ABSTRACT
BACKGROUND AND AIMS: Understanding anatomical variations across plant phylogenies and environmental gradients is vital for comprehending plant evolution and adaptation. Previous studies on tropical woody plants have paid limited attention to quantitative differences in major xylem tissues, which serve specific roles in mechanical support (fibres), carbohydrate storage and radial conduction (radial parenchyma, rays), wood capacitance (axial parenchyma) and water transport (vessels). To address this gap, we investigate xylem fractions in 173 tropical tree species spanning 134 genera and 53 families along a 2200-m elevational gradient on Mount Cameroon, West Africa. METHODS: We determined how elevation, stem height and wood density affect interspecific differences in vessel, fibre, and specific axial (AP) and radial (RP) parenchyma fractions. We focus on quantifying distinct subcategories of homogeneous or heterogeneous rays and apotracheal, paratracheal and banded axial parenchyma. KEY RESULTS: Elevation-related cooling correlated with reduced AP fractions and vessel diameters, while fibre fractions increased. Lower elevations exhibited elevated AP fractions due to abundant paratracheal and wide-banded parenchyma in tall trees from coastal and lowland forests. Vasicentric and aliform AP were predominantly associated with greater tree height and wider vessels, which might help cope with high evaporative demands via elastic wood capacitance. In contrast, montane trees featured a higher fibre proportion, scarce axial parenchyma, smaller vessel diameters and higher vessel densities. The lack of AP in montane trees was often compensated for by extended uniseriate ray sections with upright or squared ray cells or the presence of living fibres. CONCLUSIONS: Elevation gradient influenced specific xylem fractions, with lower elevations showing elevated AP due to abundant paratracheal and wide-banded parenchyma, securing greater vessel-to-parenchyma connectivity and lower embolism risk. Montane trees featured a higher fibre proportion and smaller vessel diameters, which may aid survival under greater environmental seasonality and fire risk.
Subject(s)
Trees , Tropical Climate , Wood , Xylem , Wood/anatomy & histology , Wood/physiology , Trees/anatomy & histology , Trees/physiology , Xylem/anatomy & histology , Xylem/physiology , Cameroon , AltitudeABSTRACT
BACKGROUND AND AIMS: Understanding the mutual co-ordination of vegetative and reproductive growth is important in both agricultural and ecological settings. A competitive relationship between vegetative growth and fruiting is often highlighted, resulting in an apparent trade-off between structural growth and fruit production. However, our understanding of factors driving this relationship is limited. METHODS: We used four scions grafted onto a series of size-controlling rootstocks to evaluate the relationships between the annual fruit yield and radial growth of trunks, branches and roots. To assess tree radial growth, we measured ring widths on extracted tree cores, which is an approach not frequently used in a horticultural setting. KEY RESULTS: We found that the yield and radial growth were negatively related when plotted in absolute terms or as detrended and normalized indices. The relationship was stronger in low vigour trees, but only after the age-related trend was removed. In contrast, when trunk radial growth was expressed as basal area increment, the negative relationship disappeared, suggesting that the relationship between trunk radial growth and fruit yield might not be a true trade-off related to the competition between the two sinks. The effect of low yield was associated with increased secondary growth not only in trunks but also in branches and roots. In trunks, we observed that overcropping was associated with reduced secondary growth in a subsequent year, possibly due to the depletion of reserves. CONCLUSIONS: Our results show that variation in annual fruit yield due to tree ageing, weather cueing and inherent alternate bearing behaviour is reflected in the magnitude of secondary growth of fruit trees. We found little support for the competition/architecture theory of rootstock-induced growth vigour control. More broadly, our study aimed at bridging the gap between forest ecology and horticulture.
Subject(s)
Malus , Pyrus , Fruit , Plant Roots , TreesABSTRACT
Bending and torsional properties of young roots and stems were measured in nine woody angiosperms. The variation in mechanical parameters was correlated to wood anatomical traits and analysed with respect to the other two competing functions of xylem (namely storage and hydraulics). Compared with stems, roots exhibited five times greater flexibility in bending and two times greater flexibility in torsion. Lower values of structural bending and structural torsional moduli (Estr and Gstr, respectively) of roots compared with stems were associated with the presence of thicker bark and a greater size of xylem cells. Across species, Estr and Gstr were correlated with wood density, which was mainly driven by the wall thickness to lumen area ratio of fibres. Higher fractions of parenchyma did not translate directly into a lower wood density and reduced mechanical stiffness in spite of parenchyma cells having thinner, and in some cases less lignified, cell walls than fibres. The presence of wide, partially non-lignified rays contributed to low values of Estr and Gstr in Clematis vitalba. Overall, our results demonstrate that higher demands for mechanical stability in self-supporting stems put a major constraint on xylem structure, whereas root xylem can be designed with a greater emphasis on both storage and hydraulic functions.
