Sample records for pulvinus

  1. Growth dynamics and cytoskeleton organization during stem maturation and gravity-induced stem bending in Zea mays L

    NASA Technical Reports Server (NTRS)

    Collings, D. A.; Winter, H.; Wyatt, S. E.; Allen, N. S.; Davies, E. (Principal Investigator)

    1998-01-01

    Characterization of gravitropic bending in the maize stem pulvinus, a tissue that functions specifically in gravity responses, demonstrates that the pulvinus is an ideal system for studying gravitropism. Gravistimulation during the second of three developmental phases of the pulvinus induces a gradient of cell elongation across the non-growing cells of the pulvinus, with the most elongation occurring on the lower side. This cell elongation is spatially and temporally separated from normal internodal cell elongation. The three characterized growth phases in the pulvinus correspond closely to a specialized developmental sequence in which structural features typical of cells not fully matured are retained while cell maturation occurs in surrounding internodal and nodal tissue. For example, the lignification of supporting tissue and rearrangement of transverse microtubules to oblique that occur in the internode when cell elongation ceases are delayed for up to 10 d in the adjacent cells of the pulvinus, and only occurs as a pulvinus loses its capacity to respond to gravistimulation. Gravistimulation does not modify this developmental sequence. Neither wall lignification nor rearrangement of transverse microtubules occurs in the rapidly elongating lower side or non-responsive upper side of the pulvinus until the pulvinus loses the capacity to bend further. Gravistimulation does, however, lead to the formation of putative pit fields within the expanding cells of the pulvinus.

  2. Mechanical and electrical anisotropy in Mimosa pudica pulvini

    PubMed Central

    Foster, Justin C; Baker, Kara D; Markin, Vladislav S

    2010-01-01

    Thigmonastic or seismonastic movements in Mimosa pudica, such as the response to touch, appear to be regulated by electrical, hydrodynamical and chemical signal transduction. The pulvinus of Mimosa pudica shows elastic properties, and we found that electrically or mechanically induced movements of the petiole were accompanied by a change of the pulvinus shape. As the petiole falls, the volume of the lower part of the pulvinus decreases and the volume of the upper part increases due to the redistribution of water between the upper and lower parts of the pulvinus. This hydroelastic process is reversible. During the relaxation of the petiole, the volume of the lower part of the pulvinus increases and the volume of the upper part decreases. Redistribution of ions between the upper and lower parts of a pulvinus causes fast transport of water through aquaporins and causes a fast change in the volume of the motor cells. Here, the biologically closed electrochemical circuits in electrically and mechanically anisotropic pulvini of Mimosa pudica are analyzed using the charged capacitor method for electrostimulation at different voltages. Changing the polarity of electrodes leads to a strong rectification effect in a pulvinus and to different kinetics of a capacitor discharge if the applied initial voltage is 0.5 V or higher. The electrical properties of Mimosa pudica's pulvini were investigated and the equivalent electrical circuit within the pulvinus was proposed to explain the experimental data. The detailed mechanism of seismonastic movements in Mimosa pudica is discussed. PMID:20855975

  3. Mechanical and electrical anisotropy in Mimosa pudica pulvini.

    PubMed

    Volkov, Alexander G; Foster, Justin C; Baker, Kara D; Markin, Vladislav S

    2010-10-01

    Thigmonastic or seismonastic movements in Mimosa pudica, such as the response to touch, appear to be regulated by electrical, hydrodynamical, and chemical signal transduction. The pulvinus of Mimosa pudica shows elastic properties, and we found that electrically or mechanically induced movements of the petiole were accompanied by a change of the pulvinus shape. As the petiole falls, the volume of the lower part of the pulvinus decreases and the volume of the upper part increases due to the redistribution of water between the upper and lower parts of the pulvinus. This hydroelastic process is reversible. During the relaxation of the petiole, the volume of the lower part of the pulvinus increases and the volume of the upper part decreases. Redistribution of ions between the upper and lower parts of a pulvinus causes fast transport of water through aquaporins and causes a fast change in the volume of the motor cells. Here, the biologically closed electrochemical circuits in electrically and mechanically anisotropic pulvini of Mimosa pudica are analyzed using the charged capacitor method for electrostimulation at different voltages. Changing the polarity of electrodes leads to a strong rectification effect in a pulvinus and to different kinetics of a capacitor discharge if the applied initial voltage is 0.5 V or higher. The electrical properties of Mimosa pudica's pulvini were investigated and the equivalent electrical circuit within the pulvinus was proposed to explain the experimental data. The detailed mechanism of seismonastic movements in Mimosa pudica is discussed. © 2010 Landes Bioscience

  4. Pulvinus activity, leaf movement and leaf water-use efficiency of bush bean (Phaseplus vulgaris L.) in a hot environment.

    PubMed

    Raeini-Sarjaz, Mahmoud; Chalavi, Vida

    2008-11-01

    Pulvinus activity of Phaseolus species in response to environmental stimuli plays an essential role in heliotropic leaf movement. The aims of this study were to monitor the continuous daily pulvinus movement and pulvinus temperature, and to evaluate the effects of leaf movements, on a hot day, on instantaneous leaf water-use efficiency (WUEi), leaf gas exchange, and leaf temperature. Potted plants of Phaseolus vulgaris L. var. Provider were grown in Chicot sandy loam soil under well-watered conditions in a greenhouse. When the second trifoliate leaf was completely extended, one plant was selected to measure pulvinus movement using a beta-ray gauging (BRG) meter with a point source of thallium-204 (204Tl). Leaf gas exchange measurements took place on similar leaflets of three plants at an air temperature interval of 33-42 degrees C by a steady-state LI-6200 photosynthesis system. A copper-constantan thermocouple was used to monitor pulvinus temperature. Pulvinus bending followed the daily diurnal rhythm. Significant correlations were found between the leaf-incident angle and the stomatal conductance (R2 = 0.54; P < 0.01), and photosynthesis rate (R2 = 0.84; P < 0.01). With a reduction in leaf-incidence angle and increase in air temperature, WUEi was reduced. During the measurements, leaf temperature remained below air temperature and was a significant function of air temperature (r = 0.92; P < 0.01). In conclusion, pulvinus bending followed both light intensity and air temperature and influenced leaf gas exchange.

  5. How cereal grass shoots perceive and respond to gravity

    NASA Technical Reports Server (NTRS)

    Kaufman, P. B.; Brock, T. G.; Song, I.; Rho, Y. B.; Ghosheh, N. S.

    1987-01-01

    The leaf-sheath pulvinus of grasses presents a unique system for studying gravitropism, primarily because of its differences from other organs. The mature pulvinus is a discrete organ specialized for gravitropism: it is nongrowing in the absence of gravistimulation and capable of displaying a graviresponse independent of the rest of the plant. In this paper we present a model for gravitropism in pulvini based on recent findings from studies on the mechanisms of graviperception and graviresponse. According to this model, amyloplasts play an essential role in perceiving a change in the orientation of the pulvinus. The perception of this reorientation leads to the enhanced synthesis and release from conjugate of the auxin IAA, and the increased conjugation of gibberellin, on a localized basis. Because there is a graded growth promotion across the gravistimulated pulvinus, it is suggested that the observed hormonal asymmetry is actually an indication of a linear gradient of hormone concentration, as well as hormone response, across the pulvinus. It is further suggested that the linear gradient of hormone concentration may be predominantly the result of local changes in hormone level, rather than a product of hormonal movement into or across the pulvinus.

  6. Morphing structures and signal transduction in Mimosa pudica L. induced by localized thermal stress.

    PubMed

    Volkov, Alexander G; O'Neal, Lawrence; Volkova, Maia I; Markin, Vladislav S

    2013-10-15

    Leaf movements in Mimosa pudica, are in response to thermal stress, touch, and light or darkness, appear to be regulated by electrical, hydrodynamical, and chemical signal transduction. The pulvinus of the M. pudica shows elastic properties. We have found that the movements of the petiole, or pinnules, are accompanied by a change of the pulvinus morphing structures. After brief flaming of a pinna, the volume of the lower part of the pulvinus decreases and the volume of the upper part increases due to the redistribution of electrolytes between these parts of the pulvinus; as a result of these changes the petiole falls. During the relaxation of the petiole, the process goes in the opposite direction. Ion and water channel blockers, uncouplers as well as anesthetic agents diethyl ether or chloroform decrease the speed of alert wave propagation along the plant. Brief flaming of a pinna induces bidirectional propagation of electrical signal in pulvini. Transduction of electrical signals along a pulvinus induces generation of an action potential in perpendicular direction between extensor and flexor sides of a pulvinus. Inhibition of signal transduction and mechanical responses in M. pudica by volatile anesthetic agents chloroform or by blockers of voltage gated ion channels shows that the generation and propagation of electrical signals is a primary effect responsible for turgor change and propagation of an excitation. There is an electrical coupling in a pulvinus similar to the electrical synapse in the animal nerves. Copyright © 2013 Elsevier GmbH. All rights reserved.

