Xylem hydraulic characteristics of subtropical trees from contrasting habitats grown under identical environmental conditions

Five evergreen subtropical tree species growing under identical environmental conditions were investigated to establish which hydraulic properties are genotypically rigid and which show phenotypic plasticity. Maximum xylem-specific conductivity ( k _s) correlated well with the anatomical characteristics (conduit diameter and density) for the four angiosperms Tecomaria capensis , Trichilia dregeana , Cinnamomum camphora and Barringtonia racemosa ; the anatomy of the gymnosperm Podocarpus latifolius was not assessed. Huber values (functional xylem cross-sectional area : leaf area) varied inversely with k _s among species. Maximum leaf-specific conductivity was similar in the five unrelated species. Vulnerability of xylem to cavitation differed between species, as did the relationship between transpiration and water potential. Models of these parameters and isolated midday readings confirm that these trees operate at similar maximum leaf-specific conductivity ( k _l) values. The data are consistent with the hypothesis that conductivity characteristics ( k _l, k _s) are influenced by environment, whereas vulnerability to cavitation is genetically determined.     

The hydraulic architecture of Juniperus communis L. ssp. communis: shrubs and trees compared

Juniperus communis ssp. communis can grow like a shrub or it can develop a tree-like habit. In this study, the hydraulic architecture of these contrasting growth forms was compared. We analysed the hydraulic efficiency (leaf-specific conductivity, k(l); specific conductivity, k(s); Huber value, HV) and the vulnerability to cavitation (the water potential corresponding to a 50% loss of conductivity, Psi(50)), as well as anatomical parameters [mean tracheid diameter, d; mean hydraulic diameter, d(h); cell wall reinforcement (t/b)(h)(2)] of shrub shoots, tree stems and tree branches. Shrub shoots were similar to tree branches (especially to lower branches) in growth form and conductivity (k(l) = 1.93 +/- 0.11 m(2) s(-1) MPa(-1) 10(-7), k(s) = 5.71 +/- 0.19 m(2) s(-1) MPa(-1) 10(-4)), but were similar to tree stems in their vulnerability to cavitation (Psi(50) = -5.81 +/- 0.08 MPa). Tree stems showed extraordinarily high k(l) and k(s) values, and HV increased from the base up. Stem xylem was more vulnerable to cavitation than branch xylem, where Psi(50) increased from lower (Psi(50) = -6.44 +/- 0.19 MPa) to upper branches (Psi(50) = -5.98 +/- 0.13 MPa). Conduit diameters were correlated with k(l) and k(s). Data indicate that differences in hydraulic architecture correspond to changes in growth form. In some aspects, the xylem hydraulics of tree-like Juniperus communis differs from that of other coniferous tree species.     

Drought resistance of Quercus pubescens as a function of root hydraulic conductance, xylem embolism and hydraulic architecture

Water relations, xylem embolism, root and shoot hydraulic conductance of both young plants in the field and potted seedlings of Quercus pubescens have been studied with the aim of investigating whether these variables may account for the well known adaptation of this oak species to arid habitats. Our data revealed that Q. pubescens is able to maintain high leaf relative water contents under water stress conditions. In fact, relative water contents measured in summer (July) did not differ from those recorded in April. This was apparently achieved by compensating water loss by an equal amount of water uptake. Such a drought avoidance strategy was made possible by the recorded high hydraulic efficiency of stems and roots under water stress. In fact, root hydraulic conductance of field-grown plants was maintained high in summer when the percentage loss of hydraulic conductance of stems was lowest. The hydraulic architecture of young plants of Q. pubescens measured in terms of partitioning of hydraulic resistances along the water pathway revealed that the highest hydraulic resistance was located in stems of the current year's growth. This hydraulic architecture is interpreted as consistent with the adaptation of Q. pubescens to arid habitats as a consequence of the recorded seasonal changes in water relation parameters as well as in root and stem hydraulics.     