Subject(s)
Magnoliopsida/chemistry , Plant Roots/chemistry , Plant Stems/chemistry , Xylem/chemistry , Magnoliopsida/growth & development , Plant Roots/growth & development , Plant Stems/growth & development , Wood/chemistry , Wood/growth & developmentABSTRACT
Parenchyma represents a critically important living tissue in the sapwood of the secondary xylem of woody angiosperms. Considering various interactions between parenchyma and water transporting vessels, we hypothesize a structure-function relationship between both cell types. Through a generalized additive mixed model approach based on 2,332 woody angiosperm species derived from the literature, we explored the relationship between the proportion and spatial distribution of ray and axial parenchyma and vessel size, while controlling for maximum plant height and a range of climatic factors. When factoring in maximum plant height, we found that with increasing mean annual temperatures, mean vessel diameter showed a positive correlation with axial parenchyma proportion and arrangement, but not for ray parenchyma. Species with a high axial parenchyma tissue fraction tend to have wide vessels, with most of the parenchyma packed around vessels, whereas species with small diameter vessels show a reduced amount of axial parenchyma that is not directly connected to vessels. This finding provides evidence for independent functions of axial parenchyma and ray parenchyma in large vesselled species and further supports a strong role for axial parenchyma in long-distance xylem water transport.
Subject(s)
Magnoliopsida/anatomy & histology , Wood/anatomy & histology , Xylem/anatomy & histology , Climate , Models, Theoretical , Rain , TemperatureABSTRACT
BACKGROUND: Vessel-associated cells (VACs) are highly specialized, living parenchyma cells that are in direct contact with water-conducting, dead vessels. The contact may be sparse or in large tight groups of parenchyma that completely surrounds vessels. VACs differ from vessel distant parenchyma in physiology, anatomy, and function and have half-bordered pits at the vessel-parenchyma juncture. The distinct anatomy of VACs is related to the exchange of substances to and from the water-transport system, with the cells long thought to be involved in water transport in woody angiosperms, but where direct experimental evidence is lacking. SCOPE: This review focuses on our current knowledge of VACs regarding anatomy and function, including hydraulic capacitance, storage of nonstructural carbohydrates, symplastic and apoplastic interactions, defense against pathogens and frost, osmoregulation, and the novel hypothesis of surfactant production. Based on microscopy, we visually represent how VACs vary in dimensions and general appearance between species, with special attention to the protoplast, amorphous layer, and the vessel-parenchyma pit membrane. CONCLUSIONS: An understanding of the relationship between VACs and vessels is crucial to tackling questions related to how water is transported over long distances in xylem, as well as defense against pathogens. New avenues of research show how parenchyma-vessel contact is related to vessel diameter and a new hypothesis may explain how surfactants arising from VAC can allow water to travel under negative pressure. We also reinforce the message of connectivity between VAC and other cells between xylem and phloem.
Subject(s)
Magnoliopsida/cytology , Xylem/cytology , Magnoliopsida/physiology , Magnoliopsida/ultrastructure , Water/metabolism , Xylem/physiology , Xylem/ultrastructureABSTRACT
Variation in xylem vessel diameter is one of the most important parameters when evaluating plant water relations. This review provides a synthesis of the ecophysiological implications of variation in lumen diameter together with a summary of our current understanding of vessel development and its endogenous regulation. We analyzed inter-specific variation of the mean hydraulic vessel diameter (Dv ) across biomes, intra-specific variation of Dv under natural and controlled conditions, and intra-plant variation. We found that the Dv measured in young branches tends to stay below 30 µm in regions experiencing winter frost, whereas it is highly variable in the tropical rainforest. Within a plant, the widest vessels are often found in the trunk and in large roots; smaller diameters have been reported for leaves and small lateral roots. Dv varies in response to environmental factors and is not only a function of plant size. Despite the wealth of data on vessel diameter variation, the regulation of diameter is poorly understood. Polar auxin transport through the vascular cambium is a key regulator linking foliar and xylem development. Limited evidence suggests that auxin transport is also a determinant of vessel diameter. The role of auxin in cell expansion and in establishing longitudinal continuity during secondary growth deserve further study.
