- and beta-tubulin gene families in Populus.
Oakley, Rodney V; Wang, Yuh-Shuh; Ramakrishna, Wusirika; Harding, Scott A; Tsai, Chung-Jui
2007-11-01
Microtubule organization is intimately associated with cellulose microfibril deposition, central to plant secondary cell wall development. We have determined that a relatively large suite of eight alpha-TUBULIN (TUA) and 20 beta-TUBULIN (TUB) genes is expressed in the woody perennial Populus. A number of features, including gene number, alpha:beta gene representation, amino acid changes at the C terminus, and transcript abundance in wood-forming tissue, distinguish the Populus tubulin suite from that of Arabidopsis thaliana. Five of the eight Populus TUAs are unusual in that they contain a C-terminal methionine, glutamic acid, or glutamine, instead of the more typical, and potentially regulatory, C-terminal tyrosine. Both C-terminal Y-type (TUA1) and M-type (TUA5) TUAs were highly expressed in wood-forming tissues and pollen, while the Y-type TUA6 and TUA8 were abundant only in pollen. Transcripts of the disproportionately expanded TUB family were present at comparatively low levels, with phylogenetically distinct classes predominating in xylem and pollen. When tension wood induction was used as a model system to examine changes in tubulin gene expression under conditions of augmented cellulose deposition, xylem-abundant TUA and TUB genes were up-regulated. Immunolocalization of TUA and TUB in xylem and phloem fibers of stems further supported the notion of heavy microtubule involvement during cellulose microfibril deposition in secondary walls. The high degree of sequence diversity, differential expansion, and differential regulation of Populus TUA and TUB families may confer flexibility in cell wall formation that is of adaptive significance to the woody perennial growth habit.
Characterization of microRNAs Expressed during Secondary Wall Biosynthesis in Acacia mangium
Ong, Seong Siang; Wickneswari, Ratnam
2012-01-01
MicroRNAs (miRNAs) play critical regulatory roles by acting as sequence specific guide during secondary wall formation in woody and non-woody species. Although thousands of plant miRNAs have been sequenced, there is no comprehensive view of miRNA mediated gene regulatory network to provide profound biological insights into the regulation of xylem development. Herein, we report the involvement of six highly conserved amg-miRNA families (amg-miR166, amg-miR172, amg-miR168, amg-miR159, amg-miR394, and amg-miR156) as the potential regulatory sequences of secondary cell wall biosynthesis. Within this highly conserved amg-miRNA family, only amg-miR166 exhibited strong differences in expression between phloem and xylem tissue. The functional characterization of amg-miR166 targets in various tissues revealed three groups of HD-ZIP III: ATHB8, ATHB15, and REVOLUTA which play pivotal roles in xylem development. Although these three groups vary in their functions, -psRNA target analysis indicated that miRNA target sequences of the nine different members of HD-ZIP III are always conserved. We found that precursor structures of amg-miR166 undergo exhaustive sequence variation even within members of the same family. Gene expression analysis showed three key lignin pathway genes: C4H, CAD, and CCoAOMT were upregulated in compression wood where a cascade of miRNAs was downregulated. This study offers a comprehensive analysis on the involvement of highly conserved miRNAs implicated in the secondary wall formation of woody plants. PMID:23251324
Fabrication of Artificial Leaf to Develop Fluid Pump Driven by Surface Tension and Evaporation
NASA Astrophysics Data System (ADS)
Lee, Minki; Lim, Hosub; Lee, Jinkee
2017-11-01
Plants transport water from roots to leaves via xylem through transpiration, which is an evaporation process that occurs at the leaves. During transpiration, negative pressure can be generated by the porous structure of mesophyll cells in the leaves. Here, an artificial leaf mimicking structure using hydrogel, which has a nanoporous structure is fabricated. The cryogel method is used to develop a hierarchy structure on the nano- and microscale in the hydrogel media that is similar to the mesophyll cells and veins of a leaf, respectively. The theoretical model is analyzed to calculate the flow resistance in the artificial leaf, and compare the model with the experimental results. The experiment involves connecting a glass capillary tube at the bottom of the artificial leaf to observe the fluid velocity in the glass capillary tube generated by the negative pressure. The use of silicone oil as fluid instead of water to increase the flow resistance enables the measurement of negative pressure. The negative pressure of the artificial leaf is affected by several variables (e.g., pore size, wettability of the structure). Finally, by decreasing the pore size and increasing the wettability, the maximum negative pressure of the artificial leaf, -7.9 kPa is obtained.
