The accumulation pattern of ferruginol in the heartwood-forming Cryptomeria japonica xylem as determined by time-of-flight secondary ion mass spectrometry and quantity analysis

PubMed Central

Kuroda, Katsushi; Fujiwara, Takeshi; Hashida, Koh; Imai, Takanori; Kushi, Masayoshi; Saito, Kaori; Fukushima, Kazuhiko

2014-01-01

Background and Aims Heartwood formation is a unique phenomenon of tree species. Although the accumulation of heartwood substances is a well-known feature of the process, the accumulation mechanism remains unclear. The aim of this study was to determine the accumulation process of ferruginol, a predominant heartwood substance of Cryptomeria japonica, in heartwood-forming xylem. Methods The radial accumulation pattern of ferruginol was examined from sapwood and through the intermediate wood to the heartwood by direct mapping using time-of-flight secondary ion mass spectrometry (TOF-SIMS). The data were compared with quantitative results obtained from a novel method of gas chromatography analysis using laser microdissection sampling and with water distribution obtained from cryo-scanning electron microscopy. Key Results Ferruginol initially accumulated in the middle of the intermediate wood, in the earlywood near the annual ring boundary. It accumulated throughout the entire earlywood in the inner intermediate wood, and in both the earlywood and the latewood in the heartwood. The process of ferruginol accumulation continued for more than eight annual rings. Ferruginol concentration peaked at the border between the intermediate wood and heartwood, while the concentration was less in the latewood compared wiht the earlywood in each annual ring. Ferruginol tended to accumulate around the ray parenchyma cells. In addition, at the border between the intermediate wood and heartwood, the accumulation was higher in areas without water than in areas with water. Conclusions TOF-SIMS clearly revealed ferruginol distribution at the cellular level. Ferruginol accumulation begins in the middle of intermediate wood, initially in the earlywood near the annual ring boundary, then throughout the entire earlywood, and finally across to the whole annual ring in the heartwood. The heterogeneous timing of ferruginol accumulation could be related to the distribution of ray parenchyma cells

Fibril angle of loblolly pine wood as related to specific gravity, growth rate, and distance from pith

Treesearch

Charles W. McMillin

1972-01-01

Fibril angles were greater for earlywood (avg. 33.4o) than for latewood tracheids (avg. 26.9o). For earlywood, fibril angle did not differ between growth rates when the specific gravity was low (avg. 33.3o). When the specific gravity was high, wood of fast growth had a higher fibril angle (avg. 35.1...

Microtensile strength of spruce pine after exposure to acids and bases

Treesearch

Floyd G. Manwiller; Paul R. Godfrey

1972-01-01

Earlywood and latewood microtensile specimens from 12 trees of Pinus glabra Wal. were subjected to 10-percent solution of 5 acids and 3 bases at 90oC for up to 3 hours. Hydrochloric and sulfuric acids were the most damaging, lowering maximum tensile strength 27 and 17 percent in earlywood and 36 and 39 percent in latewood; they...

Microtensile strength of spruce pine after exposure to acids and bases

Treesearch

F.G. Manwiller; P.R. Godfrey

1973-01-01

Earlywood and latewood microtensile specimens from 12 trees of Pinus glabra Walt. were subjected to 10-percent solution of 5 acids and 3 bases at good for up to 3 hours. Hydrochloric and sulfuric acids were the most damaging, lowering maximum tensile strength 27 and 17 percent in earlywood and 36 and 39 percent in latewood; they reduced work to maximum load 40 percent...

Fibril angle of loblolly pine wood as related to specific gravity, growth rate, and distance from pith

Treesearch

C.W. McMillin

1973-01-01

Fibril angles were greater for earlywood (avg. 33.4°) than for latewood tracheida (avg. 26.9°). For earlywood, fibril angle did not differ between growth rates when the specific gravity was low (avg. 33.3°). When the specific gravity was high, wood of fast growth had a higher fibril angle (avg. 35.1.°) than wood of slow growth (avg. 32.0°). No differences were detected...

Elevated temperature and CO(2) concentration effects on xylem anatomy of Scots pine.

PubMed

Kilpeläinen, Antti; Gerendiain, Ane Zubizarreta; Luostarinen, Katri; Peltola, Heli; Kellomäki, Seppo

2007-09-01

We studied the effects of elevated temperature and carbon dioxide concentration ([CO(2)]) alone and together on wood anatomy of 20-year-old Scots pine (Pinus sylvestris L.) trees. The study was conducted in 16 closed chambers, providing a factorial combination of two temperature regimes and two CO(2) concentrations (ambient and elevated), with four trees in each treatment. The climate scenario included a doubling of [CO(2)] and a corresponding increase of 2-6 degrees C in temperature at the site depending on the season. Anatomical characteristics analyzed were annual earlywood, latewood and ring widths, intra-ring wood densities (earlywood, latewood and mean wood density), tracheid width, length, wall thickness, lumen diameter, wall thickness:lumen diameter ratio and mass per unit length (coarseness), and numbers of rays, resin canals and tracheids per xylem cross-sectional area. Elevated [CO(2)] increased ring width in four of six treatment years; earlywood width increased in the first two years and latewood width in the third year. Tracheid walls in both the earlywood and latewood tended to become thicker over the 6-year treatment period when temperature or [CO(2)] was elevated alone, whereas in the combined treatment they tended to become thinner relative to the tracheids of trees grown under ambient conditions. Latewood tracheid lumen diameters were larger in all the treatments relative to ambient conditions over the 6-year period, whereas lumen diameters in earlywood increased only in response to elevated [CO(2)] and were 3-6% smaller in the treatments with elevated temperature than in ambient conditions. Tracheid width, length and coarseness were greater in trees grown in elevated than in ambient temperature. The number of resin canals per mm(2) decreased in the elevated [CO(2)] treatment and increased in the elevated temperature treatments relative to ambient conditions. The treatments decreased the number of rays and tracheids per mm(2) of cross

Hydraulic and mechanical properties of young Norway spruce clones related to growth and wood structure

PubMed Central

ROSNER, SABINE; KLEIN, ANDREA; MÜLLER, ULRICH; KARLSSON, BO

2011-01-01

Summary Stem segments of eight five-year-old Norway spruce (Picea abies (L.) Karst.) clones differing in growth characteristics were tested for maximum specific hydraulic conductivity (ks100), vulnerability to cavitation and behavior under mechanical stress. The vulnerability of the clones to cavitation was assessed by measuring the applied air pressure required to cause 12 and 50% loss of conductivity (Ψ12, Ψ50) and the percent loss of conductivity at 4 MPa applied air pressure (PLC4MPa). The bending strength and stiffness and the axial compression strength and stiffness of the same stem segments were measured to characterize wood mechanical properties. Growth ring width, wood density, latewood percentage, lumen diameter, cell wall thickness, tracheid length and pit dimensions of earlywood cells, spiral grain and microfibril angles were examined to identify structure–function relationships. High ks100 was strongly and positively related to spiral grain angle, which corresponded positively to tracheid length and pit dimensions. Spiral grain may reduce flow resistance of the bordered pits of the first earlywood tracheids, which are characterized by rounded tips and an equal distribution of pits along the entire length. Wood density was unrelated to hydraulic vulnerability parameters. Traits associated with higher hydraulic vulnerability were long tracheids, high latewood percentage and thick earlywood cell walls. The positive relationship between earlywood cell wall thickness and vulnerability to cavitation suggest that air seeding through the margo of bordered pits may occur in earlywood. There was a positive phenotypic and genotypic relationship between ks100 and PLC4MPa, and both parameters were positively related to tree growth rate. Variability in mechanical properties depended mostly on wood density, but also on the amount of compression wood. Accordingly, hydraulic conductivity and mechanical strength or stiffness showed no tradeoff. PMID:17472942

Basal area growth, carbon isotope discrimination, and intrinsic ...

EPA Pesticide Factsheets

Many hectares of intensively managed Douglas-fir (Pseudotsuga menziesii Mirb. Franco) stands in western North America are fertilized with nitrogen to increase growth rates. Understanding the mechanisms of response facilitates prioritization of stands for treatment. The objective of this study was to test the hypothesis that the short-term basal area growth response to a single application of 224 kg N ha-1 as urea was associated with reduced stable carbon isotope discrimination (∆13C) and increased intrinsic water use efficiency (iWUE) in a 20-yr-old plantation of Douglas-fir in the Oregon Coast Range, USA. Increment cores were measured to estimate earlywood, latewood, and total basal area increment over a time series from 1997 to 2015. Stable carbon isotope discrimination and iWUE were estimated using earlywood and latewood stable carbon isotope concentrations in tree-ring holocellulose starting seven years before fertilization in early 2009 and ending seven years after treatment. A highly significant interaction effect between fertilization treatment and year was found for total basal area growth and earlywood basal area increment. Fertilized trees showed significant total basal area growth and earlywood basal area increment in the first (2009) and second (2010) growing seasons after fertilization in 2009. A marginally significant fertilization effect was found for latewood basal area increment only in the first growing season after treatment. A significant i

Seasonal and Perennial Changes in the Distribution of Water in the Sapwood of Conifers in a Sub-Frigid Zone1

PubMed Central

Utsumi, Yasuhiro; Sano, Yuzou; Funada, Ryo; Ohtani, Jun; Fujikawa, Seizo

2003-01-01

An analysis was made of progressive changes in patterns of cavitation in the sapwood of three species of conifer (Larix kaempferi, Abies sachalinensis, and Picea jezoensis) that were growing in a sub-frigid zone. In all three conifers, all tracheids of the newly forming outermost annual ring were filled with water or cytoplasm during the period from May to August. However, many tracheids in the transition zone from earlywood to latewood lost water in September, presumably through drought-induced cavitation. Cavitated tracheids tended to be continuously distributed in a tangential direction. Subsequently, some earlywood tracheids of the outermost annual ring lost water during the period from January to March. This was associated with freeze-thaw cycles. In the second and third annual rings from the cambium of all three conifers, the lumina of most tracheids in the transition zone from earlywood to latewood contained no water. In contrast, some latewood tracheids near the annual ring boundary and many earlywood tracheids retained water in their lumina. The third annual ring had more cavitated tracheids than the second annual ring. Our observations indicated that cavitation progressed gradually in the tracheids of the conifers and that they were never refilled once cavitation had occurred. The region involved in water transport in conifers did not include the entire sapwood and differed among annual rings. PMID:12692342

Unstable relationships between tree ring δ18O and climate variables over southwestern China: possible impacts from increasing central Pacific SSTs

NASA Astrophysics Data System (ADS)

An, Wenling; Liu, Xiaohong; Hou, Shugui; Zeng, Xiaomin; Sun, Weizhen; Wang, Wenzhi; Wang, Yu; Xu, Guobao; Ren, Jiawen

2018-05-01

In this study, we investigated the potential influence of central and eastern Pacific sea surface temperatures (SSTs) on the unstable relationship between earlywood δ18O and climatic factors in the southwestern China from 1902 to 2005. The results show that the strength of the climate signals recorded in the earlywood δ18O series has declined since the late 1970s. This reduction in signal strength may have been caused by the changes in the local hydroclimate, which is associated with the increasing SSTs in the central Pacific Ocean over recent decades. Alongside these increasing SSTs in the central Pacific, southwestern China has experienced more droughts, as well as more severe droughts through the late spring and early summer during the central Pacific (CP) El Niño years than during the eastern Pacific (EP) El Niño years in recent decades. This increased drought frequency may have weakened the response of earlywood δ18O to climate variables.

Seasonal climate signals from multiple tree ring metrics: A case study of Pinus ponderosa in the upper Columbia River Basin

NASA Astrophysics Data System (ADS)

Dannenberg, Matthew P.; Wise, Erika K.

2016-04-01

Projected changes in the seasonality of hydroclimatic regimes are likely to have important implications for water resources and terrestrial ecosystems in the U.S. Pacific Northwest. The tree ring record, which has frequently been used to position recent changes in a longer-term context, typically relies on signals embedded in the total ring width of tree rings. Additional climatic inferences at a subannual temporal scale can be made using alternative tree ring metrics such as earlywood and latewood widths and the density of tree ring latewood. Here we examine seasonal precipitation and temperature signals embedded in total ring width, earlywood width, adjusted latewood width, and blue intensity chronologies from a network of six Pinus ponderosa sites in and surrounding the upper Columbia River Basin of the U.S. Pacific Northwest. We also evaluate the potential for combining multiple tree ring metrics together in reconstructions of past cool- and warm-season precipitation. The common signal among all metrics and sites is related to warm-season precipitation. Earlywood and latewood widths differ primarily in their sensitivity to conditions in the year prior to growth. Total and earlywood widths from the lowest elevation sites also reflect cool-season moisture. Effective correlation analyses and composite-plus-scale tests suggest that combining multiple tree ring metrics together may improve reconstructions of warm-season precipitation. For cool-season precipitation, total ring width alone explains more variance than any other individual metric or combination of metrics. The composite-plus-scale tests show that variance-scaled precipitation reconstructions in the upper Columbia River Basin may be asymmetric in their ability to capture extreme events.

[Dendrochronology of Chinese pine in Mulan-Weichang, Hebei Province: a primary study].

PubMed

Cui, Ming-xing; He, Xing-yuan; Chen, Wei; Chen, Zhen-ju; Zhou, Chang-hong; Wu, Tao

2008-11-01

Dendroclimatic methods were used to investigate the relationships between the growth of Chinese pine (Pinus tabulaeformis Carr.) and the climatic parameters in Mulan-Weichang of Hebei Province. The results showed that Chinese pine presented high sensitivity to climatic changes, and its earlywood width showed the highest sensitivity. There was a significant negative correlation between the tree-ring width chronology of Chinese pine and the air temperature in May-June. The precipitation and relative humidity in June had strong positive effects on the growth of earlywood, the precipitation from September to next September had significant positive effects on Chinese pine growth, and the relative humidity in winter more strongly affected the growth of latewood than of earlywood. There was a definite correlation between the tree-ring width chronology of Chinese pine and the large scale climate fluctuation. From 1951 to 2006, the increase of air temperature in study area was significant, and the sensitivity of Chinese pine to the variations of local temperature and precipitation decreased, presenting an inverse transforming trend with increasing temperature. Greater differences were observed between the reconstructed and observed data of mean temperature in May - June in a century scale, suggesting that the tree-ring growth of Chinese pine in study area had a greater fluctuation of sensitivity to the variation of climatic factors.

Transcriptome profiling of Pinus radiata juvenile wood with contrasting stiffness identifies putative candidate genes involved in microfibril orientation and cell wall mechanics

PubMed Central

2011-01-01

Background The mechanical properties of wood are largely determined by the orientation of cellulose microfibrils in secondary cell walls. Several genes and their allelic variants have previously been found to affect microfibril angle (MFA) and wood stiffness; however, the molecular mechanisms controlling microfibril orientation and mechanical strength are largely uncharacterised. In the present study, cDNA microarrays were used to compare gene expression in developing xylem with contrasting stiffness and MFA in juvenile Pinus radiata trees in order to gain further insights into the molecular mechanisms underlying microfibril orientation and cell wall mechanics. Results Juvenile radiata pine trees with higher stiffness (HS) had lower MFA in the earlywood and latewood of each ring compared to low stiffness (LS) trees. Approximately 3.4 to 14.5% out of 3, 320 xylem unigenes on cDNA microarrays were differentially regulated in juvenile wood with contrasting stiffness and MFA. Greater variation in MFA and stiffness was observed in earlywood compared to latewood, suggesting earlywood contributes most to differences in stiffness; however, 3-4 times more genes were differentially regulated in latewood than in earlywood. A total of 108 xylem unigenes were differentially regulated in juvenile wood with HS and LS in at least two seasons, including 43 unigenes with unknown functions. Many genes involved in cytoskeleton development and secondary wall formation (cellulose and lignin biosynthesis) were preferentially transcribed in wood with HS and low MFA. In contrast, several genes involved in cell division and primary wall synthesis were more abundantly transcribed in LS wood with high MFA. Conclusions Microarray expression profiles in Pinus radiata juvenile wood with contrasting stiffness has shed more light on the transcriptional control of microfibril orientation and the mechanical properties of wood. The identified candidate genes provide an invaluable resource for further

Xylem Development of Loblolly Pine During Irrigation and Simulated Drought

Treesearch

David M. Moehring; Charles X. Grano; John R. Bassett

1975-01-01

Induced drought decreased the total number of tracheid cells produced in a season and the radial width of late wood tracheids. Transition from earlywood to late woodo ccurred in early June and was little affected by irrigation.

Wood anatomical parameters of lowland European oak and Scots pine as proxies for climate reconstructions

NASA Astrophysics Data System (ADS)

Balanzategui, Daniel; Heußner, Karl-Uwe; Wazny, Tomasz; Helle, Gerd; Heinrich, Ingo

2017-04-01

Tree-ring based temperature reconstructions from the temperate lowlands worldwide are largely missing due to diffuse climate signals so far found in tree-ring widths. This motivated us to concentrate our efforts on the wood anatomies of two common European tree species, the European oak (Quercus robur) and Scots pine (Pinus sylvestris). We combined core samples of living trees with archaeological wood from northern Germany and Poland. We measured approx. 46,000 earlywood oak vessels of 34 trees covering the period AD 1500 to 2016 and approx. 7.5 million pine tracheid cells of 41 trees covering the period AD 1300 to 2010. First climate growth analyses indicate that both oak earlywood vessel and pine tracheid parameters contain climate signals which are different and more significant than those found in tree-ring widths. Preliminary results will be presented and discussed at EGU for the first time.

An improved method and data analysis for ultrasound acoustic emissions and xylem vulnerability in conifer wood.

PubMed

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.

Severe winter rings of oak trees (Quercus robur L.) from Central European Russia.

PubMed

Khasanov, B F

2013-11-01

Oak trees were sampled in a flood plain forest in the valley of the Zapadnaya Dvina (Daugava) river (Tver region, Russia). Annual rings of the time period from 1826 to 2010 were studied. Anatomically distinct rings with a stripe of small-sized cells in the innermost part and narrow earlywood vessels located in three to four rows occurred in 1861, 1862, 1929, 1940, 1942, 1956 and 1979. Deviations of earlywood development were associated with the drop of winter temperature below -42 °C. The percentage of severe winter ring (SWR) occurrence depends upon tree age and decreases from 75.6 % in younger specimens (under 41 years old at the time of the severe winter) to 27.1 % in middle-aged ones (from 41 to 80 years) to 3.5 % in trees older than 80 years. Described anatomical features can be used in the reconstruction of severe winter frequency in the past.