  7. Gravity-stimulated changes in auxin and invertase gene expression in maize pulvinal cells

    NASA Technical Reports Server (NTRS)

    Long, Joanne C.; Zhao, Wei; Rashotte, Aaron M.; Muday, Gloria K.; Huber, Steven C.; Brown, C. S. (Principal Investigator)

    2002-01-01

    Maize (Zea mays) stem gravitropism involves differential elongation of cells within a highly specialized region, the stem internodal pulvinus. In the present study, we investigated factors that control gravitropic responses in this system. In the graviresponding pulvinus, hexose sugars (D-Glc and D-Fru) accumulated asymmetrically across the pulvinus. This correlated well with an asymmetric increase in acid invertase activity across the pulvinus. Northern analyses revealed asymmetric induction of one maize acid invertase gene, Ivr2, consistent with transcriptional regulation by gravistimulation. Several lines of evidence indicated that auxin redistribution, as a result of polar auxin transport, is necessary for gravity-stimulated Ivr2 transcript accumulation and differential cell elongation across the maize pulvinus. First, the auxin transport inhibitor, N-1-naphthylphthalamic acid, inhibited gravistimulated curvature and Ivr2 transcript accumulation. Second, a transient gradient of free indole-3-acetic acid (IAA) across the pulvinus was apparent shortly after initiation of gravistimulation. This temporarily free IAA gradient appears to be important for differential cell elongation and Ivr2 transcript accumulation. This is based on the observation that N-1-naphthylphthalamic acid will not inhibit gravitropic responses when applied to pulvinus tissue after the free IAA gradient peak has occurred. Third, IAA alone can stimulate Ivr2 transcript accumulation in non-gravistimulated pulvini. The gravity- and IAA-stimulated increase in Ivr2 transcripts was sensitive to the protein synthesis inhibitor, cycloheximide. Based on these results, a two-phase model describing possible relationships between gravitropic curvature, IAA redistribution, and Ivr2 expression is presented.

  8. Cell wall and enzyme changes during the graviresponse of the leaf-sheath pulvinus of oat (Avena sativa)

    NASA Technical Reports Server (NTRS)

    Gibeaut, David M.; Karuppiah, Nadarajah; Chang, S.-R.; Brock, Thomas G.; Vadlamudi, Babu; Kim, Donghern; Ghosheh, Najati S.; Rayle, David L.; Carpita, Nicholas C.; Kaufman, Peter B.

    1990-01-01

    The graviresponse of the leaf-sheath pulvinus of oat (Avena sativa) involves an asymmetric growth response and asymmetric processes involving degradation of starch and cell wall synthesis. Cellular and biochemical events were studied by investigation of the activities of related enzymes and changes in cell walls and their constituents. It is suggested that an osmotic potential gradient acts as the driving factor for growth, while wall extensibility is a limiting factor in pulvinus growth.

  9. TCP transcription factor, BRANCH ANGLE DEFECTIVE 1 (BAD1), is required for normal tassel branch angle formation in maize.

    PubMed

    Bai, Fang; Reinheimer, Renata; Durantini, Diego; Kellogg, Elizabeth A; Schmidt, Robert J

    2012-07-24

    In grass inflorescences, a structure called the "pulvinus" is found between the inflorescence main stem and lateral branches. The size of the pulvinus affects the angle of the lateral branches that emerge from the main axis and therefore has a large impact on inflorescence architecture. Through EMS mutagenesis we have identified three complementation groups of recessive mutants in maize having defects in pulvinus formation. All mutants showed extremely acute tassel branch angles accompanied by a significant reduction in the size of the pulvinus compared with normal plants. Two of the complementation groups correspond to mutations in the previously identified genes, RAMOSA2 (RA2) and LIGULELESS1 (LG1). Mutants corresponding to a third group were cloned using mapped-based approaches and found to encode a new member of the plant-specific TCP (TEOSINTE BRANCHED1/CYCLOIDEA/PROLIFERATING CELL NUCLEAR ANTIGEN FACTOR) family of DNA-binding proteins, BRANCH ANGLE DEFECTIVE 1 (BAD1). BAD1 is expressed in the developing pulvinus as well as in other developing tissues, including the tassels and juvenile leaves. Both molecular and genetics studies show that RA2 is upstream of BAD1, whereas LG1 may function in a separate pathway. Our findings demonstrate that BAD1 is a TCP class II gene that functions to promote cell proliferation in a lateral organ, the pulvinus, and influences inflorescence architecture by impacting the angle of lateral branch emergence.

  10. X-ray CT and histological imaging of xylem vessels organization in Mimosa pudica.

    PubMed

    Lee, Sang Joon; Song, Kahye; Kim, Hae Koo; Park, Joonghyuk

    2013-11-01

    Mimosa pudica has three distinct specialized organs, namely, pulvinus, secondary pulvinus, and pulvinule, which are respectively controlling the movements of petioles, leaflets, and pinna in response to external stimuli. Water flow is a key factor for such movements, but detailed studies on the organization of the vascular system for water transport in these organs have not been published yet. In this study, organizations of the xylem vessels and morphological features of the pulvinus, the secondary pulvinus, and the pulvinule were experimentally investigated by X-ray computed tomography and histological technique. Results showed that the xylem vessels were circularly distributed in the specialized motile organs and reorganized into distinct vascular bundles at the extremities. The number and the total cross-sectional area of the xylem vessels were increased inside the specialized motile organs. Morphological characteristics obtained in this study provided new insight to understand the functions of the vascular networks in the dynamic movements of M. pudica. Copyright © 2013 Wiley Periodicals, Inc.

  11. Cell Wall Modifications in Maize Pulvini in Response to Gravitational Stress1[W][OA

    PubMed Central

    Zhang, Qisen; Pettolino, Filomena A.; Dhugga, Kanwarpal S.; Rafalski, J. Antoni; Tingey, Scott; Taylor, Jillian; Shirley, Neil J.; Hayes, Kevin; Beatty, Mary; Abrams, Suzanne R.; Zaharia, L. Irina; Burton, Rachel A.; Bacic, Antony; Fincher, Geoffrey B.

    2011-01-01

    Changes in cell wall polysaccharides, transcript abundance, metabolite profiles, and hormone concentrations were monitored in the upper and lower regions of maize (Zea mays) pulvini in response to gravistimulation, during which maize plants placed in a horizontal position returned to the vertical orientation. Heteroxylan levels increased in the lower regions of the pulvini, together with lignin, but xyloglucans and heteromannan contents decreased. The degree of substitution of heteroxylan with arabinofuranosyl residues decreased in the lower pulvini, which exhibited increased mechanical strength as the plants returned to the vertical position. Few or no changes in noncellulosic wall polysaccharides could be detected on the upper side of the pulvinus, and crystalline cellulose content remained essentially constant in both the upper and lower pulvinus. Microarray analyses showed that spatial and temporal changes in transcript profiles were consistent with the changes in wall composition that were observed in the lower regions of the pulvinus. In addition, the microarray analyses indicated that metabolic pathways leading to the biosynthesis of phytohormones were differentially activated in the upper and lower regions of the pulvinus in response to gravistimulation. Metabolite profiles and measured hormone concentrations were consistent with the microarray data, insofar as auxin, physiologically active gibberellic acid, and metabolites potentially involved in lignin biosynthesis increased in the elongating cells of the lower pulvinus. PMID:21697508

  12. Kinetic Induction of Oat Shoot Pulvinus Invertase mRNA by Gravistimulation and Partial cDNA Cloning by the Polymerase Chain Reaction

    NASA Technical Reports Server (NTRS)

    Wu, Liu-Lai; Song, Il; Karuppiah, Nadarajah; Kaufman, Peter B.

    1993-01-01

    An asymmetric (top vs. bottom halves of pulvini) induction of invertase mRNA by gravistimulation was analyzed in oat shoot pulvini. Total RNA and poly(A)(+) RNA, isolated from oat pulvini, and two oli-gonucleotide primers, corresponding to two conserved amino acid sequences (NDPNG and WECPD) found in invertase from other species, were used for the polymerase chain reaction (PCR). A partial length cDNA (550 bp) was obtained and characterized. A 62% nucleotide sequence homology and 58% deduced amino acid sequence homology, as compared to beta-fructosidase of carrot cell wall, was found. Northern blot analysis showed that there was an obviously transient induction of invertase mRNA by gravistimulation in the oat pulvinus system. The mRNA was rapidly induced to a maximum level at 1 hour after gravistimulation treatment and gradually decreased afterwards. The mRNA level in the bottom half of the oat pulvinus was significantly higher than that in the top half of the pulvinus tissue. The kinetic induction of invertase mRNA was consistent with the transient accumulation of invertase activity during the graviresponse of the pulvinus. This indicates that the expression of the invertase gene(s) could be regulated by gravistimulation at the transcriptional level. Southern blot analysis showed that there were two to three genomic DNA fragments which hybridized with the partial-length invertase cDNA.

  13. Differentially expressed proteins of soybean (Glycine max) pulvinus in light and dark conditions

    USDA-ARS?s Scientific Manuscript database

    Soybean leaves (Glycine max) both track and avoid the sun through turgor changes of the pulvinus tissue at the base of leaves. Pulvinar response is known to be affected by both diurnally varying environmental factors and circadian patterns. Differential expression of the proteins between light and d...

  14. Competency for graviresponse in the leaf-sheath pulvinus of Avena sativa: onset to loss

    NASA Technical Reports Server (NTRS)

    Brock, T. G.; Kaufman, P. B.