Subject(s)
Xylem/anatomy & histology , Xylem/physiology , Indoleacetic Acids/metabolism , Magnoliopsida/anatomy & histology , Magnoliopsida/physiology , Plant Development , Species SpecificityABSTRACT
Parenchyma is an important tissue in secondary xylem of seed plants, with functions ranging from storage to defence and with effects on the physical and mechanical properties of wood. Currently, we lack a large-scale quantitative analysis of ray parenchyma (RP) and axial parenchyma (AP) tissue fractions. Here, we use data from the literature on AP and RP fractions to investigate the potential relationships of climate and growth form with total ray and axial parenchyma fractions (RAP). We found a 29-fold variation in RAP fraction, which was more strongly related to temperature than with precipitation. Stem succulents had the highest RAP values (mean ± SD: 70.2 ± 22.0%), followed by lianas (50.1 ± 16.3%), angiosperm trees and shrubs (26.3 ± 12.4%), and conifers (7.6 ± 2.6%). Differences in RAP fraction between temperate and tropical angiosperm trees (21.1 ± 7.9% vs 36.2 ± 13.4%, respectively) are due to differences in the AP fraction, which is typically three times higher in tropical than in temperate trees, but not in RP fraction. Our results illustrate that both temperature and growth form are important drivers of RAP fractions. These findings should help pave the way to better understand the various functions of RAP in plants.
Subject(s)
Mesophyll Cells/physiology , Seeds/physiology , Trees/physiology , Xylem/physiology , Climate , Databases as Topic , Magnoliopsida/growth & development , Magnoliopsida/physiology , Mesophyll Cells/cytology , Rain , Statistics, Nonparametric , Temperature , Tracheophyta/growth & development , Tracheophyta/physiology , Wood/physiology , Xylem/cytologyABSTRACT
The evolution of lignified xylem allowed for the efficient transport of water under tension, but also exposed the vascular network to the risk of gas emboli and the spread of gas between xylem conduits, thus impeding sap transport to the leaves. A well-known hypothesis proposes that the safety of xylem (its ability to resist embolism formation and spread) should trade off against xylem efficiency (its capacity to transport water). We tested this safety-efficiency hypothesis in branch xylem across 335 angiosperm and 89 gymnosperm species. Safety was considered at three levels: the xylem water potentials where 12%, 50% and 88% of maximal conductivity are lost. Although correlations between safety and efficiency were weak (r(2) < 0.086), no species had high efficiency and high safety, supporting the idea for a safety-efficiency tradeoff. However, many species had low efficiency and low safety. Species with low efficiency and low safety were weakly associated (r(2) < 0.02 in most cases) with higher wood density, lower leaf- to sapwood-area and shorter stature. There appears to be no persuasive explanation for the considerable number of species with both low efficiency and low safety. These species represent a real challenge for understanding the evolution of xylem.
Subject(s)
Cycadopsida/physiology , Magnoliopsida/physiology , Xylem/physiology , Plant Leaves/physiology , Plant Transpiration , Water/physiology , WoodABSTRACT
PREMISE OF THE STUDY: Concentrations of nonstructural carbohydrates (NSCs) are used as proxies for the net carbon balance of trees and as indicators of carbon starvation resulting from environmental stress. Woody organs are the largest NSC-storing compartments in forest ecosystems; therefore, it is essential to understand the factors that affect the size of this important storage pool. In wood, NSC are predominantly deposited in ray and axial parenchyma (RAP); however, direct links between nutrient storage and RAP anatomy have not yet been established. Here, we tested whether the NSC storage capacity of wood is influenced by the amount of RAP. METHODS: We measured NSC concentrations and RAP fractions in root and stem sapwood of 12 temperate species sampled at the onset of winter dormancy and in stem sapwood of four tropical trees growing in an evergreen lowland rainforest. The patterns of starch distribution were visualized by staining with Lugol's solution. KEY RESULTS: The concentration of NSCs in sapwood of temperate trees scales tightly with the amount of RAP and living fibers (LFs), with almost all RAP and LFs being densely packed with starch grains. In contrast, the tropical species had lower NSC concentrations despite their higher RAP and LFs fraction and had considerable interspecific differences in starch distribution. CONCLUSIONS: The differences in RAP and LFs abundance affect the ability of sapwood to store NSC in temperate trees, whereas a more diverse set of functions of RAP might be pronounced in species growing in a tropical environment with little seasonality.