Axial and radial water flow in the trunks of oak trees: a quantitative and qualitative analysis.

PubMed

Granier, A; Anfodillo, T; Sabatti, M; Cochard, H; Dreyer, E; Tomasi, M; Valentini, R; Bréda, N

1994-12-01

Axial water flow in the trunks of mature oak trees (Quercus petraea (Matt.) Liebl. and Q. robur L.) was studied by four independent techniques: water absorption from a cut trunk, sap flowmeters, heat pulse velocity (HPV) and thermoimaging. Estimation of the total water flow with sap flowmeters, HPV and water absorption yielded comparable results. We concluded from dye colorations, thermograms and axial profiles of sap flow and heat pulse velocity that, in intact trunks, most of the flow occurred in the current-year ring, where early-wood vessels in the outermost ring were still functional. Nevertheless, there was significant flow in the older rings of the xylem. Total water flow through the trunk was only slightly reduced when air embolisms were artificially induced in early-wood vessels, probably because there was little change in hydraulic conductance in the root-leaf sap pathway. Embolization of the current-year vessels reactivated transport in the older rings.

Morphological characteristics of loblolly pine wood as related to specific gravity, growth rate and distance from pith

Treesearch

Charles W. McMillin

1968-01-01

Earlywood and latewood tracheid length and transverse cellular dimensions of wood removed from stems of loblolly pine (Pinus taeda L.) and factorially aegregated by specific gravity, rings from the pith, and growth rate were determined from sample chips. The independent relationships of each factor with fiber morphology are described.

Specific heat of ovendry loblolly pine wood

Treesearch

Charles W. McMillin

1969-01-01

In the range of 333 K to 413 K, the specific heat of ovendry loblolly pine (Pinus taeda L.) wood was expressed by a linear function of temperature. No relationship was detected with specific gravity, growth rate, or distance from the pith; nor were differences found between earlywood and latewood.

Method for computing a roughness factor for veneer surfaces

Treesearch

Chung-Yun Hse

1972-01-01

Equations for determining the roughness factor (ratio of true surface to apparent area) of rotary-cut veneer were derived from an assumed tracheid model. With data measured on southern pine veneers, the equations indicated that the roughness factor of latewood was near unity, whereas that of earlywood was about 2.

Guying to prevent wind sway influences loblolly pine growth and wood properties

Treesearch

James D. Burton; Diana M. Smith

1972-01-01

Restraining young loblolly pine (Pinus taeda L.) trees from normal swaying in the wind markedly reduced radial growth in the immobilized portion of the bole and accelerated it in the upper, free-swaying portion. Guying also reduced specific gravity, number of earlywood and latewood tracheids, latewood tracheid diameter, and amount of compression wood...

Two-dimensional finite element heat transfer model of softwood. Part I, Effective thermal conductivity

Treesearch

John F. Hunt; Hongmei Gu

2006-01-01

The anisotropy of wood complicates solution of heat and mass transfer problems that require analyses be based on fundamental material properties of the wood structure. Most heat transfer models use average thermal properties across either the radial or tangential direction and do not differentiate the effects of cellular alignment, earlywood/latewood differences, or...

Mechanical properties of individual southern pine fibers. Part I. Determination of variability of stress-strain curves with respect to tree height and juvenility

Treesearch

Leslie H. Groom; Laurence Mott; Stephen Shaler

2002-01-01

This paper is the first in a three-part series investigating the mechanical properties of loblolly pine fibers. This paper outlines the experimental method and subsequent variation of latewood fiber mechanical properties in relation to tree position. Subsequent papers will deal with differences between earlywood and latewood fibers and effect of juvenility and tree...

Two dimensional finite element heat transfer models for softwood

Treesearch

Hongmei Gu; John F. Hunt

2004-01-01

The anisotropy of wood creates a complex problem for solving heat and mass transfer problems that require analyses be based on fundamental material properties of the wood structure. Most heat transfer models use average thermal properties across either the radial or tangential directions and have not differentiated the effects of cellular alignment, earlywood/latewood...

Cosmic ray event in 994 C.E. recorded in radiocarbon from Danish oak

NASA Astrophysics Data System (ADS)

Fogtmann-Schulz, A.; Østbø, S. M.; Nielsen, S. G. B.; Olsen, J.; Karoff, C.; Knudsen, M. F.

2017-08-01

We present measurements of radiocarbon in annual tree rings from the time period 980-1006 Common Era (C.E.), hereby covering the cosmic ray event in 994 C.E. The new radiocarbon record from Danish oak is based on both earlywood and latewood fractions of the tree rings, which makes it possible to study seasonal variations in 14C production. The measurements show a rapid increase of ˜10‰ from 993 to 994 C.E. in latewood, followed by a modest decline and relatively high values over the ensuing ˜10 years. This rapid increase occurs from 994 to 995 C.E. in earlywood, suggesting that the cosmic ray event most likely occurred during the period between April and June 994 C.E. Our new record from Danish oak shows strong agreement with existing Δ14C records from Japan, thus supporting the hypothesis that the 994 C.E. cosmic ray event was uniform throughout the Northern Hemisphere and therefore can be used as an astrochronological tie point to anchor floating chronologies of ancient history.

Mechanical properties of individual southern pine fibers. Part I. Determination and variability of stress-strain curves with respect to tree height and juvenility

Treesearch

Leslie H. Groom; Laurence Mott; Stephen Shaler

2002-01-01

This paper is the first in a three-part series investigating the mechanical properties of loblolly pine fibers. This paper outlines the experimental method and subsequent variation of latewood fiber mechan-ical properties in relation to tree position. Subsequent papers will deal with differences between early-wood and latewood fibers and effect of juvenility and tree...

Simultaneous bond degradation and bond formation during phenol-formaldehyde curing with wood

Treesearch

Daniel J. Yelle; John Ralph

2016-01-01

Bonding of wood using phenol–formaldehyde adhesive develops highly durable bonds. Phenol– formaldehyde is believed to form primary bonds with wood cell wall polymers (e.g., lignin). However, it is unclear how this adhesive interacts and bonds to lignin. Through wood solubilisation methodologies, earlywood and latewood bonded assemblies were characterized using two-...

Tool Forces and Chip Formation In Orthogonal Cutting Of Loblolly Pine

Treesearch

George E. Woodson; Peter Koch

1970-01-01

Specimens of earlywood and latewood of Pinus taeda L. were excised so that length along the grain was 3 inches and thickness was 0.1 inch. These specimens were cut orthogonally-as with a carpenter's plane-in the three major directions. Cutting velocity was 2 inches per minute. When cutting was in the planing (90-O) direction, thin chips,...

Ash content of loblolly pine wood as related to specific gravity, growth rate, and distance from pith

Treesearch

Charles W. McMillin

1968-01-01

In earlywood of Pinus taeda L. grown in central Louisiana, ash content generally decreased with increasing distance from the pith and icnreased with increases in rate of tree growth (as measured in rings per inch). Latewood ash content was unrelated to the gross wood factors of distance, from the pith, specific gravity, and growth rate. The ash...

Effect of wood grain and veneer side on loblolly pine veneer wettability

Treesearch

Todd E. Shupe; Chung Y. Hse; Elvin T. Choong; Leslie H. Groom

1998-01-01

Research was initiated to determine the effect of veneer side (tight or loose), and wood grain (earlywood or latewood) on the wettability of loblolly pine veneer. Contact angle measurements were performed with phenol-formaldehyde resin and distilled water. The resin and distilled water showed slightly higher contact angle mean values on the latewood portion for both...

Effect of early age woody and herbaceous competition control on wood properties of loblolly pine

Treesearch

F. Antony; L. R. Schimleck; L. Jordan; Alexander Clark; R. F. Daniels

2011-01-01

Early age competition control has been reported to significantly improve the growth and yield of plantation grown loblolly pine. The objective of this paper is to understand the changes in wood properties: basal area weighted whole disk SG, earlywood SG (EWSG), latewood SG (LWSG) and latewood percent (LWP) of 14 year-old trees which received early age herbaceous and...

Wood and Bark Properties of Spruce Pine

Treesearch

F. G. Manwiller

1972-01-01

Weighted stem averagees were determined for wood and bard of 72 trees representing the commercial range of Pinus glabra Walt. The trees were stratified into three age classes (15, 30, and 45 years) and two growth rates (averaging 4.9 and 9.0 rings per inch). Within-stem variation was determined from 1,269 earlywood and alte wood sampling points in...

Heat of combustion of various southern pine materials

Treesearch

Elaine T. Howard

1972-01-01

As measured with an oxygen-bomb calorimeter, heats of combustion averaged about 8,600 Btu/ovendry lb. for stemwood, rootwood, and earlywood, and latewood. In stemwood, 54 percent of the variation was related to extractive content. Values for bark were greatest in samples from the base of the tree and decreased with increasing height in tree. Values for resinous wood...

Heat of combustion of various southern pine materials

Treesearch

E.T. Howard

1973-01-01

As measured with an oxygen-bomb calorimeter, heats of combustion averaged about 8,600 Btu/ovendry lb. for stemwood, rootwood, earlywood, and latewood. In stemwood, 54 percent of the variation was related to extractive content. Values for bark were greatest in samples from the base of the tree and decreased with increasing height in tree. Values for resinous wood varied...

Erosion rates of wood during natural weathering. Part I, Effects of grain angle and surface texture

Treesearch

R. Sam Williams; Mark T. Knaebe; Peter G. Sotos; William C. Feist

2001-01-01

This is the first in a series of reports on the erosion rates of wood exposed outdoors near Madison, Wisconsin. The specimens were oriented vertically, facing south; erosion was measured annually for the first several years and biannually for the remainder of the exposure. In the work reported here, the erosion rates of earlywood and latewood were determined for smooth...

Two-dimensional finite element heat transfer model of softwood. Part II, Macrostructural effects

Treesearch

Hongmei Gu; John F. Hunt

2006-01-01

A two-dimensional finite element model was used to study the effects of structural features on transient heat transfer in softwood lumber with various orientations. Transient core temperature was modeled for lumber samples “cut” from various locations within a simulated log. The effects of ring orientation, earlywood to latewood (E/L) ratio, and ring density were...

Erosion rates of wood during natural weathering. Part III, Effect of exposure angle on erosion rate

Treesearch

R. Sam Williams; Mark T. Knaebe; James W. Evans; William C. Feist

2001-01-01

This is the third in a series of reports on the erosion rates of wood exposed outdoors near Madison, Wisconsin. The specimens were exposed at an orientation of 90* or 45* facing south or horizontally (0*) for 10 years. Erosion was measured annually for the first 8 years and after 10 years. The erosion rates of earlywood (springwood) and latewood (summerwood) were...

Carbon Allocation of 13CO2-labeled Photoassimilate in Larix gmelinii Saplings - A Physiological Basis for Isotope Dendroclimatology in Eastern Siberia.

NASA Astrophysics Data System (ADS)

Kagawa, A.; Sugimoto, A.; Maximov, T. C.

2006-12-01

Tree-ring density and widths have been successfully used to reconstruct summer temperatures in high- northern latitudes, although a discrepancy between tree-growth and temperature has been found for recent decades. The so-called "reduced sensitivity" of tree rings to summer temperatures has been observed especially strongly in northern Siberia (Briffa et al., 1998) and drought stress (increased water use efficiency) arose from global warming and/or increasing CO2 are suggested as causes (Barber et al. 2000, Saurer et al. 2004). By using carbon isotope ratio as an indicator of drought stress and ring-width/density as indicators of growth, we can clarify how drought stress caused by recent global warming affects wood formation of Siberian trees. However, isotope dendroclimatology is still in its infancy and our understanding of basic physiological processes of isotope signal transfer from leaves to tree rings is insufficient. In order to understand translocation, storage, and allocation of photoassimilate to different organs of trees, we pulse- labeled ten L. gmelinii growing in a continuous permafrost zone with stable 13CO2. We studied seasonal course of carbon allocation patterns of photoassimilate among needles, branches, stem and roots and also how spring, summer, and autumn photoassimilate is later used for both earlywood and latewood formation. About half of the carbon in new needles was derived from stored material. The starch pool in non- needle parts, which can be used for xylem formation, drew about 43 percent of its carbon from previous year's photoassimilate, suggesting that carbon storage is the key mechanism behind autocorrelation in (isotope) dendroclimatology. Analysis of intra-annual 13C of the tree rings formed after the labeling revealed that earlywood contained photoassimilate from the previous summer and autumn as well as from the current spring. Latewood was mainly composed of photoassimilate from the current year's summer/autumn, although it

Force-displacement measurements of earlywood bordered pits using a mesomechanical tester

Treesearch

Samuel L. Zelinka; Keith J. Bourne; John C. Hermanson; Samuel V. Glass; Adriana Costa; Alex C. Wiedenhoeft

2015-01-01

The elastic properties of pit membranes are reported to have important implications in understanding air-seeding phenomena in gymnosperms, and pit aspiration plays a large role in wood technological applications such as wood drying and preservative treatment. Here we present force–displacement measurements for pit membranes of circular bordered pits, collected on a...

Hydraulic efficiency compromises compression strength perpendicular to the grain in Norway spruce trunkwood

PubMed Central

2011-01-01

The aim of this study was to investigate bending stiffness and compression strength perpendicular to the grain of Norway spruce (Picea abies (L.) Karst.) trunkwood with different anatomical and hydraulic properties. Hydraulically less safe mature sapwood had bigger hydraulic lumen diameters and higher specific hydraulic conductivities than hydraulically safer juvenile wood. Bending stiffness (MOE) was higher, whereas radial compression strength lower in mature than in juvenile wood. A density-based tradeoff between MOE and hydraulic efficiency was apparent in mature wood only. Across cambial age, bending stiffness did not compromise hydraulic efficiency due to variation in latewood percent and because of the structural demands of the tree top (e.g. high flexibility). Radial compression strength compromised, however, hydraulic efficiency because it was extremely dependent on the characteristics of the “weakest” wood part, the highly conductive earlywood. An increase in conduit wall reinforcement of earlywood tracheids would be too costly for the tree. Increasing radial compression strength by modification of microfibril angles or ray cell number could result in a decrease of MOE, which would negatively affect the trunk’s capability to support the crown. We propose that radial compression strength could be an easily assessable and highly predictive parameter for the resistance against implosion or vulnerability to cavitation across conifer species, which should be topic of further studies. PMID:22058609

Is blue intensity ready to replace maximum latewood density as a strong temperature proxy? A tree-ring case study on Scots pine from northern Sweden

NASA Astrophysics Data System (ADS)

Björklund, J. A.; Gunnarson, B. E.; Seftigen, K.; Esper, J.; Linderholm, H. W.

2013-09-01

At high latitudes, where low temperatures mainly limit tree-growth, measurements of wood density (e.g. Maximum Latewood Density, MXD) using the X-Ray methodology provide a temperature proxy that is superior to that of TRW. Density measurements are however costly and time consuming and have lead to experimentation with optical flatbed scanners to produce Maximum Blue Intensity (BImax). BImax is an excellent proxy for density on annual scale but very limited in skill on centennial scale. Discolouration between samples is limiting BImax where specific brightnesses can have different densities. To overcome this, the new un-exploited parameter Δ blue intensity (ΔBI) was constructed by using the brightness in the earlywood (BIEW) as background, (BImax - BIEW = ΔBI). This parameter was tested on X-Ray material (MXD - earlywood density = ΔMXD) and showed great potential both as a quality control and as a booster of climate signals. Unfortunately since the relationship between grey scale and density is not linear, and between-sample brightness can differ tremendously for similar densities, ΔBI cannot fully match ΔMXD in skill as climate proxy on centennial scale. For ΔBI to stand alone, the range of brightness/density offset must be reduced. Further studies are needed to evaluate this possibility, and solutions might include heavier sample treatment (reflux with chemicals) or image-data treatment (digitally manipulating base-line levels of brightness).

Generation and analysis of expressed sequence tags from six developing xylem libraries in Pinus radiata D. Don

PubMed Central

Li, Xinguo; Wu, Harry X; Dillon, Shannon K; Southerton, Simon G

2009-01-01

Background Wood is a major renewable natural resource for the timber, fibre and bioenergy industry. Pinus radiata D. Don is the most important commercial plantation tree species in Australia and several other countries; however, genomic resources for this species are very limited in public databases. Our primary objective was to sequence a large number of expressed sequence tags (ESTs) from genes involved in wood formation in radiata pine. Results Six developing xylem cDNA libraries were constructed from earlywood and latewood tissues sampled at juvenile (7 yrs), transition (11 yrs) and mature (30 yrs) ages, respectively. These xylem tissues represent six typical development stages in a rotation period of radiata pine. A total of 6,389 high quality ESTs were collected from 5,952 cDNA clones. Assembly of 5,952 ESTs from 5' end sequences generated 3,304 unigenes including 952 contigs and 2,352 singletons. About 97.0% of the 5,952 ESTs and 96.1% of the unigenes have matches in the UniProt and TIGR databases. Of the 3,174 unigenes with matches, 42.9% were not assigned GO (Gene Ontology) terms and their functions are unknown or unclassified. More than half (52.1%) of the 5,952 ESTs have matches in the Pfam database and represent 772 known protein families. About 18.0% of the 5,952 ESTs matched cell wall related genes in the MAIZEWALL database, representing all 18 categories, 91 of all 174 families and possibly 557 genes. Fifteen cell wall-related genes are ranked in the 30 most abundant genes, including CesA, tubulin, AGP, SAMS, actin, laccase, CCoAMT, MetE, phytocyanin, pectate lyase, cellulase, SuSy, expansin, chitinase and UDP-glucose dehydrogenase. Based on the PlantTFDB database 41 of the 64 transcription factor families in the poplar genome were identified as being involved in radiata pine wood formation. Comparative analysis of GO term abundance revealed a distinct transcriptome in juvenile earlywood formation compared to other stages of wood development

Tree-ring proxies of larch bud moth defoliation: latewood width and blue intensity are more precise than tree-ring width.