Climate-related trends in sapwood biophysical properties in two conifers: avoidance of hydraulic dysfunction through coordinated adjustments in xylem efficiency, safety and capacitance

In the Pacific north‐west, the Cascade Mountain Range blocks much of the precipitation and maritime influence of the Pacific Ocean, resulting in distinct climates east and west of the mountains. The current study aimed to investigate relationships between water storage and transport properties in populations of Douglas‐fir (Pseudotsuga menziesii) and ponderosa pine (Pinus ponderosa) adapted to both climates. Sapwood thickness, capacitance, vulnerability to embolism, and axial and radial conductivity were measured on samples collected from trunks of mature trees. The sapwood of ponderosa pine was three to four times thicker than Douglas‐fir. Radial conductivity was higher in west‐side populations of both species, but axial conductivity was higher in the east‐side populations and in Douglas‐fir. Eastern populations of both species had sapwood that was more vulnerable to embolism than west‐side populations. Sapwood capacitance was similar between species, but was about twice as great in east‐side populations (580 kg m^?3 MPa^?1) as in west‐side populations (274 kg m^?3 MPa^?1). Capacitance was positively correlated with both mean embolism pressure and axial conductivity across species and populations, suggesting that coordinated adjustments in xylem efficiency, safety and water storage capacity may serve to avoid embolism along a gradient of increasing aridity.     

Influence of nutrient versus water supply on hydraulic architecture and water balance in Pinus taeda

We investigated the hydraulic consequences of a major decrease in root‐to‐leaf area ratio (A_R:A_L) caused by nutrient amendments to 15‐year‐old Pinus taeda L. stands on sandy soil. In theory, such a reduction in A_R:A_L should compromise the trees’ ability to extract water from drying sand. Under equally high soil moisture, canopy stomatal conductance (G_S) of fertilized trees (F) was 50% that of irrigated/fertilized trees (IF), irrigated trees (I), and untreated control trees (C). As predicted from theory, F trees also decreased their stomatal sensitivity to vapour pressure deficit by 50%. The lower G_S in F was associated with 50% reduction in leaf‐specific hydraulic conductance (K_L) compared with other treatments. The lower K_L in F was in turn a result of a higher leaf area per sapwood area and a lower specific conductivity (conducting efficiency) of the plant and its root xylem. The root xylem of F trees was also 50% more resistant to cavitation than the other treatments. A transport model predicted that the lower A_R:A_L in IF trees resulted in a considerably restricted ability to extract water during drought. However, this deficiency was not exposed because irrigation minimized drought. In contrast, the lower A_R:A_L in F trees caused only a limited restriction in water extraction during drought owing to the more cavitation resistant root xylem in this treatment. In both fertilized treatments, approximate safety margins from predicted hydraulic failure were minimal suggesting increased vulnerability to drought‐induced dieback compared with non‐fertilized trees. However, IF trees are likely to be so affected even under a mild drought if irrigation is withheld.     

伊犁河谷退化野果林中新疆野苹果茎的水力结构

为揭示伊犁河谷退化野果林中新疆野苹果植株个体的死亡机理,对比研究3种长势(Ⅰ级,20%枯枝率;Ⅱ级,40%~60%枯枝率;Ⅲ级,>80%枯枝率)新疆野苹果茎的水力结构特征的差异以及水分运输有效性和安全性及其影响因素。结果表明: 随野果林退化程度的增加,3种长势的新疆野苹果茎的边材比导水率和叶比导水率呈下降趋势,但差异不显著;傍晚时的栓塞程度、栓塞脆弱性呈显著增加的趋势,其中Ⅰ、Ⅱ和Ⅲ级长势个体的导水率损失50%时木质部水势分别为-1.87、-1.35和-0.53 MPa;黎明前和正午叶水势、水力安全边际均为Ⅰ级>Ⅱ级>Ⅲ级;与水力学相关的木质部解剖结构以及枝叶性状在3种长势个体之间存在显著差异。相关性分析表明,新疆野苹果茎木质部水分运输的有效性和安全性之间存在弱的权衡关系。新疆野苹果茎的水力结构变化伴随着新疆野果林的衰退过程。野果林的退化会加剧新疆野苹果木质部导管的栓塞化程度,降低其抵抗空穴化栓塞的能力,新疆野苹果面临水力失衡的威胁更大。     