Subject(s)
Carbohydrates/analysis , Plant Roots/anatomy & histology , Plant Roots/chemistry , Plant Stems/anatomy & histology , Plant Stems/chemistry , Trees/anatomy & histology , Trees/chemistry , Species Specificity , Starch/metabolism , Tropical ClimateABSTRACT
Cellulose delta18O and deltaD can provide insights on climates and hydrological cycling in the distant past and how these factors differ spatially. However, most studies of plant cellulose have used only one isotope, most commonly delta18O, resulting in difficulties partitioning variation in delta18O of precipitation vs. evaporative conditions that affect leaf water isotopic enrichment. Moreover, observations of pronounced diurnal differences from conventional steady-state model predictions of leaf water isotopic fractionation have cast some doubt on single isotope modeling approaches for separating precipitation and evaporation drivers of cellulose delta18O or deltaD. We explore a dual isotope approach akin to the concept of deuterium-excess (d), to establish deuterium deviations from the global meteoric water line in leaf water (deltad(l)) as driven by relative humidity (RH). To demonstrate this concept, we survey studies of leaf water delta18O and deltaD in hardwood vs. conifer trees. We then apply the concept to cellulose delta18O and deltaD using a mechanistic model of cellulose delta18O and deltaD to reconstruct deuterium deviations from the global meteoric water line (deltad(c)) in Quercus macrocarpa, Q. robur, and Pseudotsuga menziesii. For each species, deltad(c) showed strong correlations with RH across sites. deltad(c) agreed well with steady-state predictions for Q. macrocarpa, while for Q. robur, the relationship with RH was steeper than expected. The slope of deltad(c) vs. RH of P. menziesii was also close to steady-state predictions, but deltad(c) were more enriched than predicted. This is in agreement with our leaf water survey showing conifer deltad(l) was more enriched than predicted. Our data reveal that applications of this method should be appropriate for reconstructing RH from cellulose delta18O and deltaD after accounting for differences between hardwoods and conifers. Hence, deltad(c) should be useful for understanding variability in RH associated with past climatic cycles, across regional climates, or across complex terrain where climate modeling is challenging. Furthermore, deltad(c) and inferred RH values should help in constraining variation in source water delta18O.
Subject(s)
Deuterium , Humidity , Climate , Oxygen Isotopes , Plant Leaves , Trees , WaterABSTRACT
Nitrogen availability has a strong influence on plant growth and development. In this study, we examined the effect of nitrogen availability on xylogenesis in hybrid poplar (Populus trichocarpa x deltoides H11-11). Saplings of hybrid poplar were fertilized for 33 d with either high or adequate levels of ammonium nitrate. We observed enhanced radial growth, wider vessels and fibres and thinner fibre walls in the secondary xylem of high N relative to adequate N plants. These anatomical differences translated into altered hydraulic properties with xylem being more transport efficient but also more vulnerable to drought-induced cavitation in high N plants. The changes in xylem structure and function were associated with differences in gene expression as revealed by the transcriptome analysis of the developing xylem region. We found 388 genes differentially expressed (fold change ±1.5, P-value ≤ 0.05), including a number of genes putatively involved in nitrogen and carbohydrate metabolism and various aspects of xylem cell differentiation. Several genes encoding known transcriptional regulators of secondary cell wall deposition were down-regulated in high N plants, corresponding with thinner secondary cell walls in these plants. The results of this study provide us with gene candidates potentially affecting xylem hydraulic and structural traits.