PubMed

Arbellay, Estelle; Jarvis, Ingrid; Chavardès, Raphaël D; Daniels, Lori D; Stoffel, Markus

2018-05-19

Reconstructions of defoliation by larch bud moth (LBM, Zeiraphera diniana Gn.) based on European larch (Larix decidua Mill.) tree rings have unraveled outbreak patterns over exceptional temporal and spatial scales. In this study, we conducted tree-ring analyses on 105 increment cores of European larch from the Valais Alps, Switzerland. The well-documented history of LBM outbreaks in Valais provided a solid baseline for evaluating the LBM defoliation signal in multiple tree-ring parameters. First, we used tree-ring width measurements along with regional records of LBM outbreaks to reconstruct the occurrence of these events at two sites within the Swiss Alps. Second, we measured earlywood width, latewood width and blue intensity, and compared these parameters with tree-ring width to assess the capacity of each proxy to detect LBM defoliation. A total of six LBM outbreaks were reconstructed for the two sites between AD 1850 and 2000. Growth suppression induced by LBM was, on average, highest in latewood width (59%), followed by total ring width (54%), earlywood width (51%) and blue intensity (26%). We show that latewood width and blue intensity can improve the temporal accuracy of LBM outbreak reconstructions, as both proxies systematically detected LBM defoliation in the first year it occurred, as well as the differentiation between defoliation and non-defoliation years. This study introduces blue intensity as a promising new proxy of insect defoliation and encourages its use in conjunction with latewood width.

Tree-ring anatomy and carbon isotope ratio show both direct and legacy effects of climate on bimodal xylem formation in Pinus pinea.

PubMed

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.

Stem sapwood permeability in relation to crown dominance and site quality in self-thinning fire-origin lodgepole pine stands.

PubMed

Reid, Douglas E B; Silins, Uldis; Lieffers, Victor J

2003-08-01

Stem sapwood hydraulic permeability, tree leaf area, sapwood basal area, earlywood to latewood ratio of annual rings, radial variation in hydraulic permeability and stem hydraulic capacity were examined in dominant (D), codominant (CD) and suppressed (SP) lodgepole pine (Pinus contorta Dougl. ex Loud.) trees growing on medium and poor sites. Hydraulic permeability on a sapwood area basis (ks) was lower in suppressed trees (0.71 x 10(-12) m2) compared to dominants (1.97 x 10(-12) m2) and codominants (1.79 x 10(-12) m2), and higher on medium than on poor sites. The leaf/sapwood area ratio (S) varied with crown dominance position (D > CD > SP) but not by site type. Leaf specific conductivity (kL) did not vary between crown classes or site types. The relationship between leaf area and stem hydraulic supply capacity (Q*) was strong, but differed among crown classes. Dominant trees and trees from the medium sites had a greater proportion of earlywood in outer rings of sapwood than suppressed trees. Sapwood permeability declined from the cambium to the sapwood-heartwood boundary in all samples, but the decline was more gradual in dominant trees compared to codominant and suppressed trees; differences in the radial variation in sapwood permeability may be related to differences in S. Sapwood permeability is positively related to crown dominance, whereas subdominant (CD and SP) trees have greater Q* in relation to leaf area, leading us to propose that this may give subdominant trees a survival advantage, slowing self-thinning.

Can forest dieback and tree death be predicted by prior changes in wood anatomy?

NASA Astrophysics Data System (ADS)

Colangelo, Michele; Julio Camarero, Jesus; De Micco, Veronica; Borghetti, Marco; Gentilesca, Tiziana; Sanchez-Salguero, Raul; Ripullone, Francesco

2017-04-01

Climate warming is expected to amplify drought stress resulting in more intense and widespread dieback episodes and increasing mortality rates. Studies on quantitative wood anatomy and dendrochronology have demonstrated their potential to supply useful information on the causes of tree decline, although this approach is basically observational and retrospective. Moreover, the long-term reconstruction of wood anatomical features, strictly linked to the evolution of xylem anatomy plasticity through time, allow investigating hydraulic adjustments of trees. In this study, we analyzed wood-anatomical variables in two Italian oak forests where recent episodes of dieback and mortality have been reported. We analyzed in coexisting now-dead and living trees the following wood-anatomical variables: annual tree-ring area, earlywood (EW) and latewood (LW) areas, absolute and relative (%) areas occupied by vessels in the EW and LW, EW and LW vessel areas, EW and LW vessel density and vessel diameter classification. We also calculated the hydraulic diameter (Dh) for all vessels measured within each ring by weighting individual conduit diameters to correspond to the average Hagen-Poiseuille lumen theoretical hydraulic conductivity for a vessel size. Wood-anatomical analyses showed that declining and dead trees were more sensitive to drought stress compared to non declining trees, indicating different susceptibility to water shortage between trees. Dead trees did not form earlywood vessels with smaller lumen diameter than surviving trees but tended to form wider latewood vessels with a higher percentage of vessel area. We discuss the results and implications focusing on those proved more sensitive to the phenomena of decline and mortality.

Season-specific climate signal and reconstruction from a new tree-ring network in the southwestern U.S

NASA Astrophysics Data System (ADS)

Griffin, D.; Woodhouse, C. A.; Meko, D. M.; Stahle, D. W.; Faulstich, H.; Leavitt, S. W.; Touchan, R.; Castro, C. L.; Carrillo, C.

2011-12-01

Our research group has updated existing tree-ring collections from over 50 sampling sites in the southwestern U.S. The new and archived specimens, carefully dated with dendrochronology, have been analyzed for width variations of "earlywood" and "latewood." These are the two components of annual rings in conifers that form in spring and summer, respectively. The network of primary tree-ring data has been used to develop a suite of well-replicated chronologies that extend through the 2008 growing season and are sensitive to the season-specific climate variability of the Southwest. Correlation function analysis indicates that the earlywood chronologies are closely related to cool season (October-April) precipitation variability and the chronologies derived from latewood are generally sensitive to precipitation and temperature conditions during the warm season (June-August). These proxy data originate from biological organisms and are not without bias; however, they do constitute a new means for evaluating the recent paleoclimatic history of the North American summer monsoon. The monsoon is a major component of the region's climate, impacting social and environmental systems and delivering up to 60% of the annual precipitation in the southwestern U.S. We have developed latewood-based retrodictions of monsoon precipitation that explain over half of the variance in the instrumental record, pass standard verification tests, and point to periods of persistent drought and wetness during the last 300-500 years. These reconstructions are being used to evaluate the monsoon's long-term spatiotemporal variability and its relationship to cool season climate and the major modes of ocean-atmosphere variability.

Function and dynamics of auxin and carbohydrates during earlywood/latewood transition in scots pine.

PubMed

Uggla, C; Magel, E; Moritz, T; Sundberg, B

2001-04-01

In temperate regions the annual pattern of wood development is characterized by the formation of radially narrow and thick walled latewood cells. This takes place at the later part of the growing season when cambial cell division declines. To gain new insight into the regulation of this process, micro-analytical techniques were used to visualize the distribution of indole-3-acetic acid (IAA), soluble carbohydrates, and activities of sucrose (Suc)-metabolizing enzymes across the cambial region tissues in Scots pine (Pinus sylvestris). The total amount of IAA in the cambial region did not change with latewood initiation. But its radial distribution pattern was altered, resulting in an increased concentration in the cambial meristem and its recent derivatives. Thus, initiation of latewood formation and cessation of cambial cell division is not a consequence of decreased IAA concentrations in dividing and expanding cells. Rather, IAA most likely has a role in defining the altered developmental pattern associated with latewood formation. Carbohydrates and enzyme activities showed distinctive radial distribution patterns. Suc peaked in the phloem and decreased sharply to low levels across the cambial zone, whereas fructose and glucose reached their highest levels in the maturing tracheids. Suc synthase was the dominating Suc cleaving enzyme with a peak in the secondary wall-forming tracheids and in the phloem. Soluble acid invertase peaked in dividing and expanding cells. Suc-phosphate synthase had its highest activities in the phloem. Activities of cell wall bound invertase were low. The absence of major seasonal variations indicates that carbohydrate availability is not a trigger for latewood initiation. However, steep concentration gradients of the sugars suggest a role for sugar signaling in vascular development.

Climatic Signals from Intra-annual Density Fluctuation Frequency in Mediterranean Pines at a Regional Scale

PubMed Central

Zalloni, Enrica; de Luis, Martin; Campelo, Filipe; Novak, Klemen; De Micco, Veronica; Di Filippo, Alfredo; Vieira, Joana; Nabais, Cristina; Rozas, Vicente; Battipaglia, Giovanna

2016-01-01

Tree rings provide information about the climatic conditions during the growing season by recording them in different anatomical features, such as intra-annual density fluctuations (IADFs). IADFs are intra-annual changes of wood density appearing as latewood-like cells within earlywood, or earlywood-like cells within latewood. The occurrence of IADFs is dependent on the age and size of the tree, and it is triggered by climatic drivers. The variations of IADF frequency of different species and their dependence on climate across a wide geographical range have still to be explored. The objective of this study is to investigate the effect of age, tree-ring width and climate on IADF formation and frequency at a regional scale across the Mediterranean Basin in Pinus halepensis Mill., Pinus pinaster Ait., and Pinus pinea L. The analyzed tree-ring network was composed of P. pinea trees growing at 10 sites (2 in Italy, 4 in Spain, and 4 in Portugal), P. pinaster from 19 sites (2 in Italy, 13 in Spain, and 4 in Portugal), and P. halepensis from 38 sites in Spain. The correlations between IADF frequency and monthly minimum, mean and maximum temperatures, as well as between IADF frequency and total precipitation, were analyzed. A significant negative relationship between IADF frequency and tree-ring age was found for the three Mediterranean pines. Moreover, IADFs were more frequent in wider rings than in narrower ones, although the widest rings showed a reduced IADF frequency. Wet conditions during late summer/early autumn triggered the formation of IADFs in the three species. Our results suggest the existence of a common climatic driver for the formation of IADFs in Mediterranean pines, highlighting the potential use of IADF frequency as a proxy for climate reconstructions with geographical resolution. PMID:27200052

Evidence of floods on the Potomac River from anatomical abnormalities in the wood of flood-plain trees

USGS Publications Warehouse

Yanosky, Thomas M.

1983-01-01

Ash trees along the Potomac River flood plain near Washington, D.C., were studied to determine changes in wood anatomy related to flood damage, and anomalous growth was compared to flood records for April 15 to August 31, 1930-79. Collectively, anatomical evidence was detected for 33 of the 34 growing-season floods during the study period. Evidence of 12 floods prior to 1930 was also noted, including catastrophic ones in 1889 and 1924. Trees damaged after the transition from earlywood to latewood growth typically formed ' flood rings ' of enlarged vessels within the latewood zone. Trees damaged near the beginning of the growth year developed flood rings within, or contiguous with, the earlywood. Both patterns are assumed to have developed when flood-damaged trees produced a second crop of leaves. Trees damaged by high-magnitude floods developed well formed flood rings along the entire height and around the entire circumference of the stem. Small floods were generally associated wtih diffuse or discontinuous anomalies restricted to stem apices. Frequency of flood rings was positively related to flood magnitude, and time of flood generation during the tree-growth season was estimated from the radial position of anomalous growth relative to annual ring width. Reconstructing tree heights in a year of flood-ring formation gives a minimum stage estimate along local stream reaches. Some trees provided evidence of numerous floods. Those with the greatest number of flood rings grew on frequently flooded surfaces subject to flood-flow velocities of at least 1 m/s, and more typically greater than 2 m/s. Tree size, more than age, was related to flood-ring formation. Trees kept small by frequent flood damage had more flood rings than taller trees of comparable age. (USGS)

Chronological Sequence of Leaf Phenology, Xylem and Phloem Formation and Sap Flow of Quercus pubescens from Abandoned Karst Grasslands

PubMed Central

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

Function and Dynamics of Auxin and Carbohydrates during Earlywood/Latewood Transition in Scots Pine1

PubMed Central

Uggla, Claes; Magel, Elisabeth; Moritz, Thomas; Sundberg, Björn

2001-01-01

In temperate regions the annual pattern of wood development is characterized by the formation of radially narrow and thick walled latewood cells. This takes place at the later part of the growing season when cambial cell division declines. To gain new insight into the regulation of this process, micro-analytical techniques were used to visualize the distribution of indole-3-acetic acid (IAA), soluble carbohydrates, and activities of sucrose (Suc)-metabolizing enzymes across the cambial region tissues in Scots pine (Pinus sylvestris). The total amount of IAA in the cambial region did not change with latewood initiation. But its radial distribution pattern was altered, resulting in an increased concentration in the cambial meristem and its recent derivatives. Thus, initiation of latewood formation and cessation of cambial cell division is not a consequence of decreased IAA concentrations in dividing and expanding cells. Rather, IAA most likely has a role in defining the altered developmental pattern associated with latewood formation. Carbohydrates and enzyme activities showed distinctive radial distribution patterns. Suc peaked in the phloem and decreased sharply to low levels across the cambial zone, whereas fructose and glucose reached their highest levels in the maturing tracheids. Suc synthase was the dominating Suc cleaving enzyme with a peak in the secondary wall-forming tracheids and in the phloem. Soluble acid invertase peaked in dividing and expanding cells. Suc-phosphate synthase had its highest activities in the phloem. Activities of cell wall bound invertase were low. The absence of major seasonal variations indicates that carbohydrate availability is not a trigger for latewood initiation. However, steep concentration gradients of the sugars suggest a role for sugar signaling in vascular development. PMID:11299382

A simple program to measure and analyse tree rings using Excel, R and SigmaScan

PubMed Central

Hietz, Peter

2011-01-01

I present a new software that links a program for image analysis (SigmaScan), one for spreadsheets (Excel) and one for statistical analysis (R) for applications of tree-ring analysis. The first macro measures ring width marked by the user on scanned images, stores raw and detrended data in Excel and calculates the distance to the pith and inter-series correlations. A second macro measures darkness along a defined path to identify latewood–earlywood transition in conifers, and a third shows the potential for automatic detection of boundaries. Written in Visual Basic for Applications, the code makes use of the advantages of existing programs and is consequently very economic and relatively simple to adjust to the requirements of specific projects or to expand making use of already available code. PMID:26109835

A simple program to measure and analyse tree rings using Excel, R and SigmaScan.

PubMed

Hietz, Peter

I present a new software that links a program for image analysis (SigmaScan), one for spreadsheets (Excel) and one for statistical analysis (R) for applications of tree-ring analysis. The first macro measures ring width marked by the user on scanned images, stores raw and detrended data in Excel and calculates the distance to the pith and inter-series correlations. A second macro measures darkness along a defined path to identify latewood-earlywood transition in conifers, and a third shows the potential for automatic detection of boundaries. Written in Visual Basic for Applications, the code makes use of the advantages of existing programs and is consequently very economic and relatively simple to adjust to the requirements of specific projects or to expand making use of already available code.

Effects of Invasive Winter Moth Defoliation on Tree Radial Growth in Eastern Massachusetts, USA

PubMed Central

Simmons, Michael J.; Lee, Thomas D.; Ducey, Mark J.; Elkinton, Joseph S.; Boettner, George H.; Dodds, Kevin J.

2014-01-01

Winter moth, Operophtera brumata L. (Lepidoptera: Geometridae), has been defoliating hardwood trees in eastern Massachusetts since the 1990s. Native to Europe, winter moth has also been detected in Rhode Island, Connecticut, eastern Long Island (NY), New Hampshire, and Maine. Individual tree impacts of winter moth defoliation in New England are currently unknown. Using dendroecological techniques, this study related annual radial growth of individual host (Quercus spp. and Acer spp.) trees to detailed defoliation estimates. Winter moth defoliation was associated with up to a 47% reduction in annual radial growth of Quercus trees. Latewood production of Quercus was reduced by up to 67% in the same year as defoliation, while earlywood production was reduced by up to 24% in the year following defoliation. Winter moth defoliation was not a strong predictor of radial growth in Acer species. This study is the first to document impacts of novel invasions of winter moth into New England. PMID:26462685

Changes of wood cell walls in response to hygro-mechanical steam treatment.

PubMed

Guo, Juan; Song, Kunlin; Salmén, Lennart; Yin, Yafang

2015-01-22

The effects of compression combined with steam treatment (CS-treatment), i.e. a hygro-mechanical steam treatment on Spruce wood were studied on a cell-structure level to understand the chemical and physical changes of the secondary cell wall occurring under such conditions. Specially, imaging FT-IR microscopy, nanoindentation and dynamic vapour absorption were used to track changes in the chemical structure, in micromechanical and hygroscopic properties. It was shown that CS-treatment resulted in different changes in morphological, chemical and physical properties of the cell wall, in comparison with those under pure steam treatment. After CS-treatment, the cellular structure displayed significant deformations, and the biopolymer components, e.g. hemicellulose and lignin, were degraded, resulting in decreased hygroscopicity and increased mechanical properties of the wood compared to both untreated and steam treated wood. Moreover, CS-treatment resulted in a higher degree of degradation especially in earlywood compared to a more uniform behaviour of wood treated only by steam. Copyright © 2014 Elsevier Ltd. All rights reserved.

A synchronous increase in hydraulic conductive capacity and mechanical support in conifers with relatively uniform xylem structure.

PubMed

Jagels, Richard; Visscher, George E

2006-02-01

The dual function provided by longitudinal tracheids in conifers has led to a generally held trade-off concept that increasing wall thickness and/or volume of latewood tracheids improves mechanical support, while increasing cell diameter and/or volume of earlywood tracheids enhances conductive potential. Yet, some conifers have either uniform cell structure across the growth ring or, at most, a small amount of latewood. How do these trees accomplish the needs for increasing support and conduction with height growth? We examined Metasequoia glyptostroboides, a species that we previously demonstrated improves its mechanical properties with increasing age without a change in specific gravity or secondary wall microfibril angle. In this paper, we showed that lignin and extractive contents are not contributing factors, and through composite structure analysis, we eliminated a role for tracheid length. Using micromorphometric analysis, we demonstrated that as cell diameter increases, total primary wall decreases, secondary wall increases, and strength and conductive capacity increase with no change in specific gravity. Meta-analysis using other species of Cupressaceae, Podocarpaceae, and Araucariaceae provided strong corroborative evidence for this design strategy.

Paleoclimate Reconstruction at Lamanai, Belize Using Oxygen-Isotope Tropical Dendrochronology

NASA Astrophysics Data System (ADS)

Prentice, A.; Webb, E. A.; White, C. D.; Graham, E.