DeFalco, L.A.; Detling, J.K.; Tracy, C.R.; Warren, S.D.
2001-01-01
Microbiotic crusts are important components of many aridland soils. Research on crusts typically focuses on the increase in soil fertility due to N-fixing micro-organisms, the stabilization of soils against water and wind erosion and the impact of disturbance on N-cycling. The effect of microbiotic crusts on the associated plant community has received little attention. We quantified the influence of crusts on the production, species diversity, nutrient content and water relations of winter annual plant species associated with microbiotic soil crusts in the northeast Mojave Desert. Shoot biomass of winter annuals was 37% greater and plant density was 77% greater on crusts than were biomass and density on soils lacking crust cover (=bare soils). This greater production of annuals on crusts was likely due to enhanced soil conditions including an almost two-fold increase in soil organic matter and inorganic N compared to bare soils. Crusted soils also had 53% greater volumetric water content than bare soils during November and December, the time when winter annuals become established. As plant development progressed into spring, however, soil water availability decreased: More negative plant xylem water potentials were associated with greater plant biomass on crusted soils. Plants associated with microbiotic soil crusts had lower concentrations of N in shoots (mg N g-1 dry mass). However, total shoot N (mg N m-2) was the same in plants growing on the different soil types when biomass production peaked in April. Shoots had similar patterns in their concentration and content of P. Species diversity of annuals was not statistically different between the two soil types. Yet, while native annuals comprised the greatest proportion of shoot biomass on bare soils, exotic forbs and grasses produced more biomass on crusts. Total shoot nutrient content (biomass x concentration) of the two exotic annual species examined was dramatically greater on crusts than bare soils; only one native species had greater shoot content of N and P when growing on crusts than bare soils. Microbiotic crusts appear to increase site fertility in the northeast Mojave Desert, but nutrients and water distributed within a greater biomass of annual plants growing on microbiotic crusts likely resulted in lower concentrations of nutrients in plant tissue and lower xylem pressure potentials than plants growing on bare soils. Exotic annuals growing on crusts appear to respond to the higher N availability by growing faster, potentially outcompeting native annual species.
DeFalco, Lesley; Detling, James K.; Tracy, C. Richard; Warren, Steven D.
2001-01-01
Microbiotic crusts are important components of many aridland soils. Research on crusts typically focuses on the increase in soil fertility due to N-fixing micro-organisms, the stabilization of soils against water and wind erosion and the impact of disturbance on N-cycling. The effect of microbiotic crusts on the associated plant community has received little attention. We quantified the influence of crusts on the production, species diversity, nutrient content and water relations of winter annual plant species associated with microbiotic soil crusts in the northeast Mojave Desert. Shoot biomass of winter annuals was 37% greater and plant density was 77% greater on crusts than were biomass and density on soils lacking crust cover (=bare soils). This greater production of annuals on crusts was likely due to enhanced soil conditions including an almost two-fold increase in soil organic matter and inorganic N compared to bare soils. Crusted soils also had 53% greater volumetric water content than bare soils during November and December, the time when winter annuals become established. As plant development progressed into spring, however, soil water availability decreased: More negative plant xylem water potentials were associated with greater plant biomass on crusted soils. Plants associated with microbiotic soil crusts had lower concentrations of N in shoots (mg N g−1 dry mass). However, total shoot N (mg N m−2) was the same in plants growing on the different soil types when biomass production peaked in April. Shoots had similar patterns in their concentration and content of P. Species diversity of annuals was not statistically different between the two soil types. Yet, while native annuals comprised the greatest proportion of shoot biomass on bare soils, exotic forbs and grasses produced more biomass on crusts. Total shoot nutrient content (biomass×concentration) of the two exotic annual species examined was dramatically greater on crusts than bare soils; only one native species had greater shoot content of N and P when growing on crusts than bare soils. Microbiotic crusts appear to increase site fertility in the northeast Mojave Desert, but nutrients and water distributed within a greater biomass of annual plants growing on microbiotic crusts likely resulted in lower concentrations of nutrients in plant tissue and lower xylem pressure potentials than plants growing on bare soils. Exotic annuals growing on crusts appear to respond to the higher N availability by growing faster, potentially outcompeting native annual species.