Subject(s)
Nitrogen/administration & dosage , Populus/drug effects , Xylem/drug effects , Carbohydrate Metabolism/drug effects , Cell Division , Fertilizers , Gene Expression/drug effects , Genes, Plant , Lignin/metabolism , Nitrates , Nitrogen/metabolism , Plant Proteins/metabolism , Populus/genetics , Populus/metabolism , Transcription Factors/metabolismABSTRACT
Variation in xylem structure and function has been extensively studied across different species with a wide taxonomic, geographical, and ecological coverage. In contrast, our understanding of how xylem of a single species can adjust to different growing condition remains limited. Here phenotypic and developmental plasticity in xylem traits of hybrid poplar (Populus trichocarpa×deltoides) was studied. Clonally propagated saplings were grown under experimental drought, nitrogen fertilization, and shade for >30 d. Xylem hydraulic and anatomical traits were subsequently examined in stem segments taken from two different vertical positions along the plant's main axis. The experimental treatments affected growth and development and induced changes in xylem phenotype. Across all treatments, the amount of leaf area supported by stem segments (A(L)) scaled linearly with stem native hydraulic conductivity (K (native)), suggesting that the area of assimilating leaves is constrained by the xylem transport capacity. In turn, K (native) was mainly driven by the size of xylem cross-sectional area (A(X)). Moreover, the structural and functional properties of xylem varied significantly. Vulnerability to cavitation, measured as the xylem pressure inducing 50% loss of conductivity (P50), ranged from -1.71 MPa to -0.15 MPa in saplings subjected to drought and nitrogen fertilization, respectively. Across all treatments and stem segment positions, P50 was tightly correlated with wood density. In contrast, no relationship between P50 and xylem-specific conductivity (K (S)) was observed. The results of this study enhance our knowledge of plant hydraulic acclimation and provide insights into common trade-offs that exist in xylem structure and function.
Subject(s)
Plant Transpiration , Populus/physiology , Xylem/anatomy & histology , Acclimatization , Droughts , Nitrogen/metabolism , Phenotype , Plant Leaves/anatomy & histology , Plant Leaves/physiology , Plant Stems/anatomy & histology , Plant Stems/genetics , Plant Stems/metabolism , Populus/genetics , Populus/growth & development , Sunlight , Water/metabolism , Xylem/genetics , Xylem/metabolismABSTRACT
The use of size-controlling rootstocks is central to modern high-density fruit production systems. While biological mechanisms responsible for vigor control are not fully understood, differences in water relations and carbohydrate storage ability have been suggested as two potential factors. To better understand the processes that control growth vigor, we analyzed the trunk radial variation at seasonal and diurnal timescales and measured the midday leaf water potential (ΨMD), leaf gas exchange and concentrations of non-structural carbohydrates (NSC) in apple trees of variety 'Jonagold' grafted on two rootstocks of contrasting growth vigor (dwarfing J-TE-G vs invigorating J-TE-H). The measurements were conducted during an exceptionally hot and dry summer. We found that smaller annual trunk radial increments in dwarfed trees were primarily due to an earlier cessation of trunk secondary growth. The interdiurnal trunk circumference changes (ΔC) were slightly lower in dwarfed trees, and these trees also had fewer days with positive ΔC values, particularly during the driest summer months. The trunks of dwarfed trees shrank gradually during the drought, showed less pronounced diurnal variation of trunk circumference and the maximum trunk daily shrinkage was only weakly responsive to the vapor pressure deficit. These results indicated that lower turgidity in the cambial region may have limited the trunk radial expansion in dwarfed trees during the hot and dry days. Dwarfed trees also maintained lower ΨMD and leaf gas exchange rates during the summer drought. These parameters decreased in parallel for both rootstock combinations, suggesting their similar drought sensitivity. Similar concentrations and seasonal dynamics of NSC in both rootstock combinations, together with their similar spring growth rates, suggest that NSC reserves were not directly limiting for growth. Our results support the prominent role of water relations in rootstock-induced size-controlling mechanisms and highlight the complexity of this topic.
Subject(s)
Malus , Carbon , Droughts , Plant Roots , Seasons , Trees , WaterABSTRACT
Intervessel pits act as safety valves that prevent the spread of xylem embolism. Pectin-calcium crosslinks within the pit membrane have been proposed to affect xylem vulnerability to cavitation. However, as the chemical composition of pit membranes is poorly understood, this hypothesis has not been verified. Using electron microscopy, immunolabeling, an antimonate precipitation technique, and ruthenium red staining, we studied the distribution of selected polysaccharides and calcium in the pit membranes of four angiosperm tree species. We tested whether shifts in xylem vulnerability resulting from perfusion of stems with a calcium chelating agent corresponded with the distribution of pectic homogalacturonans (HG) and/or calcium within interconduit pit membranes. No HG were detected in the main part of intervessel pit membranes, but were consistently found in the marginal membrane region known as the annulus. Calcium colocalized with HG in the annulus. In contrast to intervessel pits, the membrane of vessel-ray pits showed a high pectin content. The presence of two distinct chemical domains, the annulus and the actual pit membrane, can have substantial implications for pit membrane functioning. We propose that the annulus could affect the observed shift in xylem vulnerability after calcium removal by allowing increased pit membrane deflection.