2009-05-01

Tropical dendrochronology can be complicated because many trees growing in these areas lack distinct visible annual rings. However, the oxygen-isotope composition of wood growing in tropical regions can provide a record of seasonal fluctuations in the amount of precipitation even when visible rings are absent. Variations in the oxygen-isotope compositions of cellulose as the trees grow can be related to the relative timing of wet and dry seasons and used to identify periods of drought. In this study, the oxygen-isotope composition was determined for cellulose extracted from living trees at the site of Lamanai, Belize to assess the variation in oxygen-isotope values that result from heterogeneity within individual tree rings and seasonal fluctuations in amount of precipitation. In temperate regions, the latewood rings that form during periods of reduced growth are traditionally selected for oxygen-isotope analysis of cellulose because their oxygen-isotope compositions are more directly influenced by climate and precipitation during the growing season. However, in tropical isotope dendrochronology, when visible rings are present, detailed sampling of both the light coloured earlywood and the denser latewood is required. At Lamanai, a seasonal signal was evident in the oxygen- isotope composition of the cellulose when tree rings were sectioned in very small increments (approximately every mm), sub-sampling both earlywood and latewood. However, the visible rings did not always correspond with minimum or maximum oxygen-isotope values. As a result, the amplitude of the oxygen-isotope signal obtained by considering only latewood samples is smaller than that obtained from fine-increment sampling. Hence, the oxygen-isotope values of latewood samples alone did not provide accurate data for climate reconstruction. Multiple series of latewood samples extracted from different cross-sections of the same tree did not consistently show the same trends in oxygen isotope values

New star on the stage: amount of ray parenchyma in tree rings shows a link to climate.

PubMed

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.

Refined pipe theory for mechanistic modeling of wood development.

PubMed

Deckmyn, Gaby; Evans, Sam P; Randle, Tim J

2006-06-01

We present a mechanistic model of wood tissue development in response to changes in competition, management and climate. The model is based on a refinement of the pipe theory, where the constant ratio between sapwood and leaf area (pipe theory) is replaced by a ratio between pipe conductivity and leaf area. Simulated pipe conductivity changes with age, stand density and climate in response to changes in allocation or pipe radius, or both. The central equation of the model, which calculates the ratio of carbon (C) allocated to leaves and pipes, can be parameterized to describe the contrasting stem conductivity behavior of different tree species: from constant stem conductivity (functional homeostasis hypothesis) to height-related reduction in stem conductivity with age (hydraulic limitation hypothesis). The model simulates the daily growth of pipes (vessels or tracheids), fibers and parenchyma as well as vessel size and simulates the wood density profile and the earlywood to latewood ratio from these data. Initial runs indicate the model yields realistic seasonal changes in pipe radius (decreasing pipe radius from spring to autumn) and wood density, as well as realistic differences associated with the competitive status of trees (denser wood in suppressed trees).

Intra- and Inter-Annual Variability in Surface Hydrology in Northern Arizona from δ18O of Tree-Ring Cellulose

NASA Astrophysics Data System (ADS)

Whittaker, T. E.; Galewsky, J.; Scuderi, L. A.; Sharp, Z. D.

2010-12-01

The primary goal of our research is to better understand how the surface hydrology of semi-arid sites in the Southwestern U.S. is affected by the annual cycles of precipitation and evaporation. Both are tied to relative strength of the North American Monsoon, El Niño-Southern Oscillation and, on shorter timescales, the occasional passage of tropical cyclone remnants. To achieve this we aim to develop high-resolution stable oxygen isotope ratio (δ18O) profiles of tree-ring cellulose for much of the last 20 years that can be ground-truthed to direct meteorological observations. It is well documented that δ18O of alpha-cellulose extracted from wood reflects hydrological conditions of a trees’ environment at the time the tree grew. Primary controls on isotopic variability are changes in source waters and relative humidity during the growing season. We sampled rings from ≥ 10 Pinus ponderosa (Douglas) at six stands along an east-west transect across northern Arizona. Annual precipitation at these sites has a bimodal distribution with almost all annual rainfall occurring during the summer monsoon (Jul, Aug) and winter storms. At Flagstaff, in the center of our study area, monthly mean precipitation δ18O values are enriched ~6‰ during the monsoon relative to winter storms. P. ponderosa (Dougl.) rings display distinct early- and latewood bands. Earlywood typically forms using winter storm precipitation that has resided within the soil until the tree began growing and ought to reflect the isotopic composition of this water. Latewood δ18O reportedly reflect summer rainfall isotopic values. We investigate the eleven year period 1994-2004. This range encompasses the transition into the present ‘drought’, the intense 1997/98 El Niño, and the passage of the remnants of Hurricanes Nora (1997) and Javier (2004). Individual rings are sliced into subsamples of mass ~1.5 mg (yielding 3-13 samples/ring). Early isotopic data from these samples display three significant

Effect of growth ring orientation and placement of earlywood and latewood on MOE and MOR of very-small clear Douglas-fir beams.

Treesearch

Amy T. Grotta; Robert J. Leichti; Barbara L. Gartner; G.R. Johnson

2005-01-01

ASTM standard sizes for bending tests (either 50 x 50 mm or 25 x 25 mm in cross-section) are not always suitable for research purposes that characterize smaller sections of wood. Moreover, the ASTM standards specify loading the sample on the longitudinal-tangential surface. If specimens are small enough, then the effects of both growth-ring orientation and whether...

Environmental Biomonitoring of Cr and As in Shallow Groundwater: Do Red Oak Trees Preserve Long-Term Records of Contaminant Loading?

NASA Astrophysics Data System (ADS)

Shailer, M.; Brabander, D.

2005-05-01

The use of dendrochemical analysis has been shown to be a valuable, although controversial, tool in monitoring historical trends in trace metal deposition and mobilization in groundwater and sediments. Neutron activation analysis (NAA) is one method that has been used to determine annual dendrochemical patterns in tree rings. The use of NAA may also provide a practical tool for revealing sub-annual differences in metal concentrations between earlywood and latewood. In a variety of geochemical settings, Cr and As can be mobile in the groundwater-root environment and are subsequently taken up by trees and stored in xylem tissues specifically associated with groundwater transport. For the purposes of determining historical patterns in Cr and As bioavailability at a Woburn, MA, superfund site along the Aberjona River, Quercus rubra (red oak) sectioned tree rings were analyzed. Sub-annual dendrochemical analyses were used to identify different As and Cr loading pathways in oak stem wood. A sixty-year record of [As] and [Cr] in stem wood was obtained, and results suggest seasonally dependent correlations with Aberjona River flow and with pumping rates for a municipal well in close proximity to the sampling location. These two hydrological pathways likely dominate in providing a flux of dissolved As and Cr into oak stem wood.

Trees, History, and Isotopes - the Late Maunder Minimum (1675-1715) in the Pannonian Basin, Hungary

NASA Astrophysics Data System (ADS)

Kazmer, M.; Demeny, A.; Grynaeus, A.; Racz, L.; Varkonyi, A.

2002-05-01

First results of a comprehensive study on climate change in the Pannonian Basin during the Late Maunder Minimum (1675-1715) are presented. The Pannonian Basin has continental climate, distinctly warm and dry in summer, cold in winter, unlike the Atlantic-type climate of Western Europe. Surrounded by the arc of the Carpathians, exposed to Atlantic, Mediterranean, and Siberian influences, the regional climate displays steep gradients. More than one tree-ring chronology for oak is being built, independent of the south German series. Rethly's rich database of historical sources has been assembled, and completed with recently published letters. Ring-width series are measured on oak, and skeleton plots are logged. Study of hydrogen isotope composition of tree rings is in progress. Tree-ring width faithfully reflects historical indices on spring (i.e. earlywood growth season) precipitation. Generally, precipitation - as shown both by indices and tree-ring width - was high and temperature low during the growth season in the first half of the LMM. The second half has seen a retardation in oak growth and an increase in spring temperature. The decades of the Late Maunder Minimum was a politically turbulent era: it saw the decline and fall of the Ottoman domination in Hungary, followed by a rebellion against Austrian rule, associated with disruption of national economy.

A 2100-Year Reconstruction of July Rainfall Over Westcentral New Mexico

NASA Astrophysics Data System (ADS)

Stahle, D.; Cleaveland, M.; Therrell, M.; Grissino-Mayer, H.; Griffin, D.; Fye, F.

2007-05-01

We have developed a new 2,141-year long tree-ring chronology of latewood (LW) width from ancient Douglas-fir (Pseudotsuga menziesii) and ponderosa pine (Pinus ponderosae) at El Malpais National Monument, New Mexico. This is one of the longest precipitation-sensitive tree-ring chronologies yet constructed for the American Southwest and has been used to develop the first continuous multi-millennial tree-ring reconstruction of July precipitation in the region of the North American Monsoon System (NAMS). Monthly average precipitation increases sharply in July over western New Mexico, marking the dramatic onset to the summer monsoon season. The LW chronology explains 44 percent of the interannual variability of July precipitation in the instrumental record for New Mexico climate divisions 1 and 4 (1960-2004), after removal of the linear dependence of LW width on earlywood width following Meko and Baisan (2001), and has passed statistical tests of verification on independent July precipitation data (1895-1959). The instrumental and tree-ring reconstructed July precipitation data are correlated with the concurrent 500 mb height field over western North America and with the sea surface temperature gradient from the central to eastern North Pacific. The reconstruction exhibits several severe sustained July droughts that exceed any witnessed during the instrumental era, and has significant spectral power at periods near 3-5, 20, and 70 years.

Impact of solar activity on growth of pine trees (Pinus cembra: 1610 - 1970; Pinus pinaster: 1910 -1989)

NASA Astrophysics Data System (ADS)

Surový, P.; Dorotovič, I.; Karlovský, V.; Rodrigues, J. C.; Rybanský, M.; Fleischer, P.

2010-12-01

In this work we have focused on the analysis of the data on the annual growth of cembra pine (Pinus cembra) grown in the Kôprová dolina Valley in the High Tatra Mountains. The database covers the period of 1406 - 1970, however, the sunspot data (minima and maxima) at the NGDC web site are only available since 1610. Moreover, reliable sunspot data are only available since 1749. The results of this analysis agree with the observation made in our previous work, i.e. there is a negative impact of high SA on the pine tree growth. However, it should be noted that statistical significance of the results is low. We also applied wavelet analysis to the data on the tree growth evolution, with the results indicating growth variations' period of about 20 years (duration of approximately two solar cycles or one magnetic cycle, respectively). A negative impact of the SA was also observed in growth of a 90 year-old maritime pine tree (Pinus pinaster) grown in northern Portugal. The width of the annual rings was smaller in the years of maximum SA; furthermore, it was found that it is the latewood growth that it is affected while the earlywood growth is not, and consequently the latewood additions also show a significative negative correlation with SA.

Quantifications of dendrochronological information from contrasting microdensitometric measuring circumstances of experimental wood samples.

PubMed

Helama, S; Bégin, Y; Vartiainen, M; Peltola, H; Kolström, T; Meriläinen, J

2012-06-01

We analyzed how the pretreatment method of Scots pine (Pinus sylvestris L.) wood specimens together with X-ray methodology applied for density analyses affect resulting tree-ring data and derived proxy-based climate information. We also evaluated whether these results from two contrasting laboratory circumstances could be homogenized by applying dendroclimatic statistical methods. For this study, we measured a pair of X-ray based microdensitometry datasets using double samples of subfossil and recent wood specimens. Dendrochronological information of earlywood and latewood series was examined to determine for alterations in the resulting data. We found that the level of overall density, its trend over cambial ages and the growth amplitude altered due to the sample pretreatment/density measuring exercise, which means that comparisons of heterogeneous datasets should be, in general, regarded cautiously. Dendrochronological standardization did, however, even out several potentially biasing influences from the differing overall densities and their trends. The two latewood (maximum) density chronologies yielded paleoclimatic reconstructions which both calibrated and verified satisfactorily with the instrumental warm-season (March-September) mean temperatures. The transfer functions were found to further equalize the differences between the two proxy records. We recommend (if no strictly homogenous data are available) reconciling similar data assemblages through transfer functions with multiple independent variables. Copyright © 2012 Elsevier Ltd. All rights reserved.

Modular organization of the white spruce (Picea glauca) transcriptome reveals functional organization and evolutionary signatures.

PubMed

Raherison, Elie S M; Giguère, Isabelle; Caron, Sébastien; Lamara, Mebarek; MacKay, John J

2015-07-01

Transcript profiling has shown the molecular bases of several biological processes in plants but few studies have developed an understanding of overall transcriptome variation. We investigated transcriptome structure in white spruce (Picea glauca), aiming to delineate its modular organization and associated functional and evolutionary attributes. Microarray analyses were used to: identify and functionally characterize groups of co-expressed genes; investigate expressional and functional diversity of vascular tissue preferential genes which were conserved among Picea species, and identify expression networks underlying wood formation. We classified 22 857 genes as variable (79%; 22 coexpression groups) or invariant (21%) by profiling across several vegetative tissues. Modular organization and complex transcriptome restructuring among vascular tissue preferential genes was revealed by their assignment to coexpression groups with partially overlapping profiles and partially distinct functions. Integrated analyses of tissue-based and temporally variable profiles identified secondary xylem gene networks, showed their remodelling over a growing season and identified PgNAC-7 (no apical meristerm (NAM), Arabidopsis transcription activation factor (ATAF) and cup-shaped cotyledon (CUC) transcription factor 007 in Picea glauca) as a major hub gene specific to earlywood formation. Reference profiling identified comprehensive, statistically robust coexpressed groups, revealing that modular organization underpins the evolutionary conservation of the transcriptome structure. © 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.

[Cedrela odorata (Meliaceae) potential for dendrochronological studies in the Selva Central of Perú].

PubMed

Pereyra Espinoza, Manuel Jesús; Inga Guillen, Gaby Janet; Santos Morales, Mariano; Arisméndiz Rodríguez, Rodolfo

2014-06-01

Despite the progress made during the past 20 years, searching dendrochronological potential in tropical and subtropical tree species, tropical dendrochronology, is still in a development stage. The aim of this research was to determine the potential of C. odorata for dendrochronological studies in the Selva Central of Perú. The tree-ring anatomical characteristics were carefully examined and we were able to develop a 215 year (1795-2 009) tree-ring chronology and correlate it with precipitation records. The tree-ring chronology was developed based on 47 series of 27 trees. Tree rings are clearly delimited by large pore diameters in earlywood and small ones in latewood associated with marginal and paratracheal parenchyma. The tree-ring chronology was related to precipitation records from Satipo and significant correlations were found with the previous rainy season and late dry season of the current growth period. Moreover, we found close relationship between tree growth and total precipitations of the hydrological period (December to September) for the interval 1990-2009. These results demonstrate the influence of rainfall at different stages of C. odorata radial growth. The good discrimination of annual rings, strong relationship with precipitation, the wide range and longevity of trees (200 years) make C. odorata a very promising species for dendrochronological studies in tropical and subtropical forest of America.

Changes in leaf organisation, photosynthetic performance and wood formation during ex vitro acclimatisation of black mulberry (Morus nigra L.).

PubMed

Misalová, A; Durkovic, J; Mamonová, M; Priwitzer, T; Lengyelová, A; Hladká, D; Lux, A

2009-09-01

Changes in anatomical organisation of the leaf, photosynthetic performance and wood formation were examined to evaluate the temporal and spatial patterns of acclimatisation of micropropagated slow-growing black mulberry (Morus nigra L.) plantlets to the ex vitro environment. Leaf structure differentiation, the rates of net photosynthesis (P(n)), transpiration (E) and stomatal conductance (g(s)), and secondary xylem growth were determined in the course of a 56-day acclimatisation. Differentiation of palisade parenchyma was observed 7 days after transfer. At this stage, the rates of P(n), E and g(s) reached maximum values, after which the rates of all three gas exchange parameters gradually decreased. The highest proportion of woody area occupied by vessels was also observed 7 days after transfer. An important feature of developing woody tissue is the difference in patterns of vessel distribution from the characteristic differentiation patterns of earlywood and latewood vessels in mature wood of ring-porous trees. Vessels with lumen areas over 3000 microm(2) were only differentiated in acclimatised plantlets, whereas vessels in stems sampled on days 0 and 7 had very small lumen areas of up to 560 microm(2). Full acclimatisation, observed 56 days after transfer to the ex vitro environment, was associated with the rapid growth of new in vivo formed leaves, very low rates of E and g(s), and much increased secondary xylem tissue within the stem area.

New paleoclimatic database for the Iberian Peninsula since AD 1700 inferred from tree-ring records and documentary evidence: advances in temperature and drought variability reconstructions

NASA Astrophysics Data System (ADS)

Tejedor, Ernesto; Ángel Saz, Miguel; de Luis, Martín; Esper, Jan; Barriendos, Mariano; Serrano-Notivoli, Roberto; Novak, Klemen; Longares, Luis Alberto; Martínez-del Castillo, Edurne; María Cuadrat, José

2017-04-01

A substantial increase of surface air temperatures in the upcoming decades, particularly significant in the Mediterranean basin, has been reported by the IPCC (IPCC, 2013). It is therefore particularly important to study past climate extremes and variability in this region, which will in turn support the accuracy of future climate scenarios. Yet, our knowledge of past climate variability and trends is limited by the shortage of instrumental data prior to the twentieth century, which prompts to the need of discovering new sources with which to reconstruct past climate. We here present a new paleoclimatic database for the northeast of the Iberian Peninsula based on tree-ring records, documentary evidence and instrumental data. The network includes 774 tree-ring, earlywood and latewood width series from Pinus uncinata, Pinus sylvestris and Pinus nigra trees in the Pyrenees and Iberian Range reaching back to AD 1510. Three reconstructions are developed using these samples; an annual drought reconstruction since AD 1694, a summer drought reconstruction since AD 1734, and a maximum temperature reconstruction since AD 1604. Additionally, the documentary records from 16 locations in the Ebro Valley are examined focusing on climate-related 'rogations'. We differentiated three types of rogations, considering the importance of religious acts, to identify the severity of drought and pluvial events. Finally, an attempt to explore the links between documentary and tree-ring based reconstructions is presented.

Tree-ring analysis of ancient baldcypress trees and subfossil wood

NASA Astrophysics Data System (ADS)

Stahle, David W.; Burnette, Dorian J.; Villanueva, Jose; Cerano, Julian; Fye, Falko K.; Griffin, R. Daniel; Cleaveland, Malcolm K.; Stahle, Daniel K.; Edmondson, Jesse R.; Wolff, Kathryn P.

2012-02-01

Ancient baldcypress trees found in wetland and riverine environments have been used to develop a network of exactly dated annual ring-width chronologies extending from the southeastern United States, across Mexico, and into western Guatemala. These chronologies are sensitive to growing season precipitation in spite of frequently flooded site conditions, and have been used to reconstruct moisture levels the southeastern United States and Mexico for over 1000 years. The El Nino/Southern Oscillation (ENSO) is a major influence on the climate reconstructions derived from these baldcypress chronologies, especially in Mexico where some of the most extreme reconstructed droughts occurred during El Nino events. In the Southeast, the ENSO influence on climate and tree growth changes sign from spring to summer, and this change in dynamical forcing is recorded by sub-seasonal chronologies of earlywood and latewood width. Most existing baldcypress chronologies have been extended with tree-ring data from "subfossil" wood recovered from surface and submerged deposits. Well-preserved subfossil logs have also been recovered in quantity from buried deposits of great age, and may permit development of long continuously dated Holocene chronologies and discontinuous "floating" Pleistocene chronologies. The extensive subfossil baldcypress swamp discovered 6 m below the streets of Washington D.C. was overrun by a transgression of the Potomac estuary, possibly during the previous super interglacial (marine OIS 5e), and provides direct evidence for one potential impact of unmitigated anthropogenic warming and sea level rise.