Investigating water transport through the xylem network in vascular plants.
Kim, Hae Koo; Park, Joonghyuk; Hwang, Ildoo
2014-04-01
Our understanding of physical and physiological mechanisms depends on the development of advanced technologies and tools to prove or re-evaluate established theories, and test new hypotheses. Water flow in land plants is a fascinating phenomenon, a vital component of the water cycle, and essential for life on Earth. The cohesion-tension theory (CTT), formulated more than a century ago and based on the physical properties of water, laid the foundation for our understanding of water transport in vascular plants. Numerous experimental tools have since been developed to evaluate various aspects of the CTT, such as the existence of negative hydrostatic pressure. This review focuses on the evolution of the experimental methods used to study water transport in plants, and summarizes the different ways to investigate the diversity of the xylem network structure and sap flow dynamics in various species. As water transport is documented at different scales, from the level of single conduits to entire plants, it is critical that new results be subjected to systematic cross-validation and that findings based on different organs be integrated at the whole-plant level. We also discuss the functional trade-offs between optimizing hydraulic efficiency and maintaining the safety of the entire transport system. Furthermore, we evaluate future directions in sap flow research and highlight the importance of integrating the combined effects of various levels of hydraulic regulation.
Hochberg, Uri; Albuquerque, Caetano; Rachmilevitch, Shimon; Cochard, Herve; David-Schwartz, Rakefet; Brodersen, Craig R; McElrone, Andrew; Windt, Carel W
2016-09-01
The 'hydraulic vulnerability segmentation' hypothesis predicts that expendable distal organs are more susceptible to water stress-induced embolism than the main stem of the plant. In the current work, we present the first in vivo visualization of this phenomenon. In two separate experiments, using magnetic resonance imaging or synchrotron-based microcomputed tomography, grapevines (Vitis vinifera) were dehydrated while simultaneously scanning the main stems and petioles for the occurrence of emboli at different xylem pressures (Ψx ). Magnetic resonance imaging revealed that 50% of the conductive xylem area of the petioles was embolized at a Ψx of -1.54 MPa, whereas the stems did not reach similar losses until -1.9 MPa. Microcomputed tomography confirmed these findings, showing that approximately half the vessels in the petioles were embolized at a Ψx of -1.6 MPa, whereas only few were embolized in the stems. Petioles were shown to be more resistant to water stress-induced embolism than previously measured with invasive hydraulic methods. The results provide the first direct evidence for the hydraulic vulnerability segmentation hypothesis and highlight its importance in grapevine responses to severe water stress. Additionally, these data suggest that air entry through the petiole into the stem is unlikely in grapevines during drought. © 2015 John Wiley & Sons Ltd.
Barnard, David M; Meinzer, Frederick C; Lachenbruch, Barbara; McCulloh, Katherine A; Johnson, Daniel M; Woodruff, David R
2011-04-01
In the Pacific north-west, the Cascade Mountain Range blocks much of the precipitation and maritime influence of the Pacific Ocean, resulting in distinct climates east and west of the mountains. The current study aimed to investigate relationships between water storage and transport properties in populations of Douglas-fir (Pseudotsuga menziesii) and ponderosa pine (Pinus ponderosa) adapted to both climates. Sapwood thickness, capacitance, vulnerability to embolism, and axial and radial conductivity were measured on samples collected from trunks of mature trees. The sapwood of ponderosa pine was three to four times thicker than Douglas-fir. Radial conductivity was higher in west-side populations of both species, but axial conductivity was higher in the east-side populations and in Douglas-fir. Eastern populations of both species had sapwood that was more vulnerable to embolism than west-side populations. Sapwood capacitance was similar between species, but was about twice as great in east-side populations (580 kg m⁻³ MPa⁻¹) as in west-side populations (274 kg m⁻³ MPa⁻¹). Capacitance was positively correlated with both mean embolism pressure and axial conductivity across species and populations, suggesting that coordinated adjustments in xylem efficiency, safety and water storage capacity may serve to avoid embolism along a gradient of increasing aridity. © 2011 Blackwell Publishing Ltd.