Subject(s)
Calcium/metabolism , Epitopes/immunology , Magnoliopsida/immunology , Pectins/immunology , Xylem/immunology , Antibody Specificity/immunology , Esterification , Glucans/immunology , Magnoliopsida/metabolism , Magnoliopsida/ultrastructure , Methylation , Ruthenium Red/metabolism , Species Specificity , Staining and Labeling , Xylans/immunology , Xylem/metabolism , Xylem/ultrastructureABSTRACT
The effect of shading on xylem hydraulic traits and xylem anatomy was studied in hybrid poplar (Populus trichocarpa x deltoides, clone H11-11). Hydraulic measurements conducted on stem segments of 3-month-old saplings grown in shaded (SH) or control light (C) conditions indicated that shading resulted in more vulnerable and less efficient xylem. Air is thought to enter vessels through pores in inter-vessel pit membranes, thereby nucleating cavitation. Therefore, we tested if the ultrastructure and/or chemistry of pit membranes differed in SH and C plants. Transmission electron micrographs revealed that pit membranes were thinner in SH, which was paralleled by lower compound middle lamella thickness. Immunolabelling with JIM5 and JIM7 monoclonal antibodies surprisingly indicated that pectic homogalacturonans were not present in the mature pit membrane regardless of the light treatment. Porosity measurements conducted with scanning electron microscopy were significantly affected by the method used for sample dehydration. Drying through a gradual ethanol series seems to be a better alternative to drying directly from a hydrated state for pit membrane observations in poplar. Scanning electron microscopy based estimates of pit membrane porosity probably overestimated real porosity as suggested by the results from the 'rare pit' model.
Subject(s)
Cell Membrane/ultrastructure , Light , Xylem/radiation effects , Cell Wall/ultrastructure , Microscopy, Electron, Scanning , Microscopy, Electron, Transmission , Pectins/analysis , Plant Leaves/cytology , Plant Leaves/physiology , Plant Leaves/radiation effects , Populus/cytology , Populus/physiology , Populus/radiation effects , Porosity , Xylem/cytology , Xylem/physiologyABSTRACT
Hemiparasitic plants gain virtually all mineral nutrients and water from their host plant whilst organic carbon is provided, at least in part, by their own photosynthetic activity, although their rates of assimilation are substantially lower than that found in non-parasitic plants. Hence, hemiparasites must gain at least some of their organic carbon heterotrophically from the host plant. Despite this, heterotrophic carbon gain by root hemiparasites has been investigated only for a few genera. We investigated heterotrophic carbon gain by two root hemiparasites, Rhinanthus minor L. and Euphrasia rostkoviana Hayne (Orobanchaceae), using natural abundance stable isotope (delta(13)C) profiles of both parasites attached to C(3) (wheat) and C(4) (maize) hosts coupled to a linear two-source isotope-mixing model to estimate the percentage of carbon in the parasite that was derived from the host. Both R. minor and E. rostkoviana attached to maize hosts were significantly more enriched in (13)C than those attached to wheat hosts with R. minor becoming more enriched in (13)C than E. rostkoviana. The natural abundance (13)C profiles of both parasites were not significantly different from their wheat hosts, but were less enriched in (13)C than maize hosts. Using a linear two-source isotope-mixing model, we estimated that R. minor and E. rostkoviana adult plants derive c. 50 and 25% of their carbon from their hosts, respectively. In light of these results, we hypothesise that repeatedly observed negative effect of competition for light on hemiparasites acts predominantly in early ontogenetic stages when parasites grow unattached or the abstraction of host nutrients is less effective.