    1988-01-01

    The development of the leaf-sheath pulvinus of oat (Avena sativa L. cv. Victory) was studied in terms of its competency to respond to gravistimulation. Stages of onset of competency, maximum competency and loss of competency were identified, using the length of the supertending internode as a developmental marker. During the early phases in the onset of competency, the latency period between stimulus and graviresponse decreased and the steady state response rate increased significantly. When fully competent, the latency period remained constant as the plant continued to develop, suggesting that the latency period is relatively insensitive to quantitative changes (e.g., in carbohydrate or nutrient availability) at the cell level within the plant. In contrast, the response rate was found to increase with plant development, indicating that graviresponse rate is more strongly influenced by quantitative cellular changes. The total possible graviresponse of a single oat pulvinus was confirmed to be significantly less than the original presentation angle. This was shown to not result from a loss of competency, since the graviresponse could be reinitiated by increasing the presentation angle. As a result of the low overall graviresponse of individual pulvini, two or more pulvini are required to bring the plant apex to the vertical. This was determined to occur though the sequential, rather than simultaneous, action of successive pulvini, since a given pulvinus lost competency to gravirespond shortly after the next pulvinus became fully competent.

  15. A pulse of blue light induces a transient increase in activity of apoplastic K+ in laminar pulvinus of Phaseolus vulgaris L.

    PubMed

    Okazaki, Y; Azuma, K; Nishizaki, Y

    2000-02-01

    A pulse of blue light induced both a transient increase in activity of apoplastic K+ and membrane depolarization in laminar pulvinus of Phaseolus vulgaris L. This shows that blue-light-induced net efflux of K+ from motor cells is closely related to membrane depolarization.

  16. Gravity perception and asymmetric growth in plants - A model derived from the grass pulvinus

    NASA Technical Reports Server (NTRS)

    Dayanandan, P.; Franklin, C. I.; Kaufman, P. B.

    1982-01-01

    It is pointed out that gravitropic responses in plants involve asymmetric growth. On the basis of the geometry of growth response in grass leaf sheath pulvinus, a general model is proposed for gravitropism in multicellular plant organs. The negative gravitropic response of a pulvinus is a result of cell elongation involving all but the uppermost region of a horizontally placed organ. Whereas the uppermost region does not grow, the lowermost region elongates maximally. The regions between elongate to intermediate extents. An expression is given relating the angle of curvature of the organ to the diameter and initial and final lengths of the organ. It is shown that the response of the individual cells can be expressed as inherent sensitivity to gravitational stimulus according to a particular equation.

  17. Actin dynamics mediates the changes of calcium level during the pulvinus movement of Mimosa pudica

    PubMed Central

    Yao, Heng; Xu, Qiangyi

    2008-01-01

    The bending movement of the pulvinus of Mimosa pudica is caused by a rapid change in volume of the abaxial motor cells, in response to various environmental stimuli. We investigated the relationship between the actin cytoskeleton and changes in the level of calcium during rapid contractile movement of the motor cells that was induced by electrical stimulation. The bending of the pulvinus was retarded by treatments with actin-affecting reagents and calcium channel inhibitors. The actin filaments in the motor cells were fragmented in response to electrical stimulation. Further investigations were performed using protoplasts from the motor cells of M. pudica pulvini. Calcium-channel inhibitors and EGTA had an inhibitory effect on contractile movement of the protoplasts. The level of calcium increased and became concentrated in the tannin vacuole after electrical stimulation. Ruthenium Red inhibited the increase in the level of calcium in the tannin vacuole and the contractile movement of the protoplasts. However, treatment with latrunculin A abolished the inhibitory effect of Ruthenium Red. Phalloidin inhibited the contractile movement and the increase in the level of calcium in the protoplasts. Our study demonstrates that depolymerization of the actin cytoskeleton in pulvinus motor cells in response to electrical signals results in increased levels of calcium. PMID:19513198

  18. Transient dissociation of polyribosomes and concurrent recruitment of calreticulin and calmodulin transcripts in gravistimulated maize pulvini

    NASA Technical Reports Server (NTRS)

    Heilmann, I.; Shin, J.; Huang, J.; Perera, I. Y.; Davies, E.

    2001-01-01

    The dynamics of polyribosome abundance were studied in gravistimulated maize (Zea mays) stem pulvini. During the initial 15 min of gravistimulation, the amount of large polyribosomes transiently decreased. The transient decrease in polyribosome levels was accompanied by a transient decrease in polyribosome-associated mRNA. After 30 min of gravistimulation, the levels of polyribosomes and the amount of polyribosome-associated mRNA gradually increased over 24 h up to 3- to 4-fold of the initial value. Within 15 min of gravistimulation, total levels of transcripts coding for calreticulin and calmodulin were elevated 5-fold in maize pulvinus total RNA. Transcripts coding for calreticulin and calmodulin were recruited into polyribosomes within 15 min of gravistimulation. Over 4 h of gravistimulation, a gradual increase in the association of calreticulin and calmodulin transcripts with polyribosomes was seen predominantly in the lower one-half of the maize pulvinus; the association of transcripts for vacuolar invertase with polyribosomes did not change over this period. Our results suggest that within 15 min of gravistimulation, the translation of the majority of transcripts associated with polyribosomes decreased, resembling a general stress response. Recruitment of calreticulin and calmodulin transcripts into polyribosomes occurred predominantly in the lower pulvinus one-half during the first 4 h when the presentation time for gravistimulation in the maize pulvinus is not yet complete.

  19. Structure and immunocytochemical localization of photosynthetic enzymes in the lamina joint and sheath pulvinus of the C4 grass Arundinella hirta.

    PubMed

    Wakayama, Masataka; Ohnishi, Jun-ichi; Ueno, Osamu

    2013-03-01

    The C(4) grass Arundinella hirta exhibits a unique C(4) anatomy, with isolated Kranz cells (distinctive cells) and C(4)-type expression of photosynthetic enzymes in the leaf sheath and stem as well as in the leaf blade. The border zones between these organs are pale green. Those between the leaf blade and sheath and between the sheath and stem are called the lamina joint and sheath pulvinus, respectively, and are involved in gravity sensing. We investigated the structure and localization of C(3) and C(4) photosynthetic enzymes in these tissues. In both zones the epidermis lacked stomata. The inner tissue was composed of parenchyma cells and vascular bundles. The parenchyma cells were densely packed with small intercellular spaces and contained granal chloroplasts with large starch grains. No C(4)-type cellular differentiation was recognized. Western blot analysis showed that the lamina joint and pulvinus accumulated substantial amounts of phosphoenolpyruvate carboxylase (PEPC), pyruvate,Pi dikinase (PPDK), and ribulose 1,5-bisphosphate carboxylase/oxygenase (rubisco). Immunogold electron microscopy revealed PEPC in the cytosol and both PPDK and rubisco in the chloroplasts of parenchyma cells, suggesting the occurrence of C(3) and C(4) enzymes within a single type of chlorenchyma cell. These data indicate that the lamina joint and pulvinus have unique expression patterns of C(3) and C(4) enzymes, unlike those in C(4)-type anatomy.

  20. Altered growth response to exogenous auxin and gibberellic acid by gravistimulation in pulvini of Avena sativa

    NASA Technical Reports Server (NTRS)

    Brock, T. G.; Kaufman, P. B.

    1988-01-01

    Pulvini of excised segments from oats (Avena sativa L. cv Victory) were treated unilaterally with indoleacetic acid (IAA) or gibberellic acid (GA3) with or without gravistimulation to assess the effect of gravistimulation on hormone action. Optimum pulvinus elongation growth (millimeters) and segment curvature (degrees) over 24 hours were produced by 100 micromolar IAA in vertical segments. The curvature response to IAA at levels greater than 100 micromolar, applied to the lower sides of gravistimulated (90 degrees) pulvini, was significantly less than the response to identical levels in vertical segments. Furthermore, the bending response of pulvini to 100 micromolar IAA did not vary significantly over a range of presentation angles between 0 and 90 degrees. In contrast, the response to IAA at levels less than 10 micromolar, with gravistimulation, was approximately the sum of the responses to gravistimulation alone and to IAA without gravistimulation. This was observed over a range of presentation angles. Also, GA3 (0.3-30 micromolar) applied to the lower sides of horizontal segments significantly enhanced pulvinus growth and segment curvature, although exogenous GA3 over a range of concentrations had no effect on pulvinus elongation growth or segment curvature in vertical segments. The response to GA3 (10 micromolar) plus IAA (1.0 or 100 micromolar) was additive for either vertical or horizontal segments. These results indicate that gravistimulation produces changes in pulvinus responsiveness to both IAA and GA3 and that the changes are unique for each growth regulator. It is suggested that the changes in responsiveness may result from processes at the cellular level other than changes in hormonal sensitivity.

  1. The role of gravity in leaf blade curvatures

    NASA Technical Reports Server (NTRS)

    Hayes, A. B.

    1984-01-01

    In the past year we have gained useful information on several aspects of leaf blade growth. The most important observations are as follows: The C(14)-1AA moves preferentially in a gravipositive dorsiventral direction through the blade. This movement is inhibited by inversion of the blade. The responding cells in leaf blade hyponasty are in the lower epidermis and bundle sheath cells. Two additional responses in the leaf were characterized. In addition to blade curvature, the leaf shows petiole curvature and changes in the liminal angle subtended by the pulvinus. Ethylene production was studied under a number of conditions. The blade, rather than the petiole or pulvinus, is the principal site of auxin-promoted ethylene synthesis. The effects of a variety of agents on the blade, including gibberellic acid, abscisic acid, vanadate, low pH buffers, and blue light were reviewed.