Christensen-Dalsgaard, Karen K; Tyree, Melvin T
2013-11-01
Frost damage to the xylem conduits of trees is a phenomenon of eco-physiological importance. It is often documented in terms of the percentage loss of conductivity (PLC), an indicator of air filling of the conduits. However, trees that refill their conduits in spring could be impacted more by damage to the conduits that reduce cavitation resistance, making them more susceptible to future drought events. We investigated whether ice formation, dynamic flexing of frozen branches or freeze-thaw events could reduce the cavitation resistance (cause "frost fatigue") in first-year shoots of apple (Malus domestica) and clonal hybrid cottonwood (Walker). Frost fatigue was measured in terms of P50 (the negative xylem pressure required to cause a 50 % loss of conductivity). All treatment groups showed significant frost fatigue, with the exception of the pre-flushed, constantly frozen poplar branches. The P50 following freeze treatments was approximately 50 % of the pre-freeze values. The effect tended to be greater in freeze-thawed branches. Dynamic bending of the branches had no effect on either PLC or P50. In three out of four cases, there was a significant correlation between P50 and PLC. Frost fatigue occurred in both apple and poplar, two unrelated species with different drought and frost tolerances, suggesting that it may be a widespread phenomenon that could impact the ecophysiology of temperate forests.
Duan, Honglang; Duursma, Remko A; Huang, Guomin; Smith, Renee A; Choat, Brendan; O'Grady, Anthony P; Tissue, David T
2014-07-01
It has been reported that elevated temperature accelerates the time-to-mortality in plants exposed to prolonged drought, while elevated [CO(2)] acts as a mitigating factor because it can reduce stomatal conductance and thereby reduce water loss. We examined the interactive effects of elevated [CO(2)] and temperature on the inter-dependent carbon and hydraulic characteristics associated with drought-induced mortality in Eucalyptus radiata seedlings grown in two [CO(2)] (400 and 640 μL L(-1)) and two temperature (ambient and ambient +4 °C) treatments. Seedlings were exposed to two controlled drying and rewatering cycles, and then water was withheld until plants died. The extent of xylem cavitation was assessed as loss of stem hydraulic conductivity. Elevated temperature triggered more rapid mortality than ambient temperature through hydraulic failure, and was associated with larger water use, increased drought sensitivities of gas exchange traits and earlier occurrence of xylem cavitation. Elevated [CO(2)] had a negligible effect on seedling response to drought, and did not ameliorate the negative effects of elevated temperature on drought. Our findings suggest that elevated temperature and consequent higher vapour pressure deficit, but not elevated [CO(2)], may be the primary contributors to drought-induced seedling mortality under future climates. © 2013 John Wiley & Sons Ltd.
Magnetic resonance imaging of water ascent in embolized xylem vessels of grapevine stem segments
Mingtao Wang; Melvin T. Tyree; Roderick E. Wasylishen
2013-01-01
Temporal and spatial information about water refilling of embolized xylem vessels and the rate of water ascent in these vessels is critical for understanding embolism repair in intact living vascular plants. High-resolution 1H magnetic resonance imaging (MRI) experiments have been performed on embolized grapevine stem segments while they were...
Soil water and xylem chemistry in declining sugar maple stands in Pennsylvania
David R. DeWalle; Bryan R. Swistock; William E. Sharpe
1999-01-01
Evidence is accumulating that decline of sugar maple, Acer saccharum Marsh., in northern Pennsylvania may be related to overall site fertility as reflected in the chemistry of soil water and bolewood xylem. In this paper we discuss factors related to varying site fertility, including effects of soil liming, past glacialion, topographic position and...
Steven L. Voelker; Barbara Lachenbruch; Frederick C. Meinzer; Peter Kitin; Steven H. Strauss
2011-01-01
We studied xylem anatomy and hydraulic architecture in 14 transgenic insertion events and a control line of hybrid poplar (Populus spp.) that varied in lignin content. Transgenic events had different levels of down-regulation of two genes encoding 4-coumarate:coenzyme A ligase (4CL). Two-year-old trees were characterized after...