Subject(s)
Carbon/metabolism , Heterotrophic Processes/physiology , Orobanchaceae/physiology , Plant Roots/parasitology , Triticum/parasitology , Zea mays/parasitology , Biomass , Carbon Isotopes , Euphrasia/physiology , Host-Parasite Interactions , Photosynthesis , Plantago/growth & development , Plantago/parasitology , Triticum/growth & development , Zea mays/growth & developmentABSTRACT
We studied the influence of nitrogen (N) on hydraulic traits and aquaporin (AQP) expression in the stem xylem of hybrid poplar saplings (Populus trichocarpa (Torr. & Gray) x deltoides Bartr. ex Marsh clone H11-11). Plants were grown in a controlled environment and were kept well watered throughout the experiments. Hydraulic measurements were done on basal and distal stem segments of plants receiving high N fertilization (high N plants) versus plants receiving only adequate N fertilization (adequate N plants). High N plants grew faster and exhibited more leaf area than adequate N controls. These morphological differences were paralleled by wider vessels and higher specific conductivities (K(S)) in high N plants. However, stems of high N plants were more vulnerable to xylem cavitation, at least in one of two experiments, and showed lower wood densities than stems of adequate N plants. Leaf area was strongly correlated with cross-sectional xylem area in both plant groups. Since higher K(S) in high N plants was accompanied by concomitant increases in leaf area, leaf-specific conductivities were similar in both plant groups. Influences of N on hydraulic traits were paralleled by changes in AQP expression. Seven AQPs were upregulated in the stem xylem of high N plants, five of which have been identified recently as water transporters. The enhanced growth of secondary xylem of high N plants has been shown to result from both increased cambial activity as well as increased cell size. We suggest that some of these water-transporting AQPs could play a role in xylogenesis, facilitating the influx of water into the zone of differentiating and maturing cells in secondary xylem, including expanding vessels.
Subject(s)
Aquaporins/metabolism , Gene Expression Regulation, Plant/drug effects , Nitrogen/pharmacology , Populus/drug effects , Populus/metabolism , Xylem/physiology , Aquaporins/genetics , Populus/genetics , Water/metabolism , Xylem/drug effectsABSTRACT
The study aims to assess variability in leaf water isotopic enrichment occurring in the field under natural conditions. We focused on seasonal variation and difference between sun-exposed and shaded leaves. Isotopic composition (δ18O, δ2H) of leaf water was monitored in a beech tree (Fagus sylvatica L.) growing in the forest-meadow ecotone together with δ18O (2H) of water compartments which are in close relation to this signal, namely twig and soil water. The sampling was carried out in approximately two-week intervals during five consecutive vegetation seasons. The δ18O (2H) data showed a distinct seasonal pattern and a consistency in relative differences between the seasons and sample categories. Leaf water was the most isotopically enriched water compartment. The leaf water enrichment decreased toward the autumn reflecting the change in δ18O (2H) of source water and evaporative demands. The soil and twig water isotopic signal was depleted against current precipitation as it partly retained the isotopic signature from winter precipitation however the seasonal pattern of soil and twig water followed that of precipitation. No significant differences between sun-exposed and shaded samples were detected. Nevertheless, the observed strong seasonal pattern of isotope composition of leaf, twig and soil water should be taken into account when using leaf water enrichment for further calculations or modeling.
Subject(s)
Deuterium/analysis , Fagus/metabolism , Oxygen Isotopes/analysis , Plant Leaves/chemistry , Water/metabolism , Deuterium/metabolism , Fagus/chemistry , Fagus/physiology , Oxygen Isotopes/metabolism , Plant Leaves/metabolism , Seasons , SunlightABSTRACT
The release of water from storage compartments to the transpiration stream is an important functional mechanism that provides the buffering of sudden fluctuations in water potential. The ability of tissues to release water per change in water potential, referred to as hydraulic capacitance, is assumed to be associated with the anatomy of storage tissues. However, information about how specific anatomical parameters determine capacitance is limited. In this study, we measured sapwood capacitance (C) in terminal branches and roots of five temperate tree species (Fagus sylvatica L., Picea abies L., Quercus robur L., Robinia pseudoacacia L., Tilia cordata Mill.). Capacitance was calculated separately for water released mainly from capillary (CI; open vessels, tracheids, fibres, intercellular spaces and cracks) and elastic storage compartments (CII; living parenchyma cells), corresponding to two distinct phases of the moisture release curve. We found that C was generally higher in roots than branches, with CI being 3-11 times higher than CII Sapwood density and the ratio of dead to living xylem cells were most closely correlated with C In addition, the magnitude of CI was strongly correlated with fibre/tracheid lumen area, whereas CII was highly dependent on the thickness of axial parenchyma cell walls. Our results indicate that water released from capillary compartments predominates over water released from elastic storage in both branches and roots, suggesting the limited importance of parenchyma cells for water storage in juvenile xylem of temperate tree species. Contrary to intact organs, water released from open conduits in our small wood samples significantly increased CI at relatively high water potentials. Linking anatomical parameters with the hydraulic capacitance of a tissue contributes to a better understanding of water release mechanisms and their implications for plant hydraulics.