  2. Signal transduction in Mimosa pudica: biologically closed electrical circuits.

    PubMed

    Volkov, Alexander G; Foster, Justin C; Markin, Vladislav S

    2010-05-01

    Biologically closed electrical circuits operate over large distances in biological tissues. The activation of such circuits can lead to various physiological and biophysical responses. Here, we analyse the biologically closed electrical circuits of the sensitive plant Mimosa pudica Linn. using electrostimulation of a petiole or pulvinus by the charged capacitor method, and evaluate the equivalent electrical scheme of electrical signal transduction inside the plant. The discharge of a 100 microF capacitor in the pulvinus resulted in the downward fall of the petiole in a few seconds, if the capacitor was charged beforehand by a 1.5 V power supply. Upon disconnection of the capacitor from Ag/AgCl electrodes, the petiole slowly relaxed to the initial position. The electrical properties of the M. pudica were investigated, and an equivalent electrical circuit was proposed that explains the experimental data.

  3. Real-time imaging of pulvinus bending in Mimosa pudica.

    PubMed

    Song, Kahye; Yeom, Eunseop; Lee, Sang Joon

    2014-09-25

    Mimosa pudica is a plant that rapidly shrinks its body in response to external stimuli. M. pudica does not perform merely simple movements, but exhibits a variety of movements that quickly change depending on the type of stimuli. Previous studies have investigated the motile mechanism of the plants from a biochemical perspective. However, an interdisciplinary study on the structural characteristics of M. pudica should be accompanied by biophysical research to explain the principles underlying such movements. In this study, the structural characteristics and seismonastic reactions of M. pudica were experimentally investigated using advanced bio-imaging techniques. The results show that the key factors for the flexible movements by the pulvinus are the following: bendable xylem bundle, expandable/shrinkable epidermis, tiny wrinkles for surface modification, and a xylem vessel network for efficient water transport. This study provides new insight for better understanding the M. pudica motile mechanism through structural modification.

  4. Mimosa pudica: Electrical and mechanical stimulation of plant movements.

    PubMed

    Volkov, Alexander G; Foster, Justin C; Ashby, Talitha A; Walker, Ronald K; Johnson, Jon A; Markin, Vladislav S

    2010-02-01

    Thigmonastic movements in the sensitive plant Mimosa pudica L., associated with fast responses to environmental stimuli, appear to be regulated through electrical and chemical signal transductions. The thigmonastic responses of M. pudica can be considered in three stages: stimulus perception, electrical signal transmission and induction of mechanical, hydrodynamical and biochemical responses. We investigated the mechanical movements of the pinnae and petioles in M. pudica induced by the electrical stimulation of a pulvinus, petiole, secondary pulvinus or pinna by a low electrical voltage and charge. The threshold value was 1.3-1.5 V of applied voltage and 2 to 10 microC of charge for the closing of the pinnules. Both voltage and electrical charge are responsible for the electro-stimulated closing of a leaf. The mechanism behind closing the leaf in M. pudica is discussed. The hydroelastic curvature mechanism closely describes the kinetics of M. pudica leaf movements.

  5. Real-time imaging of pulvinus bending in Mimosa pudica

    PubMed Central

    Song, Kahye; Yeom, Eunseop; Lee, Sang Joon

    2014-01-01

    Mimosa pudica is a plant that rapidly shrinks its body in response to external stimuli. M. pudica does not perform merely simple movements, but exhibits a variety of movements that quickly change depending on the type of stimuli. Previous studies have investigated the motile mechanism of the plants from a biochemical perspective. However, an interdisciplinary study on the structural characteristics of M. pudica should be accompanied by biophysical research to explain the principles underlying such movements. In this study, the structural characteristics and seismonastic reactions of M. pudica were experimentally investigated using advanced bio-imaging techniques. The results show that the key factors for the flexible movements by the pulvinus are the following: bendable xylem bundle, expandable/shrinkable epidermis, tiny wrinkles for surface modification, and a xylem vessel network for efficient water transport. This study provides new insight for better understanding the M. pudica motile mechanism through structural modification. PMID:25253083

  6. Molecular Basis of the Increase in Invertase Activity Elicited by Gravistimulation of Oat-Shoot Pulvini

    NASA Technical Reports Server (NTRS)

    Wu, Liu-Lai; Song, Il; Kim, Donghern; Kaufman, Peter B.

    1993-01-01

    An asymmetric (top vs. bottom) increase in invertase activity is elicited by gravistimulation in oatshoot pulvini starting within 3h after treatment. In order to analyze the regulation of invertase gene expression in this system, we examined the effect of gravistimulation on invertase mRNA induction. Total RNA and poly(A)(+)RNA, isolated from oat pulvini, and two oligonucleotide primers, corresponding to two conserved amino-acid sequences (NDPNG and WECPD) found in invertase from other species, were used for the Polymerase Chain Reaction (PCR). A partial-length cDNA (550 base pairs) was obtained and characterized. There was a 52 % deduced amino-acid sequence homology to that of carrot beta-fructosi- dase and a 48 % homology to that of tomato invertase. Northern blot analysis showed that there was an obvious transient accumulation of invertase mRNA elicited by gravistimulation of oat pulvini. The mRNA was rapidly induced to a maximum level at 1h following gravistimulation treatment and gradually decreased afterwards. The mRNA level in the bottom half of the oat pulvinus was significantly higher (five-fold) than that in the top half of the pulvinus tissue. The induction of invertase mRNA was consistent with the transient enhancement of invertase activity during the graviresponse of the pulvinus. These data indicate that the expression of the invertase gene(s) could be regulated by gravistimulation at the transcriptional and/or translational levels. Southern blot analysis showed that there were four genomic DNA fragments hybridized to the invertase cDNA. This suggests that an invertase gene family may exist in oat plants.

  7. Localization and pattern of graviresponse across the pulvinus of barley Hordeum vulgare

    NASA Technical Reports Server (NTRS)

    Brock, T. G.; Lu, C. R.; Ghosheh, N. S.; Kaufman, P. B.

    1989-01-01

    Pulvini of excised stem segments from barley (Hordeum vulgare cv Larker') were pretreated with 1 millimolar coumarin before gravistimulation to reduce longitudinal cell expansion and exaggerate radial cell enlargement. The cellular localization and pattern of graviresponse across individual pulvini were then evaluated by cutting the organ in cross-section, photographing the cross-section, and then measuring pulvinus thickness and the radial width of cortical and epidermal cells in enlargements of the photomicrographs. With respect to orientation during gravistimulation, we designated the uppermost point of the cross-section 0 degrees and the lowermost point 180 degrees. A gravity-induced increase in pulvinus thickness was observable within 40 degrees of the vertical in coumarin-treated pulvini. In upper halves of coumarin-treated gravistimulated pulvini, cells in the inner cortex and inner epidermis had increased radial widths, relative to untreated gravistimulated pulvini. In lower halves of coumarin-treated pulvini, cells in the central and outer cortex and in the outer epidermis showed the greatest increase in radial width. Cells comprising the vascular bundles also increased in radial width, with this pattern following that of the central cortex. These results indicate (a) that all cell types are capable of showing a graviresponse, (b) that the graviresponse occurs in both the top and the bottom of the responding organ, and (c) that the magnitude of the response increases approximately linearly from the uppermost point to the lowermost. These results are also consistent with models of gravitropism that link the pattern and magnitude of the graviresponse to graviperception via statolith sedimentation.

  8. Conserved genetic determinant of motor organ identity in Medicago truncatula and related legumes

    PubMed Central

    Chen, Jianghua; Moreau, Carol; Liu, Yu; Kawaguchi, Masayoshi; Hofer, Julie; Ellis, Noel; Chen, Rujin

    2012-01-01

    Plants exhibit various kinds of movements that have fascinated scientists and the public for centuries. Physiological studies in plants with the so-called motor organ or pulvinus suggest that cells at opposite sides of the pulvinus mediate leaf or leaflet movements by swelling and shrinking. How motor organ identity is determined is unknown. Using a genetic approach, we isolated a mutant designated elongated petiolule1 (elp1) from Medicago truncatula that fails to fold its leaflets in the dark due to loss of motor organs. Map-based cloning indicated that ELP1 encodes a putative plant-specific LOB domain transcription factor. RNA in situ analysis revealed that ELP1 is expressed in primordial cells that give rise to the motor organ. Ectopic expression of ELP1 resulted in dwarf plants with petioles and rachises reduced in length, and the epidermal cells gained characteristics of motor organ epidermal cells. By identifying ELP1 orthologs from other legume species, namely pea (Pisum sativum) and Lotus japonicus, we show that this motor organ identity is regulated by a conserved molecular mechanism. PMID:22689967

  9. Early membrane events induced by salicylic acid in motor cells of the Mimosa pudica pulvinus.

    PubMed

    Saeedi, Saed; Rocher, Françoise; Bonmort, Janine; Fleurat-Lessard, Pierrette; Roblin, Gabriel

    2013-04-01

    Salicylic acid (o-hydroxy benzoic acid) (SA) induced a rapid dose-dependent membrane hyperpolarization (within seconds) and a modification of the proton secretion (within minutes) of Mimosa pudica pulvinar cells at concentrations higher than 0.1mM. Observations on plasma membrane vesicles isolated from pulvinar tissues showed that SA acted directly at the membrane level through a protonophore action as suggested by the inhibition of the proton gradient and the lack of effect on H(+)-ATPase catalytic activity. Comparative data obtained with protonophores (carbonylcyanide-m-chlorophenylhydrazone and 2,4-dinitrophenol) and inhibitors of ATPases (vanadate, N,N'-dicyclohexylcarbodiimide, and diethylstilbestrol) corroborated this conclusion. Consequently, the collapse of the proton motive force led to an impairment in membrane functioning. This impairment is illustrated by the inhibition of the ion-driven turgor-mediated seismonastic reaction of the pulvinus following SA treatment. SA acted in a specific manner as its biosynthetic precursor benzoic acid induced much milder effects and the m- and p-OH benzoic acid derivatives did not trigger similar characteristic effects. Therefore, SA may be considered both a membrane signal molecule and a metabolic effector following its uptake in the cells.