Association Genetics of Wood Physical Traits in the Conifer White Spruce and Relationships With Gene Expression

PubMed Central

Beaulieu, Jean; Doerksen, Trevor; Boyle, Brian; Clément, Sébastien; Deslauriers, Marie; Beauseigle, Stéphanie; Blais, Sylvie; Poulin, Pier-Luc; Lenz, Patrick; Caron, Sébastien; Rigault, Philippe; Bicho, Paul; Bousquet, Jean; MacKay, John

2011-01-01

Marker-assisted selection holds promise for highly influencing tree breeding, especially for wood traits, by considerably reducing breeding cycles and increasing selection accuracy. In this study, we used a candidate gene approach to test for associations between 944 single-nucleotide polymorphism markers from 549 candidate genes and 25 wood quality traits in white spruce. A mixed-linear model approach, including a weak but nonsignificant population structure, was implemented for each marker–trait combination. Relatedness among individuals was controlled using a kinship matrix estimated either from the known half-sib structure or from the markers. Both additive and dominance effect models were tested. Between 8 and 21 single-nucleotide polymorphisms (SNPs) were found to be significantly associated (P ≤ 0.01) with each of earlywood, latewood, or total wood traits. After controlling for multiple testing (Q ≤ 0.10), 13 SNPs were still significant across as many genes belonging to different families, each accounting for between 3 and 5% of the phenotypic variance in 10 wood characters. Transcript accumulation was determined for genes containing SNPs associated with these traits. Significantly different transcript levels (P ≤ 0.05) were found among the SNP genotypes of a 1-aminocyclopropane-1-carboxylate oxidase, a β-tonoplast intrinsic protein, and a long-chain acyl-CoA synthetase 9. These results should contribute toward the development of efficient marker-assisted selection in an economically important tree species. PMID:21385726

Wood properties of Scots pines (Pinus sylvestris) grown at elevated temperature and carbon dioxide concentration.

PubMed

Kilpeläinen, Antti; Peltola, Heli; Ryyppö, Aija; Sauvala, Kari; Laitinen, Kaisa; Kellomäki, Seppo

2003-09-01

Impacts of elevated temperature and carbon dioxide concentration ([CO2]) on wood properties of 15-year-old Scots pines (Pinus sylvestris L.) grown under conditions of low nitrogen supply were investigated in open-top chambers. The treatments consisted of (i) ambient temperature and ambient [CO2] (AT+AC), (ii) ambient temperature and elevated [CO2] (AT+EC), (iii) elevated temperature and ambient [CO2] (ET+AC) and (iv) elevated temperature and elevated [CO2] (ET+EC). Wood properties analyzed for the years 1992-1994 included ring width, early- and latewood width and their proportions, intra-ring wood density (minimum, maximum and mean, as well as early- and latewood densities), mean fiber length and chemical composition of the wood (cellulose, hemicellulose, lignin and acetone extractive concentration). Absolute radial growth over the 3-year period was 54% greater in AT+EC trees and 30 and 25% greater in ET+AC and ET+EC trees, respectively, than in AT+AC trees. Neither elevated temperature nor elevated [CO2] had a statistically significant effect on ring width, early- and latewood widths or their proportions. Both latewood density and maximum intra-ring density were increased by elevated [CO2], whereas fiber length was increased by elevated temperature. Hemicellulose concentration decreased and lignin concentration increased significantly in response to elevated temperature. There were no statistically significant interaction effects of elevated temperature and elevated [CO2] on the wood properties, except on earlywood density.

Wood anatomy and carbon-isotope discrimination support long-term hydraulic deterioration as a major cause of drought-induced dieback.

PubMed

Pellizzari, Elena; Camarero, J Julio; Gazol, Antonio; Sangüesa-Barreda, Gabriel; Carrer, Marco

2016-06-01

Hydraulic impairment due to xylem embolism and carbon starvation are the two proposed mechanisms explaining drought-induced forest dieback and tree death. Here, we evaluate the relative role played by these two mechanisms in the long-term by quantifying wood-anatomical traits (tracheid size and area of parenchyma rays) and estimating the intrinsic water-use efficiency (iWUE) from carbon isotopic discrimination. We selected silver fir and Scots pine stands in NE Spain with ongoing dieback processes and compared trees showing contrasting vigour (declining vs nondeclining trees). In both species earlywood tracheids in declining trees showed smaller lumen area with thicker cell wall, inducing a lower theoretical hydraulic conductivity. Parenchyma ray area was similar between the two vigour classes. Wet spring and summer conditions promoted the formation of larger lumen areas, particularly in the case of nondeclining trees. Declining silver firs presented a lower iWUE than conspecific nondeclining trees, but the reverse pattern was observed in Scots pine. The described patterns in wood anatomical traits and iWUE are coherent with a long-lasting deterioration of the hydraulic system in declining trees prior to their dieback. Retrospective quantifications of lumen area permit to forecast dieback in declining trees 2-5 decades before growth decline started. Wood anatomical traits provide a robust tool to reconstruct the long-term capacity of trees to withstand drought-induced dieback. © 2016 John Wiley & Sons Ltd.

The Tunguska event in 1908: evidence from tree-ring anatomy.

PubMed

Vaganov, Evgenii A; Hughes, Malcolm K; Silkin, Pavel P; Nesvetailo, Valery D

2004-01-01

We analyzed tree rings in wood samples collected from some of the few surviving trees found close to the epicenter (within 4-5 km) of the Tunguska event that occurred on the last day of June 1908. Tree-ring growth shows a depression starting in the year after the event and continuing during a 4-5-year period. The most remarkable traces of the event were found in the rings' anatomical structure: (1) formation of "light" rings and a reduction of maximum density in 1908; (2) non-thickened tracheids (the cells that make up most of the wood volume) in the transition and latewood zones (the middle and last-formed parts of the ring, respectively); and (3) deformed tracheids, which are located on the 1908 annual ring outer boundary. In the majority of samples, normal earlywood and latewood tracheids were formed in all annual rings after 1908. The observed anomalies in wood anatomy suggest two main impacts of the Tunguska event on surviving trees--(1) defoliation and (2) direct mechanical stress on active xylem tissue. The mechanical stress needed to fell trees is less than the stress needed to cause the deformation of differentiating tracheids observed in trees close to the epicenter. In order to resolve this apparent contradiction, work is suggested on possible topographic modification of the overpressure experienced by these trees, as is an experimental test of the effects of such stresses on precisely analogous growing trees.

Nine Hundred Years of Weekly Streamflows: Stochastic Downscaling of Ensemble Tree-Ring Reconstructions

NASA Astrophysics Data System (ADS)

Sauchyn, David; Ilich, Nesa

2017-11-01

We combined the methods and advantages of stochastic hydrology and paleohydrology to estimate 900 years of weekly flows for the North and South Saskatchewan Rivers at Edmonton and Medicine Hat, Alberta, respectively. Regression models of water-year streamflow were constructed using historical naturalized flow data and a pool of 196 tree-ring (earlywood, latewood, and annual) ring-width chronologies from 76 sites. The tree-ring models accounted for up to 80% of the interannual variability in historical naturalized flows. We developed a new algorithm for generating stochastic time series of weekly flows constrained by the statistical properties of both the historical record and proxy streamflow data, and by the necessary condition that weekly flows correlate between the end of a year and the start of the next. A second innovation, enabled by the density of our tree-ring network, is to derive the paleohydrology from an ensemble of 100 statistically significant reconstructions at each gauge. Using paleoclimatic data to generate long series of weekly flow estimates augments the short historical record with an expanded range of hydrologic variability, including sequences of wet and dry years of greater length and severity. This unique hydrometric time series will enable evaluation of the reliability of current water supply and management systems given the range of hydroclimatic variability and extremes contained in the stochastic paleohydrology. It also could inform evaluation of the uncertainty in climate model projections, given that internal hydroclimatic variability is the dominant source of uncertainty.

Assessment of Methods to Determine Tree Ring Response to Large Magnitude Mississippi River Floods

NASA Astrophysics Data System (ADS)

Therrell, M. D.; Meko, M. D.; Bialecki, M.; Remo, J. W.

2017-12-01

Riparian trees that experience prolonged inundation can record major flood events as inter-and intra-annual variability in size, shape and arrangement of vessels in the annual xylem growth increment. As part of an NSF-funded project to develop tree-ring records of past flooding, we have made collections of several oak species (e.g., Quercus lyrata, Q. macrocarpa) at six sites in the Mississippi River Basin. At each of these sites sampled trees exhibit notably anomalous anatomy of growth increments formed in years coinciding with major recorded floods. We have used these "flood rings" to develop individual site chronologies as well as a regional chronology of spring flood events in the basin for the past several hundred years. We have also analyzed earlywood vessel diameter as a proxy for flooding and find that although this variable reflects only a fraction of the annual-growth increment it strongly reflects tree response to flooding at all the sites so far examined. We compare both these chronologies with the instrumental and historical record of flooding and find that our chronologies are recording nearly all large observed Mississippi River floods in the 20th century, and provide a new record of similar events in the 18th and 19th centuries. These results suggest that tree-rings can be effectively used to develop and improve pre-instrumental flood records throughout the basin and potentially other similar systems.

Growth response and sapwood hydraulic properties of young lodgepole pine following repeated fertilization.

PubMed

Amponsah, Isaac G; Lieffers, Victor J; Comeau, Philip G; Brockley, Robert P

2004-10-01

We examined how tree growth and hydraulic properties of branches and boles are influenced by periodic (about 6 years) and annual fertilization in two juvenile lodgepole pine (Pinus contorta Dougl. var. latifolia Engelm.) stands in the interior of British Columbia, Canada. Mean basal area (BA), diameter at breast height (DBH) and height increments and percent earlywood and sapwood hydraulic parameters of branches and boles were measured 7 or 8 years after the initial treatments at Sheridan Creek and Kenneth Creek. At Sheridan Creek, fertilization significantly increased BA and DBH increments, but had no effect on height increment. At Kenneth Creek, fertilization increased BA, but fertilized trees had significantly lower height increments than control trees. Sapwood permeability was greater in lower branches of repeatedly fertilized trees than in those of control trees. Sapwood permeabilities of the lower branches of trees in the control, periodic and annual treatments were 0.24 x 10(-12), 0.35 x 10(-12) and 0.45 x 10(-12) m2 at Kenneth Creek; and 0.41 x 10(-12), 0.54 x 10(-12) and 0.65 x 10(-12) m2 at Sheridan Creek, respectively. Annual fertilization tended to increase leaf specific conductivities and Huber values of the lower branches of trees at both study sites. We conclude that, in trees fertilized annually, the higher flow capacity of lower branches may reduce the availability of water to support annual growth of the leader and upper branches.

Spatial distribution of xylem embolisms in the stems of Pinus thunbergii at the threshold of fatal drought stress.

PubMed

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.

Size Matters a Lot: Drought-Affected Italian Oaks Are Smaller and Show Lower Growth Prior to Tree Death

PubMed Central

Colangelo, Michele; Camarero, Jesús J.; Borghetti, Marco; Gazol, Antonio; Gentilesca, Tiziana; Ripullone, Francesco

2017-01-01

Hydraulic theory suggests that tall trees are at greater risk of drought-triggered death caused by hydraulic failure than small trees. In addition the drop in growth, observed in several tree species prior to death, is often interpreted as an early-warning signal of impending death. We test these hypotheses by comparing size, growth, and wood-anatomy patterns of living and now-dead trees in two Italian oak forests showing recent mortality episodes. The mortality probability of trees is modeled as a function of recent growth and tree size. Drift-diffusion-jump (DDJ) metrics are used to detect early-warning signals. We found that the tallest trees of the anisohydric Italian oak better survived drought contrary to what was predicted by the theory. Dead trees were characterized by a lower height and radial-growth trend than living trees in both study sites. The growth reduction of now-dead trees started about 10 years prior to their death and after two severe spring droughts during the early 2000s. This critical transition in growth was detected by DDJ metrics in the most affected site. Dead trees were also more sensitive to drought stress in this site indicating different susceptibility to water shortage between trees. Dead trees did not form earlywood vessels with smaller lumen diameter than surviving trees but tended to form wider latewood vessels with a higher percentage of vessel area. Since living and dead trees showed similar competition we did not expect that moderate thinning and a reduction in tree density would increase the short-term survival probability of trees. PMID:28270816

Xylogenesis: Coniferous Trees of Temperate Forests Are Listening to the Climate Tale during the Growing Season But Only Remember the Last Words!1

PubMed Central

2016-01-01

The complex inner mechanisms that create typical conifer tree-ring structure (i.e. the transition from large, thin-walled earlywood cells to narrow, thick-walled latewood cells) were recently unraveled. However, what physiological or environmental factors drive xylogenesis key processes remain unclear. Here, we aim to quantify the influence of seasonal variations in climatic factors on the spectacular changes in the kinetics of wood cell differentiation and in the resulting tree-ring structure. Wood formation was monitored in three sites over 3 years for three coniferous species (Norway spruce [Picea abies], Scots pine [Pinus sylvestris], and silver fir [Abies alba]). Cell differentiation rates and durations were calculated and related to tracheid final dimensions and corresponding climatic conditions. On the one hand, we found that the kinetics of cell enlargement and the final size of the tracheids were not explained by the seasonal changes in climatic factors. On the other hand, decreasing temperatures strongly constrained cell wall deposition rates during latewood formation. However, the influence of temperature was permanently written into tree-ring structure only for the very last latewood cells, when the collapse of the rate of wall deposition was no longer counterbalanced by the increase of its duration. Our results show that the formation of the typical conifer tree-ring structure, in normal climatic conditions, is only marginally driven by climate, suggesting strong developmental control of xylogenesis. The late breakage of the compensatory mechanism at work in the wall deposition process appears as a clue to understand the capacity of the maximum latewood density to record past temperature conditions. PMID:27208048

Functional Multi-Locus QTL Mapping of Temporal Trends in Scots Pine Wood Traits

PubMed Central

Li, Zitong; Hallingbäck, Henrik R.; Abrahamsson, Sara; Fries, Anders; Gull, Bengt Andersson; Sillanpää, Mikko J.; García-Gil, M. Rosario

2014-01-01

Quantitative trait loci (QTL) mapping of wood properties in conifer species has focused on single time point measurements or on trait means based on heterogeneous wood samples (e.g., increment cores), thus ignoring systematic within-tree trends. In this study, functional QTL mapping was performed for a set of important wood properties in increment cores from a 17-yr-old Scots pine (Pinus sylvestris L.) full-sib family with the aim of detecting wood trait QTL for general intercepts (means) and for linear slopes by increasing cambial age. Two multi-locus functional QTL analysis approaches were proposed and their performances were compared on trait datasets comprising 2 to 9 time points, 91 to 455 individual tree measurements and genotype datasets of amplified length polymorphisms (AFLP), and single nucleotide polymorphism (SNP) markers. The first method was a multilevel LASSO analysis whereby trend parameter estimation and QTL mapping were conducted consecutively; the second method was our Bayesian linear mixed model whereby trends and underlying genetic effects were estimated simultaneously. We also compared several different hypothesis testing methods under either the LASSO or the Bayesian framework to perform QTL inference. In total, five and four significant QTL were observed for the intercepts and slopes, respectively, across wood traits such as earlywood percentage, wood density, radial fiberwidth, and spiral grain angle. Four of these QTL were represented by candidate gene SNPs, thus providing promising targets for future research in QTL mapping and molecular function. Bayesian and LASSO methods both detected similar sets of QTL given datasets that comprised large numbers of individuals. PMID:25305041

Latitudinal Gradients in Tree Ring Stable Carbon and Oxygen Isotopes Reveal Differential Climate Influences of the North American Monsoon System.

NASA Astrophysics Data System (ADS)

Szejner, P.; Wright, W. E.; Babst, F.; Belmecheri, S.; Trouet, V.; Leavitt, S. W.; Ehleringer, J. R.; Monson, R. K.

2016-12-01

The arrival of the North American Monsoon System (NAMS) terminates a presummer hyperarid period in the southwestern United States (U.S.), providing summer moisture that is favorable for forest growth. Montane forests in this region rely on winter snowpack to drive much of their growth; the extent to which they use NAMS moisture is uncertain. We addressed this by studying stable carbon and oxygen isotopes in earlywood and latewood from 11 sites along a latitudinal gradient extending from Arizona and New Mexico to Utah. This study provides the first regional perspective on the relative roles of winter versus summer precipitation as an ecophysiological resource. Here we present evidence that Ponderosa pine uses NAMS moisture differentially across this gradient. 13C/12C ratios suggest that photosynthetic water use efficiency during latewood formation is more sensitive to summer precipitation at the northern than at the southern sites. This is likely due to the fact that NAMS moisture provides sufficiently favorable conditions for tree photosynthesis and growth during most years in the southern sites, whereas the northern sites experience larger summer moisture variability, which in some years is limiting growth. Cellulose δ18O and δ13C values revealed that photoassimilates in the southern sites were produced under higher vapor pressure deficit conditions during spring compared to summer, demonstrating a previously underappreciated effect of seasonal differences in atmospheric humidity on tree ring isotope ratios. Our findings suggest that future changes in NAMS will potentially alter productivity and photosynthetic water use dynamics differentially along latitudinal gradients in southwestern U.S. montane forests.

Latitudinal gradients in tree ring stable carbon and oxygen isotopes reveal differential climate influences of the North American Monsoon System

NASA Astrophysics Data System (ADS)

Szejner, Paul; Wright, William E.; Babst, Flurin; Belmecheri, Soumaya; Trouet, Valerie; Leavitt, Steven W.; Ehleringer, James R.; Monson, Russell K.

