Plant water relations at elevated CO2 -- implications for water-limited environments

Long-term exposure of plants to elevated [CO2] leads to a number of growth and physiological effects, many of which are interpreted in the context of ameliorating the negative impacts of drought. However, despite considerable study, a clear picture in terms of the influence of elevated [CO2] on plant water relations and the role that these effects play in determining the response of plants to elevated [CO2] under water-limited conditions has been slow to emerge. In this paper, four areas of research are examined that represent critical, yet uncertain, themes related to the response of plants to elevated [CO2] and drought. These include (1) fine-root proliferation and implications for whole-plant water uptake; (2) enhanced water-use efficiency and consequences for drought tolerance; (3) reductions in stomatal conductance and impacts on leaf water potential; and (4) solute accumulation, osmotic adjustment and dehydration tolerance of leaves. A survey of the literature indicates that the growth of plants at elevated [CO2] can lead to conditions whereby plants maintain higher (less negative) leaf water potentials. The mechanisms that contribute to this effect are not fully known, although CO2-induced reductions in stomatal conductance, increases in whole-plant hydraulic conductance and osmotic adjustment may be important. Less understood are the interactive effects of elevated [CO2] and drought on fine-root production and water-use efficiency, and the contribution of these processes to plant growth in water-limited environments. Increases in water-use efficiency and reductions in water use can contribute to enhanced soil water content under elevated [CO2]. Herbaceous crops and grasslands are most responsive in this regard. The conservation of soil water at elevated [CO2] in other systems has been less studied, but in terms of maintaining growth or carbon gain during drought, the benefits of CO2-induced improvements in soil water content appear relatively minor. Nonetheless, because even small effects of elevated [CO2] on plant and soil water relations can have important implications for ecosystems, we conclude that this area of research deserves continued investigation. Future studies that focus on cellular mechanisms of plant response to elevated [CO2] and drought are needed, as are whole-plant investigations that emphasize the integration of processes throughout the soil--plant--atmosphere continuum. We suggest that the hydraulic principles that govern water transport provide an integrating framework that would allow CO2-induced changes in stomatal conductance, leaf water potential, root growth and other processes to be uniquely evaluated within the context of whole-plant hydraulic conductance and water transport efficiency.     

Effects of tree height on branch hydraulics, leaf structure and gas exchange in California redwoods

We examined changes in branch hydraulic, leaf structure and gas exchange properties in coast redwood ( Sequoia sempervirens ) and giant sequoia ( Sequoiadendron giganteum ) trees of different sizes. Leaf-specific hydraulic conductivity ( k _L) increased with height in S. sempervirens but not in S. giganteum , while xylem cavitation resistance increased with height in both species. Despite hydraulic adjustments, leaf mass per unit area (LMA) and leaf carbon isotope ratios ( δ ¹³C) increased, and maximum mass-based stomatal conductance ( g _mass) and photosynthesis ( A _mass) decreased with height in both species. As a result, both A _mass and g _mass were negatively correlated with branch hydraulic properties in S. sempervirens and uncorrelated in S. giganteum . In addition, A _mass and g _mass were negatively correlated with LMA in both species, which we attributed to the effects of decreasing leaf internal CO₂ conductance ( g _i). Species-level differences in wood density, LMA and area-based gas exchange capacity constrained other structural and physiological properties, with S. sempervirens exhibiting increased branch water transport efficiency and S. giganteum exhibiting increased leaf-level water-use efficiency with increasing height. Our results reveal different adaptive strategies for the two redwoods that help them compensate for constraints associated with growing taller, and reflect contrasting environmental conditions each species faces in its native habitat.     

Leafminers help us understand leaf hydraulic design

Leaf hydraulics of Aesculus hippocastanum L. were measured over the growing season and during extensive leaf mining by the larvae of an invasive moth (Cameraria ohridella Deschka et Dimic) that specifically destroy the palisade tissue. Leaves showed seasonal changes in hydraulic resistance (R_lamina) which were related to ontogeny. After leaf expansion was complete, the hydraulic resistance of leaves and the partitioning of resistances between vascular and extra‐vascular compartments remained unchanged despite extensive disruption of the palisade by leafminers (up to 50%). This finding suggests that water flow from the petiole to the evaporation sites might not directly involve the palisade cells. The analysis of the temperature dependence of R_lamina in terms of Q₁₀ revealed that at least one transmembrane step was involved in water transport outside the leaf vasculature. Anatomical analysis suggested that this symplastic step may be located at the bundle sheath where the apoplast is interrupted by hydrophobic thickening of cell walls. Our findings offer some support to the view of a compartmentalization of leaves into well‐organized water pools so that the transpiration stream would involve veins, bundle sheath and spongy parenchyma, while the palisade tissue would be largely by‐passed with the possible advantage of protecting cells from short‐term fluctuations in water status.     