  10. Effect of dark pretreatment on the kinetics of response of barley pulvini to gravistimulation and hormones

    NASA Technical Reports Server (NTRS)

    Brock, T. G.; Kaufman, P. B.

    1988-01-01

    Starch in pulvinus amyloplasts of barley (Hordeum vulgare cv Larker) disappears when 45-day-old, light-grown plants are given 5 days of continuous darkness. The effect of this loss on the pulvinus graviresponse was evaluated by following changes in the kinetics of response during the 5-day dark period. Over 5 days of dark pretreatment, the lag to initial graviresponse and the subsequent half-time to maximum steady state bending rate increased significantly while the maximum bending rate did not change. The change in response to applied indoleacetic acid (100 micromolar) plus gibberellic acid (10 micromolar) without gravistimulation, under identical dark pretreatments, was used as a model system for the response component of gravitropism. Dark pretreatment did not change the lag to initial response following hormone application to vertical pulvini, but both the maximum bending rate and the half-time to the maximum rate were significantly reduced. Also, after dark pretreatment, significant bending responses following hormone application were observed in vertical segments with or without added sucrose, while gravistimulation produced a response only if segments were given sucrose. These results indicate that starch-filled amyloplasts are required for the graviresponse of barley pulvini and suggest that they function in the stimulus perception and signal transduction components of gravitropism.

  11. A Role for the Stele in Intertissue Signaling in the Initiation of Abscission in Bean Leaves (Phaseolus vulgaris L.).

    PubMed Central

    Thompson, D. S.; Osborne, D. J.

    1994-01-01

    A combination of microdissection and viscometric endo-[beta]-1,4-glucanhydrolase assays was used to investigate if the early appearance of the abscission-related isoelectric point-9.5 endo-[beta]-1,4-glucanhydrolase in the stele of the pulvinus and abscission zone of the foliar abscission zone of Phaseolus vulgaris L. prior to cell separation (reported by E. del Campillo, P.D. Reid, R. Sexton, L.N.Lewis [1990] Plant Cell 2: 245-254) indicates that the vascular tissue of this region has a specific role in abscission. We find that no endo-[beta]-1,4-glucanhydrolase activity or cell separation is detectable in the abscission zone cortex if the abscission zone cortex is separated from the stele tissue. If the stele is separated from the abscission zone cortex after a lag period but again before any endo-[beta]-1,4-glucanhydrolase activity is present in the abscission zone cortex, then the enzyme is produced in the cortex and abscission ensues. We conclude that the cortex of the abscission zone is able to abscind independently of the vascular tissue only after the vascular tissue has begun to respond to abscission-promoting signals. We suggest that ethylene promotes formation of an abscission-permitting signal in the stele of the abscission zone and pulvinus, and that this signal is an essential elicitor for the synthesis of cell separation enzymes in the target cells of the abscission zone cortex. PMID:12232206

  12. Studies on Acidification of Media by Avena Stem Segments in the Presence and Absence of Gibberellic Acid 1

    PubMed Central

    Hebard, Frederick V.; Amatangelo, Steven J.; Dayanandan, P.; Kaufman, Peter B.

    1976-01-01

    The rate of acidification of media by Avena stem segments was studied with a titrimeter. GA3 increased this rate by an average of 17% if supplied to the segments 90 min prior to measurement. GA3 inhibited the rate by 15% if supplied 10 min prior to measurement. After 90 min incubation, stimulation of elongation had started; at 10 min, GA3 had not yet started to stimulate elongation in the segments. The acidification rates of the nodes (including the sheath-pulvinus), leaf sheath bases, and the internode bases of the stem segments were determined for plus and minus GA3-treated segments. The internode fraction contributes most to modification of the acidification rate, the node-pulvinus fraction less so, and the nongrowing sheath not at all. Acidification rates were measured for segments in different stages of elongation (lag, log, and plateau phases of growth). Segments in these growth stages were obtained from intact plants and from segments preincubated in sucrose and sucrose + GA3. Segments from all sources which are in the log phase of growth have the highest rates, those in the plateau phase the lowest. For lag and log growth phases, segments preincubated in sucrose + GA3 show the highest rates, those preincubated in sucrose the lowest rates. The opposite occurs for segments in the plateau phase of growth. Segments stimulated to grow by GA3 cause the pH of their incubation media to drop to pH 5.15 from an initial pH of 6.5. Nonstimulated segments cause a drop to pH 5.6. Long term growth of the segments is maximal in media buffered to pH 5 in the presence and absence of GA3. Our results support the idea that GA3 stimulates an active acidification process in Avena stem segments just after GA3 starts to stimulate growth in the segments, and that such an acidification process could play an important role in wall-loosening during active growth of the internode. PMID:16659741

  13. Leaf movement in Calathea lutea (Marantaceae).

    PubMed

    Herbert, Thomas J; Larsen, Parry B

    1985-09-01

    Calathea lutea is a broad-leaved, secondary successional plant which shows complex leaf movements involving both elevation and folding of the leaf surface about the pulvinus. In the plants studied, mean leaf elevation increased from approximately 34 degrees in the early morning to 70 degrees at noon while the angle of leaf folding increased from 13 degrees to 50 degrees over the same time period. During the period from early morning to noon, these movements resulted in a significant decrease in the cosine of the angle of incidence, a measure of the direct solar radiation intercepted. The observed changes in elevational angle significantly reduce the cosine of angle of incidence while folding does not significantly reduce the fraction of direct solar radiation intercepted during the period of direct exposure of the leaf surface to the solar beam. Since elevational changes seem to account for the reduction in exposure to direct solar radiation, the role of folding remains unclear.

  14. PHANTASTICA regulates leaf polarity and petiole identity in Medicago truncatula

    PubMed Central

    Ge, Liangfa; Chen, Rujin

    2014-01-01

    Establishment of proper polarities along the adaxial-abaxial, proximodistal, and medial-lateral axes is a critical step for the expansion of leaves from leaf primordia. It has been shown that the MYB domain protein, ASYMMETRIC LEAVES1/ROUGH SHEATH2/PHANTASTICA (collectively named ARP) plays an important role in this process. Loss of function of ARP leads to severe leaf polarity defects, such as abaxialized or needle-like leaves. In addition to its role in leaf polarity establishment, we have recently shown that the Medicago truncatula ARP gene, MtPHAN, also plays a role in leaf petiole identity regulation. We show that a mutation of MtPHAN results in petioles acquiring characteristics of the motor organ, pulvinus, including small epidermal cells with extensive cell surface modifications and altered vascular tissue development. Taken together, our results reveal a previously unidentified function of ARP in leaf development. PMID:24603499

  15. Mechanical stress caused by wind on leaves of Theobroma cacao: Photosynthetic, molecular, antioxidative and ultrastructural responses.

    PubMed

    Reis, Graciele Santos Monteiro; Almeida, Alex-Alan Furtado de; Mangabeira, Pedro Antônio Oliveira; Santos, Ivanildes Conceição Dos; Pirovani, Carlos Priminho; Ahnert, Dário

    2018-01-01

    Theobroma cacao is cultivated in the shade, in a so-called 'Cabruca' system, in intercropped with Erithryna or other tree species of economic value, and in full sun as a monoculture in irrigated or chemically-irrigated systems. Since it is a species quite intolerant to wind, it is practically impossible to implant cacao crops under full exposure to the sun, or in areas of frequent winds, without the protection of windbreaks, using arboreal species around the area of culture in the form of box. Wind can cause mechanical stimuli in plants, affecting their growth and development. The objective of this work was to evaluate the photosynthetic changes in mature leaves and the molecular, biochemical and ultrastructural changes in young and mature leaves of the CCN 51 cloned genotype of T. cacao subjected to intermittent (IW) and constant (CW) wind, with velocities of 2.5, 3.5 and 4.5 m s-1, during 3, 6 and 12 h of exposure. It was verified that CW and IW, considering different exposure times, interfered directly in stomatal conductance (gs), transpiration (E) and water use efficiency (WUE), causing a reduction of the photosynthetic rate (A) in mature leaves. In addition, the pulvinus and blade of young and mature leaves, exposed to IW and CW with different exposure times (3 and 12 h), showed marked macroscopic and microscopic mechanical injuries resulting from the constant leaf movement. At both speeds, there was rupture of the cell nuclear membrane in pulvinus and the mesophyll tissues, mainly in the young leaves. On the other hand, in young and mature leaves exposed to CW and IW at different speeds and exposure times, there was lipid peroxidation, increased activity of guaiacol (GPX) and ascorbate (APX) peroxidases in most treatments; and altered expression of transcripts of psba and psbo genes related to the phothosynthetic apparatus and Cu-Zn-sod and per genes related to antioxidative enzymes at the rate of 4.5 m s-1. Younger leaves were more intolerant to mechanical stress caused by the wind, since presented greater macro and microscopic damages and, consequently, greater molecular, biochemical and ultrastructural changes. High wind speeds can seriously compromise the development of young leaves of T. cacao plants and affect their productivity.