2016-07-01

The arrival of the North American Monsoon System (NAMS) terminates a presummer hyperarid period in the southwestern United States (U.S.), providing summer moisture that is favorable for forest growth. Montane forests in this region rely on winter snowpack to drive much of their growth; the extent to which they use NAMS moisture is uncertain. We addressed this by studying stable carbon and oxygen isotopes in earlywood and latewood from 11 sites along a latitudinal gradient extending from Arizona and New Mexico to Utah. This study provides the first regional perspective on the relative roles of winter versus summer precipitation as an ecophysiological resource. Here we present evidence that Ponderosa pine uses NAMS moisture differentially across this gradient. 13C/12C ratios suggest that photosynthetic water use efficiency during latewood formation is more sensitive to summer precipitation at the northern than at the southern sites. This is likely due to the fact that NAMS moisture provides sufficiently favorable conditions for tree photosynthesis and growth during most years in the southern sites, whereas the northern sites experience larger summer moisture variability, which in some years is limiting growth. Cellulose δ18O and δ13C values revealed that photoassimilates in the southern sites were produced under higher vapor pressure deficit conditions during spring compared to summer, demonstrating a previously underappreciated effect of seasonal differences in atmospheric humidity on tree ring isotope ratios. Our findings suggest that future changes in NAMS will potentially alter productivity and photosynthetic water use dynamics differentially along latitudinal gradients in southwestern U.S. montane forests.

Combining wood anatomy and stable isotope variations in a 600-year multi-parameter climate reconstruction from Corsican black pine

NASA Astrophysics Data System (ADS)

Szymczak, Sonja; Hetzer, Timo; Bräuning, Achim; Joachimski, Michael M.; Leuschner, Hanns-Hubert; Kuhlemann, Joachim

2014-10-01

We present a new multi-parameter dataset from Corsican black pine growing on the island of Corsica in the Western Mediterranean basin covering the period AD 1410-2008. Wood parameters measured include tree-ring width, latewood width, earlywood width, cell lumen area, cell width, cell wall thickness, modelled wood density, as well as stable carbon and oxygen isotopes. We evaluated the relationships between different parameters and determined the value of the dataset for climate reconstructions. Correlation analyses revealed that carbon isotope ratios are influenced by cell parameters determining cell size, whereas oxygen isotope ratios are influenced by cell parameters determining the amount of transportable water in the xylem. A summer (June to August) precipitation reconstruction dating back to AD 1185 was established based on tree-ring width. No long-term trends or pronounced periods with extreme high/low precipitation are recorded in our reconstruction, indicating relatively stable moisture conditions over the entire time period. By comparing the precipitation reconstruction with a summer temperature reconstruction derived from the carbon isotope chronologies, we identified summers with extreme climate conditions, i.e. warm-dry, warm-wet, cold-dry and cold-wet. Extreme climate conditions during summer months were found to influence cell parameter characteristics. Cold-wet summers promote the production of broad latewood composed of wide and thin-walled tracheids, while warm-wet summers promote the production of latewood with small thick-walled cells. The presented dataset emphasizes the potential of multi-parameter wood analysis from one tree species over long time scales.

[Vertical variability of Pinus sylvestris var. mongolica tree ring delta13C and its relationship with tree ring width in northern Daxing' an Mountains of Northeast China].

PubMed

Shang, Zhi-Yuan; Wang, Jian; Zhang, Wen; Li, Yan-Yan; Cui, Ming-Xing; Chen, Zhen-Ju; Zhao, Xing-Yun

2013-01-01

A measurement was made on the vertical direction tree ring stable carbon isotope ratio (delta13C) and tree ring width of Pinus sylvestris var. mongolica in northern Daxing' an Mountains of Northeast China, with the relationship between the vertical direction variations of the tree ring delta13C and tree ring width analyzed. In the whole ring of xylem, earlywood (EW) and bark endodermis, the delta13C all exhibited an increasing trend from the top to the base at first, with the maximum at the bottom of tree crown, and then, decreased rapidly to the minimum downward. The EW and late-wood (LW) had an increasing ratio of average tree ring width from the base to the top. The average annual sequence of the delta13C in vertical direction had an obvious reverse correspondence with the average annual sequence of tree ring width, and had a trend comparatively in line with the average annual sequence of the tree ring width ratio of EW to LW above tree crown. The variance analysis showed that there existed significant differences in the sequences of tree ring delta13C and ring width in vertical direction, and the magnitude of vertical delta13C variability was basically the same as that of the inter-annual delta13C variability. The year-to-year variation trend of the vertical delta13C sequence was approximately identical. For each sample, the delta13C sequence at the same heights was negatively correlated with the ring width sequence, but the statistical significance differed with tree height.

Size Matters a Lot: Drought-Affected Italian Oaks Are Smaller and Show Lower Growth Prior to Tree Death.

PubMed

Colangelo, Michele; Camarero, Jesús J; Borghetti, Marco; Gazol, Antonio; Gentilesca, Tiziana; Ripullone, Francesco

2017-01-01

Hydraulic theory suggests that tall trees are at greater risk of drought-triggered death caused by hydraulic failure than small trees. In addition the drop in growth, observed in several tree species prior to death, is often interpreted as an early-warning signal of impending death. We test these hypotheses by comparing size, growth, and wood-anatomy patterns of living and now-dead trees in two Italian oak forests showing recent mortality episodes. The mortality probability of trees is modeled as a function of recent growth and tree size. Drift-diffusion-jump (DDJ) metrics are used to detect early-warning signals. We found that the tallest trees of the anisohydric Italian oak better survived drought contrary to what was predicted by the theory. Dead trees were characterized by a lower height and radial-growth trend than living trees in both study sites. The growth reduction of now-dead trees started about 10 years prior to their death and after two severe spring droughts during the early 2000s. This critical transition in growth was detected by DDJ metrics in the most affected site. Dead trees were also more sensitive to drought stress in this site indicating different susceptibility to water shortage between trees. Dead trees did not form earlywood vessels with smaller lumen diameter than surviving trees but tended to form wider latewood vessels with a higher percentage of vessel area. Since living and dead trees showed similar competition we did not expect that moderate thinning and a reduction in tree density would increase the short-term survival probability of trees.

Hydraulic efficiency and safety of branch xylem increases with height in Sequoia sempervirens (D. Don) crowns.

PubMed

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.

Functional multi-locus QTL mapping of temporal trends in Scots pine wood traits.

PubMed

Li, Zitong; Hallingbäck, Henrik R; Abrahamsson, Sara; Fries, Anders; Gull, Bengt Andersson; Sillanpää, Mikko J; García-Gil, M Rosario

2014-10-09

Quantitative trait loci (QTL) mapping of wood properties in conifer species has focused on single time point measurements or on trait means based on heterogeneous wood samples (e.g., increment cores), thus ignoring systematic within-tree trends. In this study, functional QTL mapping was performed for a set of important wood properties in increment cores from a 17-yr-old Scots pine (Pinus sylvestris L.) full-sib family with the aim of detecting wood trait QTL for general intercepts (means) and for linear slopes by increasing cambial age. Two multi-locus functional QTL analysis approaches were proposed and their performances were compared on trait datasets comprising 2 to 9 time points, 91 to 455 individual tree measurements and genotype datasets of amplified length polymorphisms (AFLP), and single nucleotide polymorphism (SNP) markers. The first method was a multilevel LASSO analysis whereby trend parameter estimation and QTL mapping were conducted consecutively; the second method was our Bayesian linear mixed model whereby trends and underlying genetic effects were estimated simultaneously. We also compared several different hypothesis testing methods under either the LASSO or the Bayesian framework to perform QTL inference. In total, five and four significant QTL were observed for the intercepts and slopes, respectively, across wood traits such as earlywood percentage, wood density, radial fiberwidth, and spiral grain angle. Four of these QTL were represented by candidate gene SNPs, thus providing promising targets for future research in QTL mapping and molecular function. Bayesian and LASSO methods both detected similar sets of QTL given datasets that comprised large numbers of individuals. Copyright © 2014 Li et al.

Scots pine responses to elevated temperature and carbon dioxide concentration: growth and wood properties.

PubMed

Kilpeläinen, Antti; Peltola, Heli; Ryyppö, Aija; Kellomäki, Seppo

2005-01-01

Growth and wood properties of 20-year-old Scots pine (Pinus sylvestris L.) trees were studied for 6 years in 16 closed chambers providing a factorial combination of two temperature regimes (ambient and elevated) and two carbon dioxide concentrations ([CO2]) (ambient and twice ambient). The elevation of temperature corresponded to the predicted effect at the site of a doubling in atmospheric [CO2]. Annual height and radial growth and wood properties were analyzed during 1997-2002. Physical wood properties analyzed included early- and latewood widths and their proportions, intra-ring wood densities, early- and latewood density and mean fiber length. Chemical wood properties analyzed included concentrations of acetone-soluble extractives, lignin, cellulose and hemicellulose. There were no significant treatment effects on height growth during the 6-year study. Elevated [CO2] increased ring width by 66 and 47% at ambient and elevated temperatures, respectively. At ambient [CO2], elevated temperature increased ring width by 19%. Increased ring width in response to elevated [CO2] resulted from increases in both early- and latewood width; however, there was no effect of the treatments on early- and latewood proportions. Mean wood density, earlywood density and fiber length increased in response to elevated temperature. The chemical composition of wood was affected by elevated [CO2], which reduced the cellulose concentration, and by elevated temperature, which reduced the concentration of acetone-soluble extractives. Thus, over the 6-year period, radial growth was significantly increased by elevated [CO2], and some wood properties were significantly affected by elevated temperature or elevated [CO2], or both, indicating that climate change may affect the material properties of wood.

Latitudinal Gradients in the Stable Carbon and Oxygen Isotopes of Tree-Ring Cellulose Reveal Differential Climate Influences of the North American Monsoon

NASA Astrophysics Data System (ADS)

Szejner, P.; Wright, W. E.; Babst, F.; Belmecheri, S.; Trouet, V.; Ehleringer, J. R.; Leavitt, S. W.; Monson, R. K.

2015-12-01

Summer rainfall plays an important role sustaining different types of ecosystems in the Southwestern US. The arrival of the monsoon breaks the early summer hyper-arid period in the region providing unique seasonal conditions for these ecosystems to thrive. It is unknown to what extent monsoon rainfall is used by Ponderosa pine forests, which occupy many mountain ecosystems in the Western US. While these forests clearly rely on winter snowpack to drive much of their annual net primary productivity, the extent to which they supplement winter moisture, with summer monsoon moisture needs to be clarified. It is likely that there are north-south gradients in the degree to which forests rely on monsoon moisture, as the summer monsoon system tends to become diminished as it moves progressively northward. We addressed these gaps in our knowledge about the monsoon by studying stable Carbon and Oxygen isotopes in earlywood and latewood α-cellulose from cores taken from trees in eleven sites along a latitudinal gradient extending from Southern Arizona and New Mexico toward Utah. Here we show evidence that Ponderosa pine trees from most of these sites use monsoon water to support growth during the late summer, and the fractional use of monsoon precipitation is strongest in the southernmost sites. This study provides new physiological evidence on the influence of the North American monsoon and winter precipitation on tree growth in montane ecosystems of the Western US. Using these results, we predict differences in the susceptibility of southern and northern montane forests to future climate change. ACKNOWLEDGMENTS: This work was funded by an NSF Macrosystems Grant #1065790

Xylogenesis: Coniferous Trees of Temperate Forests Are Listening to the Climate Tale during the Growing Season But Only Remember the Last Words!

PubMed

Cuny, Henri E; Rathgeber, Cyrille B K

2016-05-01

The complex inner mechanisms that create typical conifer tree-ring structure (i.e. the transition from large, thin-walled earlywood cells to narrow, thick-walled latewood cells) were recently unraveled. However, what physiological or environmental factors drive xylogenesis key processes remain unclear. Here, we aim to quantify the influence of seasonal variations in climatic factors on the spectacular changes in the kinetics of wood cell differentiation and in the resulting tree-ring structure. Wood formation was monitored in three sites over 3 years for three coniferous species (Norway spruce [Picea abies], Scots pine [Pinus sylvestris], and silver fir [Abies alba]). Cell differentiation rates and durations were calculated and related to tracheid final dimensions and corresponding climatic conditions. On the one hand, we found that the kinetics of cell enlargement and the final size of the tracheids were not explained by the seasonal changes in climatic factors. On the other hand, decreasing temperatures strongly constrained cell wall deposition rates during latewood formation. However, the influence of temperature was permanently written into tree-ring structure only for the very last latewood cells, when the collapse of the rate of wall deposition was no longer counterbalanced by the increase of its duration. Our results show that the formation of the typical conifer tree-ring structure, in normal climatic conditions, is only marginally driven by climate, suggesting strong developmental control of xylogenesis. The late breakage of the compensatory mechanism at work in the wall deposition process appears as a clue to understand the capacity of the maximum latewood density to record past temperature conditions. © 2016 American Society of Plant Biologists. All Rights Reserved.

Widening of xylem conduits in a conifer tree depends on the longer time of cell expansion downwards along the stem

PubMed Central

Anfodillo, Tommaso; Deslauriers, Annie; Menardi, Roberto; Tedoldi, Laura; Petit, Giai; Rossi, Sergio

2012-01-01

The diameter of vascular conduits increases towards the stem base. It has been suggested that this profile is an efficient anatomical feature for reducing the hydraulic resistance when trees grow taller. However, the mechanism that controls the cell diameter along the plant is not fully understood. The timing of cell differentiation along the stem was investigated. Cambial activity and cell differentiation were investigated in a Picea abies tree (11.5 m in height) collecting microsamples at nine different heights (from 1 to 9 m) along the stem with a 4 d time interval. Wood sections (8–12 μm thick) were stained and observed under a light microscope with polarized light to differentiate the developing xylem cells. Cell wall lignification was detected using cresyl violet acetate. The first enlarging cells appeared almost simultaneously along the tree axis indicating that cambium activation is not height-dependent. A significant increase in the duration of the cell expansion phase was observed towards the tree base: at 9 m from the ground, xylem cells expanded for 7 d, at 6 m for 14 d, and at 3 m for 19 d. The duration of the expansion phase is positively correlated with the lumen area of the tracheids (r2=0.68, P < 0.01) at the same height. By contrast, thickness of the cell wall of the earlywood did not show any trend with height. The lumen area of the conduits down the stem appeared linearly dependent on time during which differentiating cells remained in the expansion phase. However, the inductive signal of such long-distance patterned differentiation remains to be identified. PMID:22016427

Widening of xylem conduits in a conifer tree depends on the longer time of cell expansion downwards along the stem.

PubMed

Anfodillo, Tommaso; Deslauriers, Annie; Menardi, Roberto; Tedoldi, Laura; Petit, Giai; Rossi, Sergio

2012-01-01

The diameter of vascular conduits increases towards the stem base. It has been suggested that this profile is an efficient anatomical feature for reducing the hydraulic resistance when trees grow taller. However, the mechanism that controls the cell diameter along the plant is not fully understood. The timing of cell differentiation along the stem was investigated. Cambial activity and cell differentiation were investigated in a Picea abies tree (11.5 m in height) collecting microsamples at nine different heights (from 1 to 9 m) along the stem with a 4 d time interval. Wood sections (8-12 μm thick) were stained and observed under a light microscope with polarized light to differentiate the developing xylem cells. Cell wall lignification was detected using cresyl violet acetate. The first enlarging cells appeared almost simultaneously along the tree axis indicating that cambium activation is not height-dependent. A significant increase in the duration of the cell expansion phase was observed towards the tree base: at 9 m from the ground, xylem cells expanded for 7 d, at 6 m for 14 d, and at 3 m for 19 d. The duration of the expansion phase is positively correlated with the lumen area of the tracheids (r(2)=0.68, P < 0.01) at the same height. By contrast, thickness of the cell wall of the earlywood did not show any trend with height. The lumen area of the conduits down the stem appeared linearly dependent on time during which differentiating cells remained in the expansion phase. However, the inductive signal of such long-distance patterned differentiation remains to be identified.

Non-stationary influence of El Niño-Southern Oscillation and winter temperature on oak latewood growth in NW Iberian Peninsula.

PubMed

Rozas, Vicente; García-González, Ignacio

2012-09-01

The properties of El Niño-Southern Oscillation (ENSO), such as period, amplitude, and teleconnection strength to extratropical regions, have changed since the mid-1970s. ENSO affects the regional climatic regime in SW Europe, thus tree performance in the Iberian Peninsula could be affected by recent ENSO dynamics. We established four Quercus robur chronologies of earlywood and latewood widths in the NW Iberian Peninsula. The relationship between tree growth and the Southern Oscillation Index (SOI), the atmospheric expression of ENSO, showed that only latewood growth was correlated negatively with the SOI of the previous summer-autumn-winter. This relationship was non-stationary, with significant correlations only during the period 1952-1980; and also non-linear, with enhanced latewood growth only in La Niña years, i.e. years with a negative SOI index for the previous autumn. Non-linear relationship between latewood and SOI indicates an asymmetric influence of ENSO on tree performance, biassed towards negative SOI phases. During La Niña years, climate in the study area was warmer and wetter than during positive years, but only for 1952-1980. Winter temperatures became the most limiting factor for latewood growth since 1980, when mean regional temperatures increased by 1°C in comparison to previous periods. As a result, higher winter respiration rates, and the extension of the growing season, would probably cause an additional consumption of stored carbohydrates. The influence of ENSO and winter temperatures proved to be of great importance for tree growth, even at lower altitudes and under mild Atlantic climate in the NW Iberian Peninsula.

Assessment of tree response to drought: validation of a methodology to identify and test proxies for monitoring past environmental changes in trees.

PubMed

Tene, A; Tobin, B; Dyckmans, J; Ray, D; Black, K; Nieuwenhuis, M

2011-03-01

A thinning experiment stand at Avoca, Ballinvalley, on the east coast of the Republic of Ireland was used to test a developed methodology aimed at monitoring drought stress, based on the analysis of growth rings obtained by coring. The stand incorporated six plots representing three thinning regimes (light, moderate and heavy) and was planted in the spring of 1943 on a brown earth soil. Radial growth (early- and latewood) was measured for the purpose of this study. A multidisciplinary approach was used to assess historic tree response to climate: specifically, the application of statistical tools such as principal component and canonical correlation analysis to dendrochronology, stable isotopes, ring density proxy, blue reflectance and forest biometrics. Results showed that radial growth was a good proxy for monitoring changes to moisture deficit, while maximum density and blue reflectance were appropriate for assessing changes in accumulated temperature for the growing season. Rainfall also influenced radial growth changes but not significantly, and was a major factor in stable carbon and oxygen discrimination, mostly in the latewood formation phase. Stable oxygen isotope analysis was more accurate than radial growth analysis in drought detection, as it helped detect drought signals in both early- and latewood while radial growth analysis only detected the drought signal in earlywood. Many studies have shown that tree rings provide vital information for marking past climatic events. This work provides a methodology to better identify and understand how commonly measured tree proxies relate to environmental parameters, and can best be used to characterize and pinpoint drought events (variously described using parameters such as like moisture deficit, accumulated temperature, rainfall and potential evaporation).