Sexes show contrasting patterns of leaf and crown carbon gain in a dioecious rainforest shrub

The sexes of dioecious species may differ in a range of vegetative and reproductive traits as well as in physiological traits. In Siparuna grandiflora, a Neotropical dioecious shrub, we examined differences in leaf-level photosynthesis of different classes of leaf age and, using simulation models, explored whether differences in leaf-level carbon gain led to sex differences in whole-plant daily carbon gain. Male plants had higher photosynthetic capacity at the leaf level. As leaves of both sexes aged their photosynthetic capacity and specific leaf area declined as expected. Simulations of daily carbon gain using the architecturally explicit model Y-Plant and a non-architectural model incorporating a wide range of realistic light environments revealed that the difference in leaf-level photosynthetic capacity did not translate into greater crown-level carbon gain for males. Rather, differences in patterns of allocation to leaf area allow females to achieve higher crown-level carbon gain. The results demonstrate that sex differences at the leaf level do not necessarily predict patterns at the whole-plant level.     

PIP1 plasma membrane aquaporins in tobacco: from cellular effects to function in plants

The molecular functions of several aquaporins are well characterized (e.g., by analysis of aquaporin-expressing Xenopus oocytes). However, their significance in the physiology of water transport in multicellular organisms remains uncertain. The tobacco plasma membrane aquaporin NtAQP1 was used to elucidate this issue. By comparing antisense plants that were inhibited in NtAQP1 expression with control plants, we found evidence for NtAQP1 function in cellular and whole-plant water relations. The consequences of a decrease in cellular water permeability were determined by measurement of transpiration rate and stem and leaf water potential as well as growth experiments under extreme soil water depletion. Plants impaired in NtAQP1 expression showed reduced root hydraulic conductivity and lower water stress resistance. In conclusion, our results emphasize the importance of symplastic aquaporin-mediated water transport in whole-plant water relations.     

Scaling of xylem vessels and veins within the leaves of oak species

General models of plant vascular architecture, based on scaling of pipe diameters to remove the length dependence of hydraulic resistance within the xylem, have attracted strong interest. However, these models have neglected to consider the leaf, an important hydraulic component; they assume all leaves to have similar hydraulic properties, including similar pipe diameters in the petiole. We examine the scaling of the leaf xylem in 10 temperate oak species, an important hydraulic component. The mean hydraulic diameter of petiole xylem vessels varied by 30% among the 10 oak species. Conduit diameters narrowed from the petiole to the midrib to the secondary veins, consistent with resistance minimization, but the power function scaling exponent differed from that predicted for stems. Leaf size was an organizing trait within and across species. These findings indicate that leaf vasculature needs to be included in whole-plant scaling models, for these to accurately reflect and predict whole-plant transport and its implications for performance and ecology.     

Size-dependent mortality in a Neotropical savanna tree: the role of height-related adjustments in hydraulic architecture and carbon allocation

Size-related changes in hydraulic architecture, carbon allocation and gas exchange of Sclerolobium paniculatum (Leguminosae), a dominant tree species in Neotropical savannas of central Brazil (Cerrado), were investigated to assess their potential role in the dieback of tall individuals. Trees greater than ∼6-m-tall exhibited more branch damage, larger numbers of dead individuals, higher wood density, greater leaf mass per area, lower leaf area to sapwood area ratio (LA/SA), lower stomatal conductance and lower net CO₂ assimilation than small trees. Stem-specific hydraulic conductivity decreased, while leaf-specific hydraulic conductivity remained nearly constant, with increasing tree size because of lower LA/SA in larger trees. Leaves were substantially more vulnerable to embolism than stems. Large trees had lower maximum leaf hydraulic conductance ( K _leaf) than small trees and all tree sizes exhibited lower K _leaf at midday than at dawn. These size-related adjustments in hydraulic architecture and carbon allocation apparently incurred a large physiological cost: large trees received a lower return in carbon gain from their investment in stem and leaf biomass compared with small trees. Additionally, large trees may experience more severe water deficits in dry years due to lower capacity for buffering the effects of hydraulic path-length and soil water deficits.     