  16. Circadian variations in biologically closed electrochemical circuits in Aloe vera and Mimosa pudica.

    PubMed

    Volkov, Alexander G; Baker, Kara; Foster, Justin C; Clemmons, Jacqueline; Jovanov, Emil; Markin, Vladislav S

    2011-04-01

    The circadian clock regulates a wide range of electrophysiological and developmental processes in plants. This paper presents, for the first time, the direct influence of a circadian clock on biologically closed electrochemical circuits in vivo. Here we show circadian variation of the plant responses to electrical stimulation. The biologically closed electrochemical circuits in the leaves of Aloe vera and Mimosa pudica, which regulate their physiology, were analyzed using the charge stimulation method. The electrostimulation was provided with different timing and different voltages. Resistance between Ag/AgCl electrodes in the leaf of Aloe vera was higher during the day than at night. Discharge of the capacitor in Aloe vera at night was faster than during the day. Discharge of the capacitor in a pulvinus of Mimosa pudica was faster during the day. The biologically closed electrical circuits with voltage gated ion channels in Mimosa pudica are also activated the next day, even in the darkness. These results show that the circadian clock can be maintained endogenously and has electrochemical oscillators, which can activate ion channels in biologically closed electrochemical circuits. We present the equivalent electrical circuits in both plants and their circadian variation to explain the experimental data. Copyright © 2011 Elsevier B.V. All rights reserved.

  17. Five pectinase gene expressions highly responding to heat stress in rice floral organs revealed by RNA-seq analysis.

    PubMed

    Wu, Liquan; Taohua, Zhou; Gui, Wenbin; Xu, Lisen; Li, Juan; Ding, YanFeng

    2015-07-31

    Heat stress hurts rice, and floral organs are mostly sensitive to heat stress. We aimed to unravel molecular responses to heat stress in rice floral organs using Illumina/Solexa sequencing technology for addressing the increasing concern of globle warming. At meiophase of the pollen mother cell (pulvinus flat), the plants were stressed for 3 d at 38 C, and RNA was extracted from the stressed pistil and stamen for RNA-Seq sequencing to build the heat stress transcriptom library. A total of 7178 defferentially expressed genes (DEGs) between the normal and heat stress libraries were significant, 61% up-regulated and 39% down-regulated. The 7178 DEGs were significantly classified to 34 gene ontology (GO) categories, and 11 of the GO categories were significantly enriched. The GO:0016787 for hydrolase activity of molecular function was mostly enriched with the least probability, and included 11 DEGs named Hy1 - Hy11. Expression levels of five DEGs, Hy4 - Hy6 and Hy9 - Hy10 for starch and sucrose metablism via pectinase, increased 12 - 14 times in response to the heat stress. Further investigation of the five DEGs for pectin metabolism and association with reported heat responsive genes may help develop a molecular strategy to remedy heat damage in rice. Copyright © 2015 Elsevier Inc. All rights reserved.

  18. Hormonal and gravitropic specificity in the regulation of growth and cell wall synthesis in pulvini and internodes from shoots of Avena sativa L. (oat).

    PubMed Central

    Montague, M J

    1995-01-01

    Segments can be cut from the peduncular-1 internode of oat (Avena sativa L.) shoots so as to contain the graviresponsive leaf-sheath pulvinus and gibberellin-sensitive internodal tissue. Incorporation of [14C]glucose was used to monitor cell wall synthesis in these two tissues as affected by gravistimulus, indoleacetic acid (IAA), gibberellic acid (GA3), and fusicoccin (FC). Pulvinar cell wall synthesis was promoted by IAA and FC (both within about 1 h), as well as by gravistimulus (starting between 3 and 6 h), whereas GA3 had no effect on nongravistimulated pulvini. In contrast, GA3 and FC promoted internodal cell wall synthesis (initiated between 1 and 2 h), whereas IAA and gravistimulus caused a decrease in internodal uptake. FC preferentially promoted incorporation into the matrix component of the wall in both tissues. Gravistimulus failed to increase responsiveness of pulvinar tissue to IAA, whereas GA3 partially overcame gravistimulus-promoted incorporation into pulvinar cell wall, probably because of preferential movement of label into the rapidly elongating internode. The results demonstrate that these eight stimulus/tissue combinations can be examined easily in an isolated 10-mm stem segment, providing new opportunities for the comparative study of tissue- and stimulus-specific events in gene regulation and signal transduction in agronomically important cereals. PMID:11536686

  19. DOE Office of Scientific and Technical Information (OSTI.GOV)

    Carpita, N.C.

    We have just completed the second year of a three-year project entitled Biosynthesis assembly of cell wall polysaccharides in cereal grasses.'' We made significant progress on two aspects of cell wall synthesis in grasses and greatly refined gas-liquid and high- performance liquid chromatographic techniques necessary to identify the products of synthesis in vitro and in vivo. First, Dr. David Gibeaut, a post-doctoral associate, devised a convenient procedure for the enrichment of Golgi membranes by flotation centrifugation following initial downward rate-zonal separation. Based on comparison of the IDPase marker enzyme, flotation centrifugation enriched the Golgi apparatus almost 7-fold after the initialmore » downward separation. This system is now used in our studies of the synthesis in vitro of the mixed-linkage {beta}-D-glucan. Second, Gibeaut and I have devised a simple technique to feed radioactive sugars into intact growing seedlings and follow incorporation of radioactivity into and turnover from specific cell wall polysaccharides. The project has also provided a few spin-off projects that have been productive as well. First, in collaboration with the group of Prof. Peter Kaufman, University of Michigan, we examined changes in cell wall structure concomitant with reaction to gravistimulation in the gravisensing oat pulvinus. Second, Dr. Gibeaut developed a simple clean-up procedure for partially methylated alditol derivatives to remove a large amount of undesirable interfering compounds that confound separation of the derivatives by gas-liquid chromatography. 5 refs.« less

  20. Loose Plant Architecture1, an INDETERMINATE DOMAIN Protein Involved in Shoot Gravitropism, Regulates Plant Architecture in Rice1[W

    PubMed Central

    Wu, Xinru; Tang, Ding; Li, Ming; Wang, Kejian; Cheng, Zhukuan

    2013-01-01

    Tiller angle and leaf angle are two important components of rice (Oryza sativa) plant architecture that play a crucial role in determining grain yield. Here, we report the cloning and characterization of the Loose Plant Architecture1 (LPA1) gene in rice, the functional ortholog of the AtIDD15/SHOOT GRAVITROPISM5 (SGR5) gene in Arabidopsis (Arabidopsis thaliana). LPA1 regulates tiller angle and leaf angle by controlling the adaxial growth of tiller node and lamina joint. LPA1 was also found to affect shoot gravitropism. Expression pattern analysis suggested that LPA1 influences plant architecture by affecting the gravitropism of leaf sheath pulvinus and lamina joint. However, LPA1 only influences gravity perception or signal transduction in coleoptile gravitropism by regulating the sedimentation rate of amyloplasts, distinct from the actions of LAZY1. LPA1 encodes a plant-specific INDETERMINATE DOMAIN protein and defines a novel subfamily of 28 INDETERMINATE DOMAIN proteins with several unique conserved features. LPA1 is localized in the nucleus and functions as an active transcriptional repressor, an activity mainly conferred by a conserved ethylene response factor-associated amphiphilic repression-like motif. Further analysis suggests that LPA1 participates in a complicated transcriptional and protein interaction network and has evolved novel functions distinct from SGR5. This study not only facilitates the understanding of gravitropism mechanisms but also generates a useful genetic material for rice breeding. PMID:23124325

  1. Loose Plant Architecture1, an INDETERMINATE DOMAIN protein involved in shoot gravitropism, regulates plant architecture in rice.

    PubMed

    Wu, Xinru; Tang, Ding; Li, Ming; Wang, Kejian; Cheng, Zhukuan

    2013-01-01

    Tiller angle and leaf angle are two important components of rice (Oryza sativa) plant architecture that play a crucial role in determining grain yield. Here, we report the cloning and characterization of the Loose Plant Architecture1 (LPA1) gene in rice, the functional ortholog of the AtIDD15/SHOOT GRAVITROPISM5 (SGR5) gene in Arabidopsis (Arabidopsis thaliana). LPA1 regulates tiller angle and leaf angle by controlling the adaxial growth of tiller node and lamina joint. LPA1 was also found to affect shoot gravitropism. Expression pattern analysis suggested that LPA1 influences plant architecture by affecting the gravitropism of leaf sheath pulvinus and lamina joint. However, LPA1 only influences gravity perception or signal transduction in coleoptile gravitropism by regulating the sedimentation rate of amyloplasts, distinct from the actions of LAZY1. LPA1 encodes a plant-specific INDETERMINATE DOMAIN protein and defines a novel subfamily of 28 INDETERMINATE DOMAIN proteins with several unique conserved features. LPA1 is localized in the nucleus and functions as an active transcriptional repressor, an activity mainly conferred by a conserved ethylene response factor-associated amphiphilic repression-like motif. Further analysis suggests that LPA1 participates in a complicated transcriptional and protein interaction network and has evolved novel functions distinct from SGR5. This study not only facilitates the understanding of gravitropism mechanisms but also generates a useful genetic material for rice breeding.