6-8-dihydroxy-3-hydroxymethyl-isocoumarin, and other phenolic metabolites of Ceratocystis minor
G.W. McGraw; R.W. Hemingway
1977-01-01
The fungus Ceratocytis minor (Hedge)Hunt is generally introduced into the phloem and xylem of southern pine trees during attack by the southern pine beetle Dendrotonus frontalis Zimmerman, and development of C. minor in the xylem is considered to be central to the death of beetle-attacked trees [1]. There are...
Spatial organization of xylem cell walls by ROP GTPases and microtubule-associated proteins.
Oda, Yoshihisa; Fukuda, Hiroo
2013-12-01
Proper patterning of cellulosic cell walls is critical for cell shaping and differentiation of plant cells. Cortical microtubule arrays regulate the deposition patterns of cellulose microfibrils by controlling the targeting and trajectory of cellulose synthase complexes. Although some microtubule-associated proteins (MAPs) regulate the arrangement of cortical microtubules, knowledge about the overall mechanism governing the spacing of cortical microtubules is still limited. Recent studies reveal that ROP GTPases and MAPs spatially regulate the assembly and disassembly of cortical microtubules in developing xylem cells, in which localized secondary cell walls are deposited. Here, we review recent insights into the regulation of xylem cell wall patterning by cortical microtubules, ROP GTPases, and MAPs. Copyright © 2013 Elsevier Ltd. All rights reserved.
Touati, Mostefa; Knipfer, Thorsten; Visnovitz, Tamás; Kameli, Abdelkrim; Fricke, Wieland
2015-07-01
The aim of the present study was to assess the mechanical and hydraulic limitation of growth in leaf epidermal cells of barley (Hordeum vulgare L.) in response to agents which affect cellular water (mercuric chloride, HgCl(2)) and potassium (cesium chloride, CsCl; tetraethylammonium, TEA) transport, pump activity of plasma membrane H(+)-ATPase and wall acidification (fusicoccin, FC). Cell turgor (P) was measured with the cell pressure probe, and cell osmotic pressure (π) was analyzed through picoliter osmometry of single-cell extracts. A wall extensibility coefficient (M) and tissue hydraulic conductance coefficient (L) were derived using the Lockhart equation. There was a significant positive linear relationship between relative elemental growth rate and P, which fit all treatments, with an overall apparent yield threshold of 0.368 MPa. Differences in growth between treatments could be explained through differences in P. A comparison of L and M showed that growth in all except the FC treatment was co-limited through hydraulic and mechanical properties, though to various extents. This was accompanied by significant (0.17-0.24 MPa) differences in water potential (ΔΨ) between xylem and epidermal cells in the leaf elongation zone. In contrast, FC-treated leaves showed ΔΨ close to zero and a 10-fold increase in L. © The Author 2015. Published by Oxford University Press on behalf of Japanese Society of Plant Physiologists. All rights reserved. For permissions, please email: journals.permissions@oup.com.
Limitations on gas exchange recovery following natural drought in Californian oak woodlands.
NASA Astrophysics Data System (ADS)
Ackerly, D.; Skelton, R. P.; Dawson, T.; Thompson, S.; Feng, X.; Weitz, A.; McLaughlin, B.
2017-12-01
Abstract Background/Question/Methods Drought can cause major damage to plant communities, but species damage thresholds and post-drought recovery of forest productivity are not yet predictable. We asked the question how should forest net primary productivity recover following exposure to severe drought? We used a natural drought period to investigate whether drought responses and post-drought recovery of canopy health could be predicted by properties of the water transport system. We aimed to test the hypothesis that recovery of gas exchange and canopy health would be most severely limited by xylem embolism in stems. To do this we monitored leaf level gas exchange and water status for multiple individuals of two deciduous and two evergreen species for four years spanning a severe drought event and following subsequent rehydration. Results/Discussion Severe drought caused major declines in leaf water potential, reduced stomatal conductance and assimilation rates and increased canopy bareness in our four canopy species. Water potential surpassed levels associated with incipient embolism in leaves of most trees. In contrast, due to hydraulic segmentation, water potential only rarely surpassed critical thresholds in the stems of the study trees. Individuals that surpassed critical thresholds of embolism in the stem displayed significant canopy dieback and mortality. Thus, recovery of plant gas exchange and canopy health was predicted by xylem safety margin in stems, but not leaves, providing strong support for stem cavitation vulnerability as an index of damage under natural drought conditions.