Water deficits and hydraulic limits to leaf water supply

Many aspects of plant water use -- particularly in response to soil drought -- may have as their basis the alteration of hydraulic conductance from soil to canopy. The regulation of plant water potential (Psi) by stomatal control and leaf area adjustment may be necessary to maximize water uptake on the one hand, while avoiding loss of hydraulic contact with the soil water on the other. Modelling the changes in hydraulic conductance with pressure gradients in the continuum allows the prediction of water use as a function of soil environment and plant architectural and xylem traits. Large differences in water use between species can be attributed in part to differences in their 'hydraulic equipment' that is presumably optimized for drawing water from a particular temporal and spatial niche in the soil environment. A number of studies have identified hydraulic limits as the cause of partial or complete foliar dieback in response to drought. The interactions between root:shoot ratio, rooting depth, xylem properties, and soil properties in influencing the limits to canopy water supply can be used to predict which combinations should optimize water use in a given circumstance. The hydraulic approach can improve our understanding of the coupling of canopy processes to soil environment, and the adaptive significance of stomatal behaviour.     

Biophysical properties and functional significance of stem water storage tissues in Neotropical savanna trees

Biophysical characteristics of sapwood and outer parenchyma water storage compartments were studied in stems of eight dominant Brazilian Cerrado tree species to assess the impact of differences in tissue capacitance on whole-plant water relations. The rate of decline in tissue water potential with relative water content (RWC) was greater in the outer parenchyma than in the sapwood for most of the species, resulting in tissue-and species-specific differences in capacitance. Sapwood capacitance on a tissue volume basis ranged from 40 to 160 kg m-3 MPa-1, whereas outer parenchyma capacitance ranged from 25 to only 60 kg m-3 MPa-1. In addition, osmotic potentials at full turgor and at the turgor loss point were more negative for the outer parenchyma compared with the sapwood, and the maximum bulk elastic modulus was higher for the outer parenchyma than for the sapwood. Sapwood capacitance decreased linearly with increasing sapwood density across species, but there was no significant correlation between outer parenchyma capacitance and tissue density. Midday leaf water potential, the total hydraulic conductance of the soil/leaf pathway and stomatal conductance to water vapour (gs) all increased with stem volumetric capacitance, or with the relative contribution of stored water to total daily transpiration. However, the difference between the pre-dawn water potential of non-transpiring leaves and the weighted average soil water potential, a measure of the water potential disequilibrium between the plant and soil, increased asymptotically with total stem capacitance across species, implying that overnight recharge of water storage compartments was incomplete in species with greater capacitance. Overall, stem capacitance contributes to homeostasis in the diurnal and seasonal water balance of Cerrado trees.     

Impacts of long‐term precipitation manipulation on hydraulic architecture and xylem anatomy of piñon and juniper in Southwest USA

Hydraulic architecture imposes a fundamental control on water transport, underpinning plant productivity, and survival. The extent to which hydraulic architecture of mature trees acclimates to chronic drought is poorly understood, limiting accuracy in predictions of forest responses to future droughts. We measured seasonal shoot hydraulic performance for multiple years to assess xylem acclimation in mature piñon (Pinus edulis ) and juniper (Juniperus monosperma ) after 3+ years of precipitation manipulation. Our treatments consisted of water addition (+20% ambient precipitation), partial precipitation‐exclusion (?45% ambient precipitation), and exclusion‐structure control. Supplemental watering elevated leaf water potential, sapwood‐area specific hydraulic conductivity, and leaf‐area specific hydraulic conductivity relative to precipitation exclusion. Shifts in allocation of leaf area to sapwood area enhanced differences between irrigated and droughted K _L in piñon but not juniper. Piñon and juniper achieved similar K _L under ambient conditions, but juniper matched or outperformed piñon in all physiological measurements under both increased and decreased precipitation treatments. Embolism vulnerability and xylem anatomy were unaffected by treatments in either species. Absence of significant acclimation combined with inferior performance for both hydraulic transport and safety suggests piñon has greater risk of local extirpation if aridity increases as predicted in the southwestern USA.     