  2. Gross Anatomy of Pampas Deer (Ozotoceros bezoarticus, Linnaeus 1758) Mouth and Pharynx.

    PubMed

    Pérez, W; Vazquez, N; Ungerfeld, R

    2017-04-01

    The aims of this study were to describe the anatomy of the mouth and pharynx of the pampas deer, and to consider its evolutionary feeding niche according to those characteristics. Gross dissections of the mouth and pharynx were performed in 15 animals, 10 adult females and five young animals under 1 year (three males and two females), all dead by causes unrelated to this anatomical region. The upper lip entered in the constitution of a pigmented nasolabial plane. The masseter muscles weighed 43.8 ± 3.5 g and represented 0.23% of body weight, which corresponds to ruminants of feeders intermediate to grazers and browsers. Parotid glands represented 0.08% of the body weight, characteristic that also categorize the pampas deer as belonging to the intermediate feeding group. The dental formula was the same of the domestic ruminants. The upper incisors and canines were absent, and instead of them, there was a dental pad (Pulvinus dentalis). The upper canine teeth were present only in the deciduous dentition. The existence of a brachydont dentition turns Ozotoceros very vulnerable to continuous use as there is no compensatory teeth growth. The particular anatomy of the mouth and lips of this animal was adapted to a very selective feeding, taking highly nutritious sprouts beyond plant category. In conclusion and in addition to previous studies of anatomy of the digestive organs in this species, pampas deer may be categorized as belonging to the intermediate type of feeding. © 2016 Blackwell Verlag GmbH.

  3. RNA silencing in the life cycle of soybean: multiple restriction systems and spatiotemporal variation associated with plant architecture.

    PubMed

    Mori, Ayumi; Sato, Hiroshi; Kasai, Megumi; Yamada, Tetsuya; Kanazawa, Akira

    2017-06-01

    The expression of transgenes introduced into a plant genome is sometimes suppressed by RNA silencing. Although local and systemic spread of RNA silencing have been studied, little is known about the mechanisms underlying spatial and temporal variation in transgene silencing between individual plants or between plants of different generations, which occurs seemingly stochastically. Here, we analyzed the occurrence, spread, and transmission of RNA silencing of the green fluorescent protein (GFP) gene over multiple generations of the progeny of a single soybean transformant. Observation of GFP fluorescence in entire plants of the T 3 -T 5 generations indicated that the initiation and subsequent spread of GFP silencing varied between individuals, although this GFP silencing most frequently began in the primary leaves. In addition, GFP silencing could spread into the outer layer of seed coat tissues but was hardly detectable in the embryos. These results are consistent with the notion that transgene silencing involves its reset during reproductive phase, initiation after germination, and systemic spread in each generation. GFP silencing was absent in the pulvinus, suggesting that its cortical cells inhibit cell-to-cell spread or induction of RNA silencing. The extent of GFP silencing could differ between the stem and a petiole or between petiolules, which have limited vascular bundles connecting them and thus deter long-distant movement of silencing. Taken together, these observations indicate that the initiation and/or spread of RNA silencing depend on specific features of the architecture of the plant in addition to the mechanisms that can be conserved in higher plants.

  4. Cytoplasmic pH dynamics in maize pulvinal cells induced by gravity vector changes

    NASA Technical Reports Server (NTRS)

    Johannes, E.; Collings, D. A.; Rink, J. C.; Allen, N. S.; Brown, C. S. (Principal Investigator)

    2001-01-01

    In maize (Zea mays) and other grasses, changes in orientation of stems are perceived by pulvinal tissue, which responds to the stimulus by differential growth resulting in upward bending of the stem. The amyloplast-containing bundle sheath cells are the sites of gravity perception, although the initial steps of gravity perception and transmission remain unclear. In columella cells of Arabidopsis roots, we previously found that cytoplasmic pH (pH(c)) is a mediator in early gravitropic signaling (A.C. Scott, N.S. Allen [1999] Plant Physiol 121: 1291-1298). The question arises whether pH(c) has a more general role in signaling gravity vector changes. Using confocal ratiometric imaging and the fluorescent pH indicator carboxy seminaphtorhodafluor acetoxymethyl ester acetate, we measured pH(c) in the cells composing the maize pulvinus. When stem slices were gravistimulated and imaged on a horizontally mounted confocal microscope, pH(c) changes were only apparent within the bundle sheath cells, and not in the parenchyma cells. After turning, cytoplasmic acidification was observed at the sides of the cells, whereas the cytoplasm at the base of the cells where plastids slowly accumulated became more basic. These changes were most apparent in cells exhibiting net amyloplast sedimentation. Parenchyma cells and isolated bundle sheath cells did not show any gravity-induced pH(c) changes although all cell types responded to external stimuli in the predicted way: Propionic acid and auxin treatments induced acidification, whereas raising the external pH caused alkalinization. The results suggest that pH(c) has an important role in the early signaling pathways of maize stem gravitropism.

  5. Gibberellins and gravitropism in maize shoots: endogenous gibberellin-like substances and movement and metabolism of (/sup 3/)gibberellin A

    DOE Office of Scientific and Technical Information (OSTI.GOV)

    Rood, S.B.; Kaufman, P.B.; Abe, H.

    1987-03-01

    (/sup 3/H)Gibberellin A/sub 20/(GA/sub 20/) of high specific radioactivity was applied equilaterally in a ring of microdrops to the internodal pulvinus of shoots of 3-week-old vertical normal maize (Zea mays L.), and to a pleiogravitropic (prostrate) maize mutant, lazy (la). All plants converted the (/sup 3/H)GA/sub 1//sup -/ and (/sup 3/H)GA/sub 29/-like metabolites as well as to several metabolites with the partitioning and chromatographic behavior of glucosyl conjugates of (/sup 3/H)GA/sub 1/(/sup 3/H)GA/sub 29/, and (/sup 3/H)GA/sub 8/. The tentative identification of these putative (/sup 3/H)GA glucosyl conjugates was further supported by the release of the free (/sup 3/H)GA moietymore » after cleavage with cellulase. Within 12 hours of the (/sup 3/H)GA/sub 20/ feed, there was a significantly higher proportion of total radioactivity in lower than in upper halves of internode and leaf sheaf pulvini in gravistimulated normal maize. Further, there was a significantly higher proportion of putative free GA metabolites of (/sup 3/H)GA/sub 20/, especially (/sup 3/H) GA/sub 1/, in the lower halves of normal maize relative to upper halves. The differential localization of the metabolites between upper and lower halves was not apparent in the pleiogravitropic mutant, la. Endogenous GA-like substances were also examined in gravistimulated maize shoots. Forty-eight hours after gravistimulation of 3-week-old maize seedlings, endogenous free GA-like substances in upper and lower leaf sheath and internode pulvini halves were extracted, chromatographed, and bioassayed using the Tanginbozu dwarf rice microdroassay. Lower halves contained higher total levels of GA-like activity.« less

  6. In vitro Antioxidant and Pharmacognostic Studies of Leaf Extracts of Cajanus cajan (L.) Millsp.

    PubMed

    Mahitha, B; Archana, P; Ebrahimzadeh, Md H; Srikanth, K; Rajinikanth, M; Ramaswamy, N

    2015-01-01

    Cajanus cajan (L.) Millsp is one of the second most dietary legume crops. The leaf extracts may be used as a potential source of natural antioxidant. The ash values, extractive values, total phenolic and flavonoid content, in vitro antioxidant activity of various leaf extracts as well as anatomical investigation of Cajanus cajan were carried out. Physicochemical parameters such as total, acid-insoluble and water-soluble ash values and moisture content of the leaf powder of C. cajan were found to be 9.50%, 1.40 g/100 g, 4.15 g/100 g drug and 6.72%, respectively. Percent yield of acetone, aqueous, ethanol, ethyl acetate and chloroform leaf extracts were 9.0, 10.6, 13.75, 8.7 and 5.8 g/100 g, respectively. Significant amount of phenolic and flavonoid content were observed. The results of the antioxidant activity were found to be concentration-dependent. The IC50 values for DPPH assay determined for aqueous and ethanol extracts were 0.69 and 0.79 mg/ml, respectively. Reducing power is increased with increasing amount of concentration in both aqueous and ethanol leaf extracts. The highest hydroxyl radical scavenging activity reached up to 83.67% in aqueous and 78.75% in ethanol extracts and in phosphomolybdenum assay the aqueous extract showed strong antioxidant capacity up to 55.97 nM gallic acid equivalents/g. It was found that the aqueous extract possessed highest antioxidant activity in all the assays tested. The antioxidant characteristics of leaf extracts are possibly because of the presence of polyphenols. Microscopic study showed the presence of collenchyma, fibres, xylem, phloem, epidermis, trichomes, palisade tissue, basal sheath, pith and cortex in leaf, petiole and pulvinus.

  7. In vitro Antioxidant and Pharmacognostic Studies of Leaf Extracts of Cajanus cajan (L.) Millsp

    PubMed Central

    Mahitha, B.; Archana, P.; Ebrahimzadeh, MD. H.; Srikanth, K.; Rajinikanth, M.; Ramaswamy, N.

    2015-01-01

    Cajanus cajan (L.) Millsp is one of the second most dietary legume crops. The leaf extracts may be used as a potential source of natural antioxidant. The ash values, extractive values, total phenolic and flavonoid content, in vitro antioxidant activity of various leaf extracts as well as anatomical investigation of Cajanus cajan were carried out. Physicochemical parameters such as total, acid-insoluble and water-soluble ash values and moisture content of the leaf powder of C. cajan were found to be 9.50%, 1.40 g/100 g, 4.15 g/100 g drug and 6.72%, respectively. Percent yield of acetone, aqueous, ethanol, ethyl acetate and chloroform leaf extracts were 9.0, 10.6, 13.75, 8.7 and 5.8 g/100 g, respectively. Significant amount of phenolic and flavonoid content were observed. The results of the antioxidant activity were found to be concentration-dependent. The IC50 values for DPPH assay determined for aqueous and ethanol extracts were 0.69 and 0.79 mg/ml, respectively. Reducing power is increased with increasing amount of concentration in both aqueous and ethanol leaf extracts. The highest hydroxyl radical scavenging activity reached up to 83.67% in aqueous and 78.75% in ethanol extracts and in phosphomolybdenum assay the aqueous extract showed strong antioxidant capacity up to 55.97 nM gallic acid equivalents/g. It was found that the aqueous extract possessed highest antioxidant activity in all the assays tested. The antioxidant characteristics of leaf extracts are possibly because of the presence of polyphenols. Microscopic study showed the presence of collenchyma, fibres, xylem, phloem, epidermis, trichomes, palisade tissue, basal sheath, pith and cortex in leaf, petiole and pulvinus. PMID:26009649

  8. Gibberellins and Gravitropism in Maize Shoots 1

    PubMed Central

    Rood, Stewart B.; Kaufman, Peter B.; Abe, Hiroshi; Pharis, Richard P.