Transcriptomic analysis of wound xylem formation in Pinus canariensis.

PubMed

Chano, V; Collada, C; Soto, A

2017-12-04

Woody plants, especially trees, usually must face several injuries caused by different agents during their lives. Healing of injuries in stem and branches, affecting the vascular cambium and xylem can take several years. In conifers, healing takes place mainly from the remaining vascular cambium in the margin of the wound. The woundwood formed in conifers during healing usually presents malformed and disordered tracheids as well as abundant traumatic resin ducts. These characteristics affect its functionality as water conductor and its technological properties. In this work we analyze for the first time the transcriptomic basis of the formation of traumatic wood in conifers, and reveal some differences with normal early- and late-wood. Microarray analysis of the differentiating traumatic wood, confirmed by quantitative RT-PCR, has revealed alterations in the transcription profile of up to 1408 genes during the first period of healing. We have grouped these genes in twelve clusters, according to their transcription profiles, and have distinguished accordingly two main phases during this first healing. Wounding induces a complete rearrangement of the transcriptional program in the cambial zone close to the injuries. At the first instance, radial growth is stopped, and a complete set of defensive genes, mostly related to biotic stress, are induced. Later on, cambial activity is restored in the lateral borders of the wound, even at a high rate. During this second stage certain genes related to early-wood formation, including genes involved in cell wall formation and transcription factors, are significantly overexpressed, while certain late-wood related genes are repressed. Additionally, significant alterations in the transcription profile of abundant non annotated genes are reported.

Climate Response of Tree Radial Growth at Different Timescales in the Qinling Mountains.

PubMed

Sun, Changfeng; Liu, Yu

2016-01-01

The analysis of the tree radial growth response to climate is crucial for dendroclimatological research. However, the response relationships between tree-ring indices and climatic factors at different timescales are not yet clear. In this study, the tree-ring width of Huashan pine (Pinus armandii) from Huashan in the Qinling Mountains, north-central China, was used to explore the response differences of tree growth to climatic factors at daily, pentad (5 days), dekad (10 days) and monthly timescales. Correlation function and linear regression analysis were applied in this paper. The tree-ring width showed a more sensitive response to daily and pentad climatic factors. With the timescale decreasing, the absolute value of the maximum correlation coefficient between the tree-ring data and precipitation increases as well as temperature (mean, minimum and maximum temperature). Compared to the other three timescales, pentad was more suitable for analysing the response of tree growth to climate. Relative to the monthly climate data, the association between the tree-ring data and the pentad climate data was more remarkable and accurate, and the reconstruction function based on the pentad climate was also more reliable and stable. We found that the major climatic factor limiting Huashan pine growth was the precipitation of pentads 20-35 (from April 6 to June 24) rather than the well-known April-June precipitation. The pentad was also proved to be a better timescale for analysing the climate and tree growth in the western and eastern Qinling Mountains. The formation of the earlywood density of Chinese pine (Pinus tabulaeformis) from Shimenshan in western Qinling was mainly affected by the maximum temperature of pentads 28-32 (from May 16 to June 9). The maximum temperature of pentads 28-33 (from May 16 to June 14) was the major factor affecting the ring width of Chinese pine from Shirenshan in eastern Qinling.

Drought-induced adaptation of the xylem in Scots pine and pubescent oak.

PubMed

Eilmann, Britta; Zweifel, Roman; Buchmann, Nina; Fonti, Patrick; Rigling, Andreas

2009-08-01

Drought impairs tree growth in the inner-Alpine valleys of Central Europe. We investigated species-specific responses to contrasting water supply, with Scots pine (Pinus sylvestris L.), threatened by drought-induced mortality, and pubescent oak (Quercus pubescens Willd.), showing no connection between drought events and mortality. The two co-occurring tree species were compared, growing either along an open water channel or at a site with naturally dry conditions. In addition, the growth response of Scots pine to a draining of a water channel was studied. We analysed the radial increment for the last 100 years and wood anatomical parameters for the last 45 years. Drought reduced the conduit area of pubescent oak, but increased the radial lumen diameter of the conduits in Scots pine. Both species decreased their radial increment under drought. In Scots pine, radial increment was generally more dependent on water availability than that in pubescent oak. Irrigated trees responded less negatively to high temperature as seen in the increase in the conduit area in pubescent oak and the removal of the limitation of cell division by high temperatures. After irrigation stopped, tree-ring width for Scots pine decreased within 1-year delay, whereas lumen diameter and cell-wall thickness responded with a 4-year delay. Scots pine seemed to optimize the carbon-per-conduit-costs under drought by increasing conduits diameter while decreasing cell numbers. This strategy might lead to a complete loss of tree rings under severe drought and thus to an impairment of water transport. In contrast, in pubescent oak tree-ring width is less affected by summer drought because parts of the earlywood are built in early spring. Thus, pubescent oak might have gradual advantages over pine in today's climate of the inner-Alpine valley.

Drought response of five conifer species under contrasting water availability suggests high vulnerability of Norway spruce and European larch.

PubMed

Lévesque, Mathieu; Saurer, Matthias; Siegwolf, Rolf; Eilmann, Britta; Brang, Peter; Bugmann, Harald; Rigling, Andreas

2013-10-01

The ability of tree species to cope with anticipated decrease in water availability is still poorly understood. We evaluated the potential of Norway spruce, Scots pine, European larch, black pine, and Douglas-fir to withstand drought in a drier future climate by analyzing their past growth and physiological responses at a xeric and a mesic site in Central Europe using dendroecological methods. Earlywood, latewood, and total ring width, as well as the δ(13) C and δ(18) O in early- and latewood were measured and statistically related to a multiscalar soil water deficit index from 1961 to 2009. At the xeric site, δ(13) C values of all species were strongly linked to water deficits that lasted longer than 11 months, indicating a long-term cumulative effect on the carbon pool. Trees at the xeric site were particularly sensitive to soil water recharge in the preceding autumn and early spring. The native species European larch and Norway spruce, growing close to their dry distribution limit at the xeric site, were found to be the most vulnerable species to soil water deficits. At the mesic site, summer water availability was critical for all species, whereas water availability prior to the growing season was less important. Trees at the mesic were more vulnerable to water deficits of shorter duration than the xeric site. We conclude that if summers become drier, trees growing on mesic sites will undergo significant growth reductions, whereas at their dry distribution limit in the Alps, tree growth of the highly sensitive spruce and larch may collapse, likely inducing dieback and compromising the provision of ecosystem services. However, the magnitude of these changes will be mediated strongly by soil water recharge in winter and thus water availability at the beginning of the growing season. © 2013 John Wiley & Sons Ltd.

Towards a better understanding of long-term wood-chemistry variations in old-growth forests: A case study on ancient Pinus uncinata trees from the Pyrenees.

PubMed

Hevia, Andrea; Sánchez-Salguero, Raúl; Camarero, J Julio; Buras, Allan; Sangüesa-Barreda, Gabriel; Galván, J Diego; Gutiérrez, Emilia

2018-06-01

Dendrochemical studies in old forests are still underdeveloped. Old trees growing in remote high-elevation areas far from direct human influence constitute a promising biological proxy for the long-term reconstructions of environmental changes using tree-rings. Furthermore, centennial-long chronologies of multi-elemental chemistry at inter- and intra-annual resolution are scarce. Here, we use a novel non-destructive method by applying Micro X-ray fluorescence (μXRF) to wood samples of old Pinus uncinata trees from two Pyrenean high-elevation forests growing on acidic and basic soils. To disentangle ontogenetic (changes in tree age and diameter) from environmental influences (e.g., climate warming) we compared element patterns in sapwood (SW) and heartwood (HW) during the pre-industrial (1700-1849) and industrial (1850-2008) periods. We quantified tree-ring growth, wood density and relative element concentrations at annual (TRW, tree-ring) to seasonal resolution (EW, earlywood; LW, latewood) and related them to climate variables (temperature and precipitation) and volcanic eruptions in the 18th and 19th centuries. We detected differences for most studied elements between SW and HW along the stem and also between EW and LW within rings. Long-term positive and negative trends were observed for Ca and K, respectively. Cl, P and S showed positive trends during the industrial period. However, differences between sites were also notable. Higher values of Mg, Al, Si and the Ca/Mn ratio were observed at the site with acidic soil. Growing-season temperatures were positively related to growth, maximum wood density and to the concentration of most elements. Peaks in S, Fe, Cl, Zn and Ca were linked to major volcanic eruptions (e.g., Tambora in 1815). Our results reveal the potential of long-term wood-chemistry studies based on the μXRF non-destructive technique to reconstruct environmental changes. Copyright © 2017 Elsevier B.V. All rights reserved.

Powdery Mildew Decreases the Radial Growth of Oak Trees with Cumulative and Delayed Effects over Years

PubMed Central

Bert, Didier; Lasnier, Jean-Baptiste; Capdevielle, Xavier; Dugravot, Aline; Desprez-Loustau, Marie-Laure

2016-01-01

Quercus robur and Q. petraea are major European forest tree species. They have been affected by powdery mildew caused by Erysiphe alphitoides for more than a century. This fungus is a biotrophic foliar pathogen that diverts photosynthetate from the plant for its own nutrition. We used a dendrochronological approach to investigate the effects of different levels of infection severity on the radial growth of young oak trees. Oak infection was monitored at individual tree level, at two sites in southwestern France, over a five-year period (2001–2005). Mean infection severity was almost 75% (infected leaf area) at the end of the 2001 growing season, at both sites, but only about 40% in 2002, and 8%, 5% and 2% in 2003, 2004 and 2005, respectively. Infection levels varied considerably between trees and were positively related between 2001 and 2002. Increment cores were taken from each tree to assess annual ring widths and increases in basal area. Annual radial growth was standardised to take the effect of tree size into account. Annual standardised radial growth was significantly and negatively correlated with infection severity in the same year, for both 2001 and 2002, and at both sites. The decrease in growth reached 70–90% for highly infected trees. The earlywood width was poorly correlated with infection severity, but the proportion of latewood in tree rings was lower in highly infected trees (60%) than in less heavily infected trees (85%). Infection in 2001 and 2002 was found to have a cumulative effect on radial growth in these years, together with a delayed effect detectable in 2003. Thus, even non-lethal pathogens like powdery mildew can have a significant impact on tree functioning. This impact should be taken into account in growth and yield models, to improve predictions of forest net primary production. PMID:27177029

[The changes in contents and composition of phenolic acids during cell xylem growth in scots pine].

PubMed

Antonova, G F; Zheliznichenko, T V; Stasova, V V

2011-01-01

The contents and composition of alcohol soluble phenolic acids were studied during cell xylem growth in the course of wood annual increment formation in the stems of Scots pine. The cells of cambium zone, of two stages of expansion growth and the outset of secondary thickening zone (before lignification) were successively gathered from the stem segments of 25-old pine trees in the period of earlywood xylem formation with constant anatomical and histochemical control. The contents of free and bound forms of phenolic acids, isolated by 80% ethanol from tissues, as well as of their ethers and esters were calculated both per dry weight and per cell. The content and relation of the fractions and the composition of phenolic acid have been found to change significantly from cambium zone to the outset of tracheid secondary thickening. The character of the variations depends on a calculation method. According to the calculation per cell the amount of free and bound phenolic acids and in their composition of esters and especially ethers increased at the first step of expansion growth zone, decreased at the second one and rose again in the outset of secondary wall deposition. In dependence on the stage of cell development the pool of bound phenolic acids exceeded of free acid pool in 2-5 times. Sinapic and ferulic acids dominated in the composition of free hydroxycinnamic acids. The content and composition of hydroxycinnamic acids in ethers and esters depended on cell development phase. In cambium p-coumaric and sinapic acids were principal aglycons in ethers, at other stages these were sinapic and caffeic acids. The esters in cambium zone included essentially p-coumaric acid and at the other stages - sinapic and ferulic acids. At the first phase of growth benzoic acid was connected principally by ester bonds. The pool of these esters decreased from the first phase of growth to the outset of cell wall thickening and in proportion to this the level of free benzoic acid rose.

Phloem Girdling of Norway Spruce Alters Quantity and Quality of Wood Formation in Roots Particularly Under Drought

PubMed Central

Rainer-Lethaus, Gina; Oberhuber, Walter

2018-01-01

Carbon (C) availability plays an essential role in tree growth and wood formation. We evaluated the hypothesis that a decrease in C availability (i) triggers mobilization of C reserves in the coarse roots of Picea abies to maintain growth and (ii) causes modification of wood structure notably under drought. The 6-year-old saplings were subjected to two levels of soil moisture (watered versus drought conditions) and root C status was manipulated by physically blocking phloem transport in the stem at three girdling dates (GDs). Stem girdling was done before the onset of bud break [day of the year (doy) 77], during vigorous aboveground shoot and radial stem growth (GD doy 138), and after cessation of shoot growth (GD doy 190). The effect of blockage of C transport on root growth, root phenology, and wood anatomical traits [cell lumen diameter (CLD) and cell wall thickness (CWT)] in earlywood (EW) and latewood (LW) was determined. To evaluate changes in belowground C status caused by girdling, non-structural carbohydrates (soluble sugars and starch) in coarse roots were determined at the time of girdling and after the growing season. Although fine root mass significantly decreased in response to blockage of phloem C transport, the phenology of root elongation growth was not affected. Surprisingly, radial root growth and CLD of EW tracheids in coarse roots were strikingly increased in drought-stressed trees, when girdling occurred before bud break or during aboveground stem growth. In watered trees, the growth response to girdling was less distinct, but the CWT of EW significantly increased. Starch reserves in the roots of girdled trees significantly decreased in both soil moisture treatments and at all GDs. We conclude that (i) radial growth and wood development in coarse roots of P. abies saplings are not only dependent on current photosynthates, and (ii) blockage of phloem transport induces physiological changes that outweigh drought effects imposed on root cambial

Cambial injury in lodgepole pine (Pinus contorta): mountain pine beetle vs fire.

PubMed

Arbellay, Estelle; Daniels, Lori D; Mansfield, Shawn D; Chang, Alice S

2017-12-01

Both mountain pine beetle (MPB) Dendroctonus ponderosae Hopkins and fire leave scars with similar appearance on lodgepole pine Pinus contorta Dougl. ex Loud. var. latifolia Engelm. that have never been compared microscopically, despite the pressing need to determine the respective effects of MPB and fire injury on tree physiology. We analysed changes in wood formation in naturally caused scars on lodgepole pine, and tested the hypotheses that (i) MPB and fire injury elicit distinct anomalies in lodgepole pine wood and (ii) anomalies differ in magnitude and/or duration between MPB and fire. Mountain pine beetle and fire injury reduced radial growth in the first year post-injury. Otherwise, radial growth and wood density increased over more than 10 years in both MPB and fire scars. We found that the general increase in radial growth was of greater magnitude (up to 27%) and of longer duration (up to 5 years) in fire scars compared with MPB scars, as shown in earlywood width. We also observed that the increase in latewood density was of greater magnitude (by 12%) in MPB scars, but of longer duration (by 4 years) in fire scars. Crystallinity decreased following MPB and fire injury, while microfibril angle increased. These changes in fibre traits were of longer duration (up to 4 years) in MPB scars compared with fire scars, as shown in microfibril angle. We found no significant changes in carbon and nitrogen concentrations. In conclusion, we stress that reduced competition and resistance to cavitation play an important role alongside cambial injury in influencing the type and severity of changes. In addition, more research is needed to validate the thresholds introduced in this study. Our findings serve as a foundation for new protocols to distinguish between bark beetle and fire disturbance, which is essential for improving our knowledge of historical bark beetle and fire regimes, and their interactions. © The Author 2017. Published by Oxford University Press. All

Comparing the intra-annual wood formation of three European species (Fagus sylvatica, Quercus petraea and Pinus sylvestris) as related to leaf phenology and non-structural carbohydrate dynamics.

PubMed

Michelot, Alice; Simard, Sonia; Rathgeber, Cyrille; Dufrêne, Eric; Damesin, Claire

2012-08-01

Monitoring cambial phenology and intra-annual growth dynamics is a useful approach for characterizing the tree growth response to climate change. However, there have been few reports concerning intra-annual wood formation in lowland temperate forests with high time resolution, especially for the comparison between deciduous and coniferous species. The main objective of this study was to determine how the timing, duration and rate of radial growth change between species as related to leaf phenology and the dynamics of non-structural carbohydrates (NSC) under the same climatic conditions. We studied two deciduous species, Fagus sylvatica L. and Quercus petraea (Matt.) Liebl., and an evergreen conifer, Pinus sylvestris L. During the 2009 growing season, we weekly monitored (i) the stem radial increment using dendrometers, (ii) the xylem growth using microcoring and (iii) the leaf phenology from direct observations of the tree crowns. The NSC content was also measured in the eight last rings of the stem cores in April, June and August 2009. The leaf phenology, NSC storage and intra-annual growth were clearly different between species, highlighting their contrasting carbon allocation. Beech growth began just after budburst, with a maximal growth rate when the leaves were mature and variations in the NSC content were low. Thus, beech radial growth seemed highly dependent on leaf photosynthesis. For oak, earlywood quickly developed before budburst, which probably led to the starch decrease quantified in the stem from April to June. For pine, growth began before the needles unfolding and the lack of NSC decrease during the growing season suggested that the substrates for radial growth were new assimilates of the needles from the previous year. Only for oak, the pattern determined from the intra-annual growth measured using microcoring differed from the pattern determined from dendrometer data. For all species, the ring width was significantly influenced by growth duration

Coexisting oak species, including rear-edge populations, buffer climate stress through xylem adjustments.