Diurnal and seasonal variation in root xylem embolism in neotropical savanna woody species: impact on stomatal control of plant water status

Vulnerability to water-stress-induced embolism and variation in the degree of native embolism were measured in lateral roots of four co-occurring neotropical savanna tree species. Root embolism varied diurnally and seasonally. Late in the dry season, loss of root xylem conductivity reached 80% in the afternoon when root water potential (psi root) was about -2.6 MPa, and recovered to 25-40% loss of conductivity in the morning when psi root was about -1.0 MPa. Daily variation in psi root decreased, and root xylem vulnerability and capacitance increased with rooting depth. However, all species experienced seasonal minimum psi root close to complete hydraulic failure independent of their rooting depth or resistance to embolism. Predawn psi root was lower than psi soil when psi soil was relatively high (> -0.7 MPa) but became less negative than psi soil, later in the dry season, consistent with a transition from a disequilibrium between plant and soil psi induced by nocturnal transpiration to one induced by hydraulic redistribution of water from deeper soil layers. Shallow longitudinal root incisions external to the xylem prevented reversal of embolism overnight, suggesting that root mechanical integrity was necessary for recovery, consistent with the hypothesis that if embolism is a function of tension, refilling may be a function of internal pressure imbalances. All species shared a common relationship in which maximum daily stomatal conductance declined linearly with increasing afternoon loss of root conductivity over the course of the dry season. Daily embolism and refilling in roots is a common occurrence and thus may be an inherent component of a hydraulic signaling mechanism enabling stomata to maintain the integrity of the hydraulic pipeline in long-lived structures such as stems.     

Hydraulic architecture and water relations of Spartium junceum branches affected by a mycoplasm disease

Plant water relations, xylem anatomy and the hydraulic architecture of 1‐year‐old twigs of Spartium junceum, both healthy and affected by a phytoplasm disease, were studied. The disease causes twigs to be either shortened (witches broom disease, WBD) or flat (fasciate disease, FD). WBD twigs show a sevenfold increase in total leaf area, smaller and shorter xylem conduits, a higher stomatal conductance (g_l) and a decline of minimum leaf water potentials ( Ψ _l) below the turgor loss point. FD twigs had nearly twice the leaf area of the healthy controls as well as high g_l values and Ψ _l values below the turgor loss point. Moreover, significant differences between healthy and affected twigs in stem stomatal conductance (g_s) and in the total stem area were recorded. Affected twigs die back under drought stress, which is explained by a pronounced loss of hydraulic conductivity of the infected stems (40 and 60%) in FD and WBD as well as by the unfavourable ratio of weighted conduit radius ( Σ r⁴) to total surface area (A_t), so that the efficiency of the stem in supplying the whole transpiring area with water is strongly reduced.     

Drought resistance of 2-year-old saplings of Mediterranean forest trees in the field: relations between water relations,hydraulics and productivity

Drought resistance was examined for 2-year-old saplings of Ceratonia siliqua L., Olea oleaster Hoffmgg. et Link., Quercus suber L. and Q. pubescens Willd. growing in the field in Sicily, with the aim of testing their possible use in the reforestation of degraded areas. To this purpose, leaf conductance to water vapour (g _L), transpiration rate (E _L), relative water content (RWC) and water potential (_L) were measured between pre-dawn and sunset, monthly from May to November. Parallel measurements of loss of hydraulic conductance of twigs of the current year (PLC) were made together with an estimate of whole-plant hydraulic conductance (K _PLANT) on the basis of the ratio of maximum E _L to (_PD-_MIN), where _PD is pre-dawn _L and _MIN is the minimum diurnal _L. C. siliqua saplings maintained high g _L throughout the study period with high RWC (over 90%) and _L. They grew rapidly and increased their foliage area (A _L) by over 60% from May to winter rest. This was accompanied by low twig PLC (about 20% in September) resulting in high K _PLANT all through the study period. In contrast, O. oleaster saplings underwent distinct dehydration in July, i.e. they showed stomatal closure due to a drop in RWC (to 75%) and _L (to the turgor-loss point). This was apparently due to twig cavitation, resulting in a strong decrease of K _PLANT. Plants, however, tolerated summer drought, and showed growth (A _L increased by 15% from May to November). Saplings of Q. suber and Q. pubescens were much more vulnerable to twig cavitation (PLC was 35–48% from June to November) than the other two species, and their average K _PLANT was lower. Saplings of Q. suber and Q. pubescens grew only during the wet spring period, and no new foliage was produced thereafter. Changes in twig hydraulic conductance played a dominant role in determining changes in K _PLANT in that the two variables were well correlated to each other (r=0.68 for P=0.001). We conclude that C. siliqua is an ideal candidate for reforestation of Sicilian degraded areas as is O. oleaster that, however, requires some additional water supply in the summer, at least during the early years after plantation. In wetter locations of Sicily, to an altitude between 0 and 500 m, Q. suber and Q. pubescens can be used for reforestation with expected higher competitiveness of the latter over the former species.     