Liu, Jinyu; Tyree, Melvin T.
2015-01-01
Cavitation of water in xylem vessels followed by embolism formation has been authenticated for more than 40 years. Embolism formation involves the gradual buildup of bubble pressure (air) to atmospheric pressure as demanded by Henry’s law of equilibrium between gaseous and liquid phases. However, the tempo of pressure increase has not been quantified. In this report, we show that the rate of pressurization of embolized vessels is controlled by both fast and slow kinetics, where both tempos are controlled by diffusion but over different spatial scales. The fast tempo involves a localized diffusion from endogenous sources: over a distance of about 0.05 mm from water-filled wood to the nearest embolized vessels; this process, in theory, should take <2 min. The slow tempo involves diffusion of air from exogenous sources (outside the stem). The latter diffusion process is slower because of the increased distance of diffusion of up to 4 mm. Radial diffusion models and experimental measurements both confirm that the average time constant is >17 h, with complete equilibrium requiring 1 to 2 d. The implications of these timescales for the standard methods of measuring percentage loss of hydraulic conductivity are discussed in theory and deserve more research in future. PMID:26468516
Wang, Yujie; Liu, Jinyu; Tyree, Melvin T
2015-12-01
Cavitation of water in xylem vessels followed by embolism formation has been authenticated for more than 40 years. Embolism formation involves the gradual buildup of bubble pressure (air) to atmospheric pressure as demanded by Henry's law of equilibrium between gaseous and liquid phases. However, the tempo of pressure increase has not been quantified. In this report, we show that the rate of pressurization of embolized vessels is controlled by both fast and slow kinetics, where both tempos are controlled by diffusion but over different spatial scales. The fast tempo involves a localized diffusion from endogenous sources: over a distance of about 0.05 mm from water-filled wood to the nearest embolized vessels; this process, in theory, should take <2 min. The slow tempo involves diffusion of air from exogenous sources (outside the stem). The latter diffusion process is slower because of the increased distance of diffusion of up to 4 mm. Radial diffusion models and experimental measurements both confirm that the average time constant is >17 h, with complete equilibrium requiring 1 to 2 d. The implications of these timescales for the standard methods of measuring percentage loss of hydraulic conductivity are discussed in theory and deserve more research in future. © 2015 American Society of Plant Biologists. All Rights Reserved.
Hester, M.W.; Mendelssohn, I.A.; McKee, K.L.
2001-01-01
Panicum hemitomon, Spartina patens, and Spartina alterniflora are wide-spread dominant grasses of fresh, brackish, and salt marsh plant communities, respectively. Our previous research identified significant intraspecific variation in salt tolerance and morphology among populations within each species. In this study our objectives were to determine shorter-term physiological/biochemical responses to salinity stress and identify potential indicators of salt tolerance, with the ultimate goal of discerning similarities and differences in the mechanisms of salinity stress resistance. We subjected a subset of six populations within each species, ranging from high to low salt tolerance, to sublethal salinity levels (4, 20, and 30 ppt, respectively, for species) and monitored physiological and growth responses after 1 week (early harvest) and 5 weeks (late harvest). In all three species sublethal salinity levels generally resulted in significantly reduced net CO2 assimilation, leaf expansion, midday leaf xylem pressure, water use efficiency, and live and total biomass; and significantly increased leaf Na+/K+ ratio, leaf proline, leaf glycine betaine, leaf sucrose, root-to-shoot ratio, and dead:total aboveground biomass ratio. All three species displayed significant population (intraspecific) variation in net CO2 assimilation, leaf expansion, water use efficiency, midday leaf xylem pressure, leaf proline, leaf glycine betaine (except Panicum, where it could not be accurately determined), leaf Na+/K+ ratio, leaf sucrose, total plant biomass, dead:total aboveground biomass ratio, and root-to-shoot ratio. General indicators of salt tolerance (regardless of species) included high net CO2 assimilation rates and water use efficiencies, and low ratios of root-to-shoot and dead:total aboveground biomass. Factor analysis and a-priori linear contrasts revealed some unique differences between species in terms of the relative importance of morphology and physiology in explaining intraspecific variation in salt tolerance. Plant morphology (size attributes) were strongly associated with salt tolerance in P. hemitomon, weakly associated with salt tolerance in S. patens, and not associated with salt tolerance in S. alterniflora. Highly salt-tolerant populations of Spartina alterniflora displayed the greatest ion selectivity (lower leaf Na+/K+ ratios), which was not displayed by the other two species. These results suggest that plant size attributes can be very important in explaining population differences in salt tolerance in glycophytes, but may be independent of salt tolerance in halophytes, which have specialized physiological (and/or anatomical) adaptations that can confer salinity stress resistance through mechanisms such as selective ion exclusion and secretion. ?? 2001 Elsevier Science B.V. All rights reserved.