    1987-01-01

    [3H]Gibberellin A20 (GA20) of high specific radioactivity (49.9 gigabecquerel per millimole) was applied equilaterally in a ring of microdrops to the internodal pulvinus of shoots of 3-week-old gravistimulated and vertical normal maize (Zea mays L.), and to a pleiogravitropic (prostrate) maize mutant, lazy (la). All plants converted the [3H]GA20 to [3H]GA1− and [3H]GA29-like metabolites as well as to several metabolites with the partitioning and chromatographic behavior of glucosyl conjugates of [3H]GA1, [3H]GA29, and [3H]GA8. The tentative identification of these putative [3H]GA glucosyl conjugates was further supported by the release of the free [3H]GA moiety after cleavage with cellulase. Within 12 hours of the [3H]GA20 feed, there was a significantly higher proportion of total radioactivity in lower than in upper halves of internode and leaf sheath pulvini in gravistimulated normal maize. Further, there was a significantly higher proportion of putative free GA metabolites of [3H]GA20, especially [3H]GA1, in the lower halves of normal maize relative to upper halves. The differential localization of the metabolites between upper and lower halves was not apparent in the pleiogravitropic mutant, la. Endogenous GA-like substances were also examined in gravistimulated maize shoots. Forty-eight hours after gravistimulation of 3-week-old maize seedlings, endogenous free GA-like substances in upper and lower leaf sheath and internode pulvini halves were extracted, chromatographed, and bioassayed using the `Tanginbozu' dwarf rice microdrop assay. Lower halves contained consistently higher total levels of GA-like activity. The qualitative elution profile of GA-like substances differed consistently, upper halves containing principally a GA20-like substance and lower halves containing mainly GA1-like and GA19-like substances. Gibberellins A1 (10 nanograms per gram) and A20 (5 nanograms per gram) were identified from these lower leaf sheath pulvini by capillary gas chromatography-selected ion monitoring. Results from all of these experiments are consistent with a role for GAs in the differential shoot growth that follows gravitropism, although the results do not eliminate the possibility that the redistribution of GAs results from the gravitropic response. Images Fig. 1 PMID:11539033

  9. Leaf movements and their relationship with the lunisolar gravitational force

    PubMed Central

    Barlow, Peter W.

    2015-01-01

    Background Observation of the diurnal ascent and descent of leaves of beans and other species, as well as experimental interventions into these movements, such as exposures to light at different times during the movement cycle, led to the concept of an endogenous ‘clock’ as a regulator of these oscillations. The physiological basis of leaf movement can be traced to processes that modulate cell volume in target tissues of the pulvinus and petiole. However, these elements of the leaf-movement process do not completely account for the rhythms that are generated following germination in constant light or dark conditions, or when plants are transferred to similar free-running conditions. Scope To develop a new perspective on the regulation of leaf-movement rhythms, many of the published time courses of leaf movements that provided evidence for the concept of the endogenous clock were analysed in conjunction with the contemporaneous time courses of the lunisolar tidal acceleration at the relevant experimental locations. This was made possible by application of the Etide program, which estimates, with high temporal resolution, local gravitational changes as a consequence of the diurnal variations of the lunisolar gravitational force due to the orbits and relative positions of Earth, Moon and Sun. In all cases, it was evident that a synchronism exists between the times of the turning points of both the lunisolar tide and of the leaftide when the direction of leaf movement changes. This finding of synchrony leads to the hypothesis that the lunisolar tide is a regulator of the leaftide, and that the rhythm of leaf movement is not necessarily of endogenous origin but is an expression of an exogenous lunisolar ‘clock’ impressed upon the leaf-movement apparatus. Conclusions Correlation between leaftide and Etide time courses holds for leaf movement rhythms in natural conditions of the greenhouse, in conditions of constant light or dark, under microgravity conditions of the International Space Station, and also holds for rhythms that are atypical, such as pendulum and relaxation rhythms whose periods are longer or shorter than usual. Even the apparently spontaneous short-period, small-amplitude rhythms recorded from leaves under unusual growth conditions are consistent with the hypothesis of a lunisolar zeitgeber. Two hypotheses that could account for the synchronism between leaftide and Etide, and which are based on either quantum considerations or on classical Newtonian physics, are presented and discussed. PMID:26205177

  10. Gibberellins and gravitropism in maize shoots: endogenous gibberellin-like substances and movement and metabolism of [3H]Gibberellin A20

    NASA Technical Reports Server (NTRS)

    Rood, S. B.; Kaufman, P. B.; Abe, H.; Pharis, R. P.

    1987-01-01

    [3H]Gibberellin A20 (GA20) of high specific radioactivity (49.9 gigabecquerel per millimole) was applied equilaterally in a ring of microdrops to the internodal pulvinus of shoots of 3-week-old gravistimulated and vertical normal maize (Zea mays L.), and to a pleiogravitropic (prostrate) maize mutant, lazy (la). All plants converted the [3H]GA20 to [3H]GA1- and [3H]GA29-like metabolites as well as to several metabolites with the partitioning and chromatographic behavior of glucosyl conjugates of [3H]GA1, [3H]GA29, and [3H]GA8. The tentative identification of these putative [3H]GA glucosyl conjugates was further supported by the release of the free [3H]GA moiety after cleavage with cellulase. Within 12 hours of the [3H]GA20 feed, there was a significantly higher proportion of total radioactivity in lower than in upper halves of internode and leaf sheath pulvini in gravistimulated normal maize. Further, there was a significantly higher proportion of putative free GA metabolites of [3H]GA20, especially [3H]GA1, in the lower halves of normal maize relative to upper halves. The differential localization of the metabolites between upper and lower halves was not apparent in the pleiogravitropic mutant, la. Endogenous GA-like substances were also examined in gravistimulated maize shoots. Forty-eight hours after gravistimulation of 3-week-old maize seedlings, endogenous free GA-like substances in upper and lower leaf sheath and internode pulvini halves were extracted, chromatographed, and bioassayed using the "Tanginbozu" dwarf rice microdrop assay. Lower halves contained consistently higher total levels of GA-like activity. The qualitative elution profile of GA-like substances differed consistently, upper halves containing principally a GA20-like substance and lower halves containing principally a GA20-like substance and lower halves containing mainly GA1-like and GA19-like substances. Gibberellins A1 (10 nanograms per gram) and A20 (5 nanograms per gram) were identified from these lower leaf sheath pulvini by capillary gas chromatography-selected ion monitoring. Results from all of these experiments are consistent with a role for GAs in the differential shoot growth that follows gravitropism, although the results do not eliminate the possibility that the redistribution of GAs results from the gravitropic response.

  11. Leaf movements and their relationship with the lunisolar gravitational force.

    PubMed

    Barlow, Peter W

    2015-08-01

    Observation of the diurnal ascent and descent of leaves of beans and other species, as well as experimental interventions into these movements, such as exposures to light at different times during the movement cycle, led to the concept of an endogenous 'clock' as a regulator of these oscillations. The physiological basis of leaf movement can be traced to processes that modulate cell volume in target tissues of the pulvinus and petiole. However, these elements of the leaf-movement process do not completely account for the rhythms that are generated following germination in constant light or dark conditions, or when plants are transferred to similar free-running conditions. To develop a new perspective on the regulation of leaf-movement rhythms, many of the published time courses of leaf movements that provided evidence for the concept of the endogenous clock were analysed in conjunction with the contemporaneous time courses of the lunisolar tidal acceleration at the relevant experimental locations. This was made possible by application of the Etide program, which estimates, with high temporal resolution, local gravitational changes as a consequence of the diurnal variations of the lunisolar gravitational force due to the orbits and relative positions of Earth, Moon and Sun. In all cases, it was evident that a synchronism exists between the times of the turning points of both the lunisolar tide and of the leaftide when the direction of leaf movement changes. This finding of synchrony leads to the hypothesis that the lunisolar tide is a regulator of the leaftide, and that the rhythm of leaf movement is not necessarily of endogenous origin but is an expression of an exogenous lunisolar 'clock' impressed upon the leaf-movement apparatus. Correlation between leaftide and Etide time courses holds for leaf movement rhythms in natural conditions of the greenhouse, in conditions of constant light or dark, under microgravity conditions of the International Space Station, and also holds for rhythms that are atypical, such as pendulum and relaxation rhythms whose periods are longer or shorter than usual. Even the apparently spontaneous short-period, small-amplitude rhythms recorded from leaves under unusual growth conditions are consistent with the hypothesis of a lunisolar zeitgeber. Two hypotheses that could account for the synchronism between leaftide and Etide, and which are based on either quantum considerations or on classical Newtonian physics, are presented and discussed. © The Author 2015. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please email: journals.permissions@oup.com.

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