Diurnal water relations of expanding and full-sized cotton fruits and subtending leaves

Water relations of cotton (Gossypium hirsutum L.) fruits have received less attention than those of leaves, although crop water status has an important influence on fruit physiology. This study was conducted to describe diurnal changes in the water relations of cotton fruits and subtending leaves. Young, expanding fruits and full-sized fruits were compared because of previously reported changes in xylem maturity with ontogeny. Diurnal changes in relative water content were greater in leaves than in the capsule walls of fruits. The capsule walls of young fruits had a higher relative water content than subtending leaves, and water content was lower in full-sized (87%) than in expanding (92%) fruits. Water potentials of subtending leaves were always approximately 0-3 MPa lower than those of capsule walls. Water potential gradients favoured passive water flow from young fruits to branches, but water potentials of branches and the capsule walls of full-sized fruits were similar (?0.7 MPa). Water potential gradients were consistent throughout the day. These results indicate that xylem transport to young fruits is unlikely, but may occur in older fruits.     

油松幼苗水分传输效率及安全性对养分和水分添加的响应

探讨油松水分传输效率和安全性对养分和水分添加的响应是揭示其适应性的基础。该试验采用新改进的离心机技术,以2年生油松幼苗当年生枝为材料,试验设置对照（CK,不施肥、自然降水）、氮磷养分添加（F,按每年120 kg/hm²纯N和60 kg/hm²纯P水平添加N和P素,自然降水）及养分和水分同时添加（FI,按每年120 kg/ hm²纯N和60 kg/hm²纯P水平添加N素和P素,且补水100 mm）3个处理,研究了油松幼苗水分传输效率和栓塞脆弱性对氮磷养分和水分添加的响应。结果表明：（1）与CK相比,氮磷养分添加（F）增加了油松幼苗地径、冠幅和地上部生物量,但对比导水率（K_s）、比叶导水率（LSC）、Huber值、抵抗栓塞能力（P₅₀）及水分传输安全阈值均影响不大。（2）氮磷和水分同时添加处理（FI）的地径、株高、冠幅和地上部生物量显著高于CK和F处理,其K_s、LSC和Huber值与CK及F处理相比并无显著差异,但P₅₀比CK和F处理增加0.2 MPa左右,且水分传输安全阈值相对变小。研究表明,养分添加对油松幼苗当年生枝的水分传输效率和安全性影响不大,水分添加对水分传输效率亦无显著影响,但降低了水分传输的安全性,原因主要与管胞长度增加及管胞壁抗爆破阻力下降有关。     

Elevated [CO2] alleviates the impacts of water deficit on xylem anatomy and hydraulic properties of maize stems

Plants can modify xylem anatomy and hydraulic properties to adjust to water status. Elevated [CO₂] can increase plant water potential via reduced stomatal conductance and water loss. This raises the question of whether elevated [CO₂], which thus improves plant water status, will reduce the impacts of soil water deficit on xylem anatomy and hydraulic properties of plants. To analyse the impacts of water and [CO₂] on maize stem xylem anatomy and hydraulic properties, we exposed potted maize plants to varying [CO₂] levels (400, 700, 900, and 1,200 ppm) and water levels (full irrigation and deficit irrigation). Results showed that at current [CO₂], vessel diameter, vessel roundness, stem cross-section area, specific hydraulic conductivity, and vulnerability to embolism decreased under deficit irrigation; yet, these impacts of deficit irrigation were reduced at elevated [CO₂]. Across all treatments, midday stem water potential was tightly correlated with xylem traits and displayed similar responses. A distinct trade-off between efficiency and safety in stem xylem water transportation in response to water deficit was observed at current [CO₂] but not observed at elevated [CO₂]. The results of this study enhance our knowledge of plant hydraulic acclimation under future climate environments and provide insights into trade-offs in xylem structure and function.