Castagneri, Daniele; Battipaglia, Giovanna; von Arx, Georg; Pacheco, Arturo; Carrer, Marco
2018-04-24
Understanding how climate affects xylem formation is critical for predicting the impact of future conditions on tree growth and functioning in the Mediterranean region, which is expected to face warmer and drier conditions. However, mechanisms of growth response to climate at different temporal scales are still largely unknown, being complicated by separation between spring and autumn xylogenesis (bimodal temporal pattern) in most species such as Mediterranean pines. We investigated wood anatomical characteristics and carbon stable isotope composition in Mediterranean Pinus pinea L. along tree-ring series at intra-ring resolution to assess xylem formation processes and responses to intra-annual climate variability. Xylem anatomy was strongly related to environmental conditions occurring a few months before and during the growing season, but was not affected by summer drought. In particular, the lumen diameter of the first earlywood tracheids was related to winter precipitation, whereas the size of tracheids produced later was influenced by mid-spring precipitation. Diameter of latewood tracheids was associated with precipitation in mid-autumn. In contrast, tree-ring carbon isotope composition was mostly related to climate of the previous seasons. Earlywood was likely formed using both recently and formerly assimilated carbon, while latewood relied mostly on carbon accumulated many months prior to its formation. Our integrated approach provided new evidence on the short-term and carry-over effects of climate on the bimodal temporal xylem formation in P. pinea. Investigations on different variables and time scales are necessary to disentangle the complex climate influence on tree growth processes under Mediterranean conditions.
Metal Complexation in Xylem Fluid 1
White, Michael C.; Decker, A. Morris; Chaney, Rufus L.
1981-01-01
Xylem fluid was analyzed for numerous solutes to characterize chemically the sap as a medium for forming and transporting metal complexes. The stem exudate was collected hourly for 8 hours from topped 31-day-old soybean (Glycine max L. Merr.) and 46-day-old tomato (Lycopersicon esculentum Mill.) plants grown in normal (0.5 micromolar) and Za-phytotoxic nutrient solutions. Soybean plants were grown in the normal and high-Zn solutions for 24 days; tomato plants were grown for 32 days. The exudate was analyzed for seven organic acids, 22 amino acids, eight inorganic solutes, apparent ionic strength, and pH. Significant changes in many solutes occurred over the 8-hour sampling period. These fluctuations depended on plant species, individual solute, and Zn treatment, and demonstrated that extrapolation of xylem-fluid analyses to whole-plant xylem sap is valid only for sap samples collected shortly after topping a plant. Exudate pH decreased over the 8-hour period for both species; exudate ionic strength increased for tomato and decreased for soybean. At the normal-Zn treatment (0 to 1 hour), the highest acid micromolar concentrations in soybean exudate were: asparagine, 2,583; citric, 1,706; malic, 890; and malonic, 264. Under the same conditions, the highest acid micromolar concentrations in tomato exudate were: maleic, 1,206; malic, 628; glutamine, 522; citric, 301; and asparagine, 242. Cysteine and methionine were above detection limits only in soybean exudate. Zinc phytotoxicity caused significant changes in many solutes. The analyses reported here provide a comprehensive data base for further studies on metal-complex equilibria in xylem fluid. PMID:16661664
DOE Office of Scientific and Technical Information (OSTI.GOV)
Fred J. Molz, III
To better understand longer-term vadose zone transport in southeastern soils, field lysimeter experiments were conducted at the Savannah River Site (SRS) near Aiken, SC, in the 1980s. Each of the three lysimeters analyzed herein contained a filter paper spiked with different Pu solutions, and they were left exposed to natural environmental conditions (including the growth of annual weed grasses) for 11 years. The resulting Pu activity measurements from each lysimeter core showed anomalous activity distributions below the source, with significant migration of Pu above the source. Such results are not explainable by adsorption phenomena alone. A transient variably saturated flowmore » model with root water uptake was developed and coupled to a soil reactive transport model. Somewhat surprisingly, the fully transient analysis