PubMed

Granda, E; Alla, A Q; Laskurain, N A; Loidi, J; Sánchez-Lorenzo, A; Camarero, J J

2018-02-01

The ability of trees to cope with climate change is a pivotal feature of forest ecosystems, especially for rear-edge populations facing warm and dry conditions. To evaluate current and future forests threats, a multi-proxy focus on the growth, anatomical and physiological responses to climate change is needed. We examined the long-term xylem adjustments to climate variability of the temperate Quercus robur L. at its rear edge and the sub-Mediterranean Quercus pyrenaica Willd. Both species coexist at a mesic (ME, humid and warmer) and a xeric (XE, dry and cooler) site in northern Spain, the latter experiencing increasing temperatures in recent decades. We compared xylem traits at each site and assessed their trends, relationships and responses to climate (1960-2008). Traits included basal area increment, earlywood vessel hydraulic diameter, density and theoretical-specific hydraulic conductivity together with latewood oxygen (δ18O) stable isotopes and δ13C-derived water-use efficiency (iWUE). Quercus robur showed the highest growth at ME, likely through enhanced cambial activity. Quercus pyrenaica had higher iWUE at XE compared with ME, but limited plasticity of anatomical xylem traits was found for the two oak species. Similar physiological performance was found for both species. The iWUE augmented in recent years especially at XE, likely explained by stomatal closure given the increasing δ18O signal in response to drier and sunnier growing seasons. Overall, traits were more correlated at XE than at ME. The iWUE improvements were linked to higher growth up to a threshold (~85 μmol mol-1) after which reduced growth was found at XE. Our results are consistent with Q. pyrenaica and Q. robur coexisting at the central and dry edge of the climatic species distribution, respectively, showing similar responses to buffer warmer conditions. In fact, the observed adjustments found for Q. robur point towards growth stability of similar rear-edge oak populations under

Seasonal and clonal variation in cellulose microfibril orientation during cell wall formation of tracheids in Cryptomeria japonica.

PubMed

Jyske, Tuula; Fujiwara, Takeshi; Kuroda, Katsushi; Iki, Taiichi; Zhang, Chunhua; Jyske, Tuomas K; Abe, Hisashi

2014-08-01

To investigate the biological mechanism by which trees control the changes in microfibril (MF) orientation among secondary cell wall layers of conifer tracheids, we studied seasonal variation in the orientation of newly deposited MFs during tracheid cell wall development in Japanese cedar (Cryptomeria japonica D. Don) trees growing in Central Japan (36°36'N, 140°39'E). Sample blocks were repeatedly collected from four 16-year-old clones of different origins during the growing season of 2010 to investigate the hypotheses that changes in cellulose MF orientation between wall layers exhibited seasonal and clonal differences. The progressive change in the orientation of newly deposited MFs on the primary and secondary cell wall layers of tracheids was detected by field-emission-scanning electron microscopy. Tracheid production and differentiation was studied by light microscopy. We observed a decreasing trend in the orientation of deposited MFs from earlywood to latewood in the S2 and S1 layers, where MFs appeared in a Z-helix. In contrast, no seasonal pattern in the orientation of the MFs in the S-helix was observed. Minor clonal variation was observed in the phenology of tracheid production and differentiation. We concluded that a seasonal decreasing trend in the orientation of the MFs in the Z-helix in S1 and S2 was present, whereas the MFs in other layers exhibited minor random variations. Thus, the orientation of the MFs in S2 was affected by seasonal factors, whereas the MFs in other layers were more intrinsically controlled. The within-ring variations in the MF orientation and thus the resulting average MF angle might also be related to genotypic differences in the tracheid production and differentiation rate. However, our results do not exclude other intrinsic and environmental regulations in the change in MF orientation, which remains a topic for future studies. © The Author 2014. Published by Oxford University Press. All rights reserved. For Permissions

Xylem and Leaf Functional Adjustments to Drought in Pinus sylvestris and Quercus pyrenaica at Their Elevational Boundary

PubMed Central

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

Xylem and Leaf Functional Adjustments to Drought in Pinus sylvestris and Quercus pyrenaica at Their Elevational Boundary.

PubMed

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

Seasonal Climate Signals in Multiple Tree-Ring Parameters: A Pilot Study of Pinus ponderosa in the Columbia River Basin

NASA Astrophysics Data System (ADS)

Dannenberg, M.; Wise, E. K.; Keung, J. H.

2014-12-01

Proxy-based reconstructions of past climate have played an integral role in assessments of historical climate change, and tree-ring widths (TRW) have a long history of use in this paleoclimate research due to their annual resolution, widespread availability, and sensitivity of growth processes to variation in temperature and water availability. Increasingly, studies have shown that additional tree-ring metrics—including earlywood and latewood widths (EW and LW, respectively), maximum latewood density, and the intensity of reflected blue light from latewood (BI)—can provide additional information on seasonal climatic variability that is not present in TRW alone due to different processes that affect growth in different parts of the growing season. Studies of these additional tree-ring metrics highlight their utility in climate reconstructions, but to date they have mostly been limited to a few tree species and regions. Here, we extend the range of previous studies on alternative tree-ring metrics by evaluating the seasonal climate signals in TRW, EW, LW, and BI of Pinus ponderosa at six semiarid sites surrounding the Columbia River basin in the U.S. Pacific Northwest (PNW). Cores from each site were cross-dated and EW, LW, and TRW were measured using standard dendrochronological procedures. BI was obtained using a high-resolution flatbed scanner and CooRecorder software. To evaluate the unique climate processes and seasonalities contributing to different dendrochronological metrics, monthly temperature and precipitation from each site were obtained from the PRISM climate model and were correlated with each of the tree-ring metrics using the MATLAB program SEASCORR. We also evaluate the potential of using multiple tree-ring metrics (rather than a single proxy) in reconstructions of precipitation in the PNW. Initial results suggest that 1) tree growth at each site is water-limited but with substantial differences among the sites in the strength and seasonality of

Impact of drought on the temporal dynamics of wood formation in Pinus sylvestris

PubMed Central

GRUBER, ANDREAS; STROBL, STEFAN; VEIT, BARBARA; OBERHUBER, WALTER

2011-01-01

Summary We determined the temporal dynamics of cambial activity and xylem cell differentiation of Scots pine (Pinus sylvestris L.) within a dry inner Alpine valley (750 m asl, Tyrol, Austria), where radial growth is strongly limited by drought in spring. Repeated micro-sampling of the developing tree ring of mature trees was carried out during 2 contrasting years at two study plots that differ in soil water availability (xeric and dry-mesic site). In 2007, when air temperature at the beginning of the growing season in April exceeded the long-term mean by 6.4 °C, cambial cell division started in early April at both study plots. A delayed onset of cambial activity of c. 2 wk was found in 2008, when average climate conditions prevailed in spring, indicating that resumption of cambial cell division after winter dormancy is temperature-controlled. Cambial cell division consistently ended about the end of June/early July in both study years. Radial enlargement of tracheids started almost 3 wk earlier in 2007 compared with 2008 at both study plots. At the xeric site, the maximum rate of tracheid production in 2007 and 2008 was reached in early and mid-May, respectively, and c. 2 wk later, at the dry-mesic site. Since in both study years, more favorable growing conditions (i.e., an increase in soil water content) were recorded during summer, we suggest a strong sink competition for carbohydrates to mycorrhizal root and shoot growth. Wood formation stopped c. 4 wk earlier at the xeric compared with the dry-mesic site in both years, indicating a strong influence of drought stress on cell differentiation. This is supported by radial widths of earlywood cells, which were found to be significantly narrower at the xeric than at the dry-mesic site (P < 0.05). Repeated cellular analyses during the two growing seasons revealed that, although spatial variability in the dynamics and duration of cell differentiation processes in Pinus sylvestris exposed to drought is strongly

Impact of drought on the temporal dynamics of wood formation in Pinus sylvestris.

PubMed

Gruber, Andreas; Strobl, Stefan; Veit, Barbara; Oberhuber, Walter

2010-04-01

We determined the temporal dynamics of cambial activity and xylem cell differentiation of Scots pine (Pinus sylvestris L.) within a dry inner Alpine valley (750 m a.s.l., Tyrol, Austria), where radial growth is strongly limited by drought in spring. Repeated micro-sampling of the developing tree ring of mature trees was carried out during two contrasting years at two study plots that differ in soil water availability (xeric and dry-mesic sites). In 2007, when air temperature at the beginning of the growing season in April exceeded the long-term mean by 6.4 degrees C, cambial cell division started in early April at both study plots. A delayed onset of cambial activity of c. 2 weeks was found in 2008, when average climate conditions prevailed in spring, indicating that resumption of cambial cell division after winter dormancy is temperature controlled. Cambial cell division consistently ended about the end of June/early July in both study years. Radial enlargement of tracheids started almost 3 weeks earlier in 2007 compared with 2008 at both study plots. At the xeric site, the maximum rate of tracheid production in 2007 and 2008 was reached in early and mid-May, respectively, and c. 2 weeks later at the dry-mesic site. Since in both study years more favorable growing conditions (i.e., an increase in soil water content) were recorded during summer, we suggest a strong sink competition for carbohydrates to mycorrhizal root and shoot growth. Wood formation stopped c. 4 weeks earlier at the xeric compared with the dry-mesic site in both years, indicating a strong influence of drought stress on cell differentiation. This is supported by radial widths of earlywood cells, which were found to be significantly narrower at the xeric than at the dry-mesic site (P < 0.05). Repeated cellular analyses during the two growing seasons revealed that, although spatial variability in the dynamics and duration of cell differentiation processes in P. sylvestris exposed to drought is

Assessing Conifer Ray Parenchyma for Ecological Studies: Pitfalls and Guidelines.

PubMed

von Arx, Georg; Arzac, Alberto; Olano, José M; Fonti, Patrick

2015-01-01

Ray parenchyma is an essential tissue for tree functioning and survival. This living tissue plays a major role for storage and transport of water, nutrients, and non-structural carbohydrates (NSC), thus regulating xylem hydraulics and growth. However, despite the importance of rays for tree carbon and water relations, methodological challenges hamper knowledge about ray intra- and inter-tree variability and its ecological meaning. In this study we provide a methodological toolbox for soundly quantifying spatial and temporal variability of different ray features. Anatomical ray features were surveyed in different cutting planes (cross-sectional, tangential, and radial) using quantitative image analysis on stem-wood micro-sections sampled from 41 mature Scots pines (Pinus sylvestris). The percentage of ray surface (PERPAR), a proxy for ray volume, was compared among cutting planes and between early- and latewood to assess measurement-induced variability. Different tangential ray metrics were correlated to assess their similarities. The accuracy of cross-sectional and tangential measurements for PERPAR estimates as a function of number of samples and the measured wood surface was assessed using bootstrapping statistical technique. Tangential sections offered the best 3D insight of ray integration into the xylem and provided the most accurate estimates of PERPAR, with 10 samples of 4 mm(2) showing an estimate within ±6.0% of the true mean PERPAR (relative 95% confidence interval, CI95), and 20 samples of 4 mm(2) showing a CI95 of ±4.3%. Cross-sections were most efficient for establishment of time series, and facilitated comparisons with other widely used xylem anatomical features. Earlywood had significantly lower PERPAR (5.77 vs. 6.18%) and marginally fewer initiating rays than latewood. In comparison to tangential sections, PERPAR was systematically overestimated (6.50 vs. 4.92%) and required approximately twice the sample area for similar accuracy. Radial cuttings

Assessing Conifer Ray Parenchyma for Ecological Studies: Pitfalls and Guidelines

PubMed Central

von Arx, Georg; Arzac, Alberto; Olano, José M.; Fonti, Patrick

2015-01-01

Ray parenchyma is an essential tissue for tree functioning and survival. This living tissue plays a major role for storage and transport of water, nutrients, and non-structural carbohydrates (NSC), thus regulating xylem hydraulics and growth. However, despite the importance of rays for tree carbon and water relations, methodological challenges hamper knowledge about ray intra- and inter-tree variability and its ecological meaning. In this study we provide a methodological toolbox for soundly quantifying spatial and temporal variability of different ray features. Anatomical ray features were surveyed in different cutting planes (cross-sectional, tangential, and radial) using quantitative image analysis on stem-wood micro-sections sampled from 41 mature Scots pines (Pinus sylvestris). The percentage of ray surface (PERPAR), a proxy for ray volume, was compared among cutting planes and between early- and latewood to assess measurement-induced variability. Different tangential ray metrics were correlated to assess their similarities. The accuracy of cross-sectional and tangential measurements for PERPAR estimates as a function of number of samples and the measured wood surface was assessed using bootstrapping statistical technique. Tangential sections offered the best 3D insight of ray integration into the xylem and provided the most accurate estimates of PERPAR, with 10 samples of 4 mm2 showing an estimate within ±6.0% of the true mean PERPAR (relative 95% confidence interval, CI95), and 20 samples of 4 mm2 showing a CI95 of ±4.3%. Cross-sections were most efficient for establishment of time series, and facilitated comparisons with other widely used xylem anatomical features. Earlywood had significantly lower PERPAR (5.77 vs. 6.18%) and marginally fewer initiating rays than latewood. In comparison to tangential sections, PERPAR was systematically overestimated (6.50 vs. 4.92%) and required approximately twice the sample area for similar accuracy. Radial cuttings

Wood Anatomy and Insect Defoliator Systems: Is there an anatomical response to sustained feeding by the western spruce budworm (Choristoneura occidentalis) on Douglas-fir (Pseudotusga menziesii)?

NASA Astrophysics Data System (ADS)

Axelson, Jodi; Gärtner, Holger; Alfaro, René; Smith, Dan

2013-04-01

), cell wall thickness (µm), lumen diameter (µm), and total cell width (µm) were measured. Preliminary results indicate that earlywood parameters remain quite stable during WSB outbreak, while latewood parameters such as secondary cell wall thickness and cell length undergo step shifts at the beginning and end of outbreaks. These parameters, tree-level data, and annual defoliation data will further be tested to determine if changes in stem wood anatomy during WSB outbreaks were statistically significant.

Seasonally Resolved Oxygen Isotope Paleoclimate Proxy in Tree-Ring Cellulose from the Southeastern U.S.

NASA Astrophysics Data System (ADS)

Miller, D. L.; Mora, C. I.; Grissino-Mayer, H. D.; Mock, C. J.

2004-12-01

Stable isotopes in precipitation reflect changes in climate, moisture source, and extreme events such as tropical cyclones, and an oxygen isotope proxy record of these changes through time and space is preserved in tree-ring cellulose. Extreme climate events such as droughts and hurricanes are formidable natural disasters in the southeastern United States, and considerable efforts have been made to understand factors controlling their frequency, whether natural or anthropogenic. Tree rings offer an unusually well-resolved, dateable record of climate events extending beyond modern or historical (documentary) records. Oxygen isotopes in alpha-cellulose of shallowly-rooted conifers predominately reflect the composition of precipitation. Tropical storm convection results in marked 18O depletion in storm precipitation, to -15‰ relative to source seawater (~0‰ ). The depletion increases towards the eyewall of the cyclone, however, isotopically depleted precipitation may extend outward many 100's of km. Storm water 18O depletion translates to soil water 18O depletion that may persist for many weeks until ameliorated by soil water evaporation. Tree growth during that time will take up the anomalous isotopic compositions. Distinctive earlywood (EW ~March-June) versus latewood (LW ~July-October) growth allows the rings to be resolved at an intra-annual (seasonal) scale. By comparison to average soil water, droughts result in 18O-enriched soil water compositions. Seasonal drought or years of continued drought will be similarly captured in the isotope compositions of tree-ring cellulose. A 227-year (1770-1997) seasonally-resolved record of tropical cyclone and drought activity was obtained from cross-sections of felled slash pines (Pinus elliottii Engelm.) and remnant longleaf pines (Pinus palustris Mill.) from southern Georgia. Interpretations of drought or hurricane events were tested by comparison with recent, detailed meteorological records. The 227-year record reveals

Influence of drought on radial stem growth of Scots pine (Pinus sylvestris) in an inner Alpine environment

NASA Astrophysics Data System (ADS)

Oberhuber, Walter; Gruber, Andreas

2010-05-01

Radial stem growth indices of trees are known to be valuable long-term measures of overall tree vigor and are frequently applied to identify the climatic factors limiting tree growth. Based on several tree-ring studies conducted within inner-Alpine dry valleys, it is well established that growth of Pinus sylvestris is primarily limited by spring precipitation (April through June) and severe drought results in abrupt growth reductions and increased tree mortality. However, the record breaking heat-wave in summer 2003 had only minor impact on growth of drought exposed coniferous trees within the dry inner-Alpine valley of the Inn river (750 m a.s.l., Tyrol, Austria), where mean annual precipitation and temperature amount to 716 mm and 7.3 °C, respectively. To examine short-term influences of drought stress on growth processes more closely, we determined the influence of meteorological factors (air temperature, precipitation) and soil moisture on intra-annual dynamics of tree ring development and stem radial growth in Pinus sylvestris at two sites differing in soil moisture characteristics (xeric and dry-mesic). Radial stem development was continuously followed during 2007 and 2008 by band dendrometers and repeated micro-sampling of the developing tree ring of mature trees. In 2007, when air temperature at the beginning of the growing season in April exceeded long-term mean by 6.4 °C, cambial cell division started in early April at both study plots. A delayed onset of cambial activity of c. 2 wk was found in 2008, when average climate conditions prevailed in spring, suggesting that resumption of cambial cell division after winter dormancy is temperature-controlled. Wood formation stopped c. 4 wk earlier at the xeric compared to dry-mesic site in both study years, which indicates a strong influence of drought stress on cell differentiation processes. This is supported by radial widths of earlywood cells, which were found to be significantly narrower at the xeric

Neutron Scattering Studies of Nano-Scale Wood-Water Interactions

NASA Astrophysics Data System (ADS)

Plaza Rodriguez, Nayomi Z.

Understanding and controlling water in wood is critical to both improving forest products moisture durability and developing new sustainable forest products-based technologies. While wood is known to be hygroscopic, there is still a lack of understanding on the nanoscale wood-water interactions necessary for increased moisture-durability and dimensional stability. My PhD thesis focuses on the development and implementation of neutron scattering methods that can provide insight on both the structural and dynamical changes associated with these interactions so that products with improved moisture durability can be developed efficiently. Using small angle neutron scattering (SANS) and a custom-built in situ relative humidity chamber I studied the anisotropic moisture-induced swelling of wood nanostructure. First, I studied the effects of sample preparation by comparing SANS patterns of wiley milled wood and intact latewood cell walls, and found that scattering from intact wood provide more information about the spatial arrangement of the wood nanostructures inside the cell wall. Comparisons between SANS patterns from earlywood and latewood, also showed that the higher cell wall density of latewood cell walls results in patterns with more pronounced anisotropic features. Then, by measuring latewood loblolly pine sections obtained from the same growth ring and prepared in each of the primary wood planes, I tracked the cellulose elementary fibril spacing as a function of humidity in both intact and partially cut cell walls. These studies showed that even though swelling at the elementary fibril spacing is responsible for the majority of the transverse swelling observed at the S2 level, it is not primary plane dependent. Additionally, there were no differences in the elementary fibril spacing between partially-cut and intact cell walls, except at high humidity where the spacing in partially-cut cells was higher. SANS was also used to study the effects of two chemical