showed results nearly identical to those of a much simpler steady flow analysis performed previously. However, all phenomena studied were unable to produce the upward Pu transport observed in the data. This result suggests another transport mechanism such as Pu uptake by roots and upward transport due to transpiration. Thus, the variably saturated flow and reactive transport model was extended to include uptake and transport of Pu within the root xylem, along with computational methodology and results. In the extended model, flow velocity in the soil was driven by precipitation input along with transpiration and drainage. Water uptake by the roots determined the flow velocity in the root xylem, and this along with uptake of Pu in the transpiration stream drove advection and dispersion of the two Pu species in the xylem. During wet periods with high potential evapotranspiration, maximum flow velocities through the xylem would approached 600 cm/hr, orders of magnitude larger that flow velocities in the soil. Values for parameters and the correct conceptual viewpoint for Pu transport in plant xylem was uncertain. This motivated further experiments devoted to Pu uptake by corn roots and xylem transport. Plants were started in wet paper wrapped around each corn seed. When the tap roots were sufficiently long, the seedlings were transplanted to a soil container with the tap root extending out the container bottom. The soil container was then placed over a nutrient solution container, and the solution served as an additional medium for root growth. To conduct an uptake study, a radioactive substance, such as Pu complexed with the bacterial siderophore DFOB, was added to the nutrient solution. After a suitable elapsed time, the corn plant was sacrificed, cut into 10 cm lengths, and the activity distribution measured. Experimental results clarified the basic nature of Pu uptake and transport in corn plants, and resulting simulations suggested that each growing season Pu in the SRS lysimeters would move into the plant shoots and be deposited on the soil surface during the Fall dieback. Subsequent isotope ratio analyses showed that this did happen. OVERALL RESULTS AND CONCLUSIONS - (1) Pu transport downward from the source is controlled by advection, dispersion and adsorption, along with surface-mediated REDOX reactions. (2) Hysteresis, extreme root distribution functions, air-content dependent oxidation rate constants, and large evaporation rates from the soil surface were not able to explain the observed upward migration of Pu. (3) Small amounts of Pu uptake by plant roots and translocation in the transpiration stream creates a realistic mechanism for upward Pu migration (4) Realistic xylem cross-sectional areas imply high flow velocities under hot, wet conditions. Such flow velocities produce the correct shape for the observed activity distributions in the top 20 cm of the lysimeter soil. (5) Simulations imply that Pu should have moved into the above-ground grass tissue each year during the duration of the experiments, resulting in an activity residual accumulating on the soil surface. An isotope ratio analysis showed that the observed surface Pu residue was from the buried sources, not atmospheric fallout. (6) The plant experiments indicate a Pu-DFOB velocity in the corn xylem of at least 174 cm/hr, much higher than ionic Pu in soil. Thus, Pu complexation with chelating agents is probably what led to the observed enhanced uptake and mobility in grasses. (7) Plant experiments show that the uptake of Fe-DFOB, Pu-DFOB and the resulting distributions are very similar. This supports the hypothesis that plant and bacterial iron-seeking chemistry mistakes Pu for Fe.« less
USDA-ARS?s Scientific Manuscript database
The aim of this work was to study the effects of Fe and Mn deficiencies on the xylem sap proteome of tomato using a shotgun proteomic approach, with the final goal of elucidating plant response mechanisms to these stresses. This approach yielded 643 proteins reliably identified and quantified with 7...
Structure-function relationships in sapwood water transport and storage.
Barbara L. Gartner; Frederick C. Meinzer
2005-01-01
Primary production by plants requires the loss of substantial quantities of water when the stomata are open for carbon assimilation. The delivery of that water to the leaves occurs through the xylem. The structure, condition, and quantity of the xylem control not only the transport efficiency but also the release of water from storage. For example, if there is high...