Loss of whole-tree hydraulic conductance during severe drought and multi-year forest die-off

Understanding the pathways through which drought stress kills woody vegetation can improve projections of the impacts of climate change on ecosystems and carbon-cycle feedbacks. Continuous in situ measurements of whole trees during drought and as trees die hold promise to illuminate physiological pathways but are relatively rare. We monitored leaf characteristics, water use efficiency, water potentials, branch hydraulic conductivity, soil moisture, meteorological variables, and sap flux on mature healthy and sudden aspen decline-affected (SAD) trembling aspen (Populus tremuloides) ramets over two growing seasons, including a severe summer drought. We calculated daily estimates of whole-ramet hydraulic conductance and modeled whole-ramet assimilation. Healthy ramets experienced rapid declines of whole-ramet conductance during the severe drought, providing an analog for what likely occurred during the previous drought that induced SAD. Even in wetter periods, SAD-affected ramets exhibited fivefold lower whole-ramet hydraulic conductance and sevenfold lower assimilation than counterpart healthy ramets, mediated by changes in leaf area, water use efficiency, and embolism. Extant differences between healthy and SAD ramets reveal that ongoing multi-year forest die-off is primarily driven by loss of whole-ramet hydraulic capability, which in turn limits assimilation capacity. Branch-level measurements largely captured whole-plant hydraulic limitations during drought and mortality, but whole-plant measurements revealed a potential role of other losses in the hydraulic continuum. Our results highlight the importance of a whole-tree perspective in assessing physiological pathways to tree mortality and indicate that the effects of mortality on these forests’ assimilation and productivity are larger than expected based on canopy leaf area differences.     

Irrigation effects on daily and seasonal variations of trunk sap flow and leaf water relations in olive trees

Irrigation effects on whole-plant sap flow and leaf-level water relations were characterised throughout a growing season in an experimental olive (Olea europaea L.) orchard. Atmospheric evaporative demand and soil moisture conditions for irrigated and non-irrigated olive trees were also monitored. Whole-plant water use in field-grown irrigated and rain fed olive trees was determined using a xylem sap flow method (compensation heat-pulse velocity). Foliage gas exchange and water potentials were determined throughout the experimental period. Physiological parameters responded diurnally and seasonally to variations in tree water status, soil moisture conditions and atmospheric evaporative demand. There was a considerable degree of agreement between daily transpiration deduced from heat-pulse velocity and that determined by calibration using the Penman–Monteith equation in the field. Summer drought caused decreasing leaf gas exchange and water potentials, and a progressive increase in hydraulic conductance (stronger in non-irrigated than irrigated trees), probably attributable to modifications in hydraulic properties at the soil-root interface. Negligible hysteresis, attributable to low plant capacitance, was observed in the relationship between leaf water potential and sap flow. A proportional decrease in maximum daily leaf conductance with increasing vapour pressure deficit was observed, while mean daytime canopy stomatal conductance decreased with the season. As a result, plant water use was limited and excessive drought stress prevented. Non-irrigated olive trees recovered after the summer drought, showing a physiological behaviour similar to that of irrigated trees. In addition to physiological and environmental factors, there are endogenous keys (chemical signals) influencing leaf level parameters. Olive trees are confirmed to be economical and sparing users of soil water, with an efficient xylem sap transport, maintenance of significant gas exchange and transpiration, even during drought stress.     

Photosynthesis of Eucalyptus globulus with Mycosphaerella leaf disease

Mycosphaerella leaf disease (MLD) is a major cause of foliage damage in Eucalyptus globulus plantations. Our study is the first to describe the physiological effects of MLD on E. globulus leaves. It involved measurements on both field and potted plants. Changes in photosynthetic parameters in response to MLD were quantified in a study using gas exchange techniques. There was a negative linear relationship between light-saturated photosynthesis (A(max)) and leaf-level damage from MLD. Reductions in A(max) were proportionally greater than might be expected from the reduction in green leaf area as a result of the disease, indicating that asymptomatic tissue also was affected by MLD. The reductions in A(max) were not related to increases in stomatal resistance, but were a result of reduced activity of ribulose bisphosphate carboxylase (Rubisco) and changes in the capacity for ribulose bisphosphate (RuBP) regeneration. Changes in mesophyll resistance to CO2 were also implicated. The effect of MLD was similar at different sites and irrespective of tree-level infection, suggesting a general leaf-level response of E. globulus to MLD.     

Homeostasis in leaf water potentials on leeward and windward sides of desert shrub crowns: water loss control vs. high hydraulic efficiency

Phenotypic plasticity in morphophysiological leaf traits in response to wind was studied in two dominant shrub species of the Patagonian steppe, used as model systems for understanding effects of high wind speed on leaf water relations and hydraulic properties of small woody plants. Morpho-anatomical traits, hydraulic conductance and conductivity and water relations in leaves of wind-exposed and protected crown sides were examined during the summer with nearly continuous high winds. Although exposed sides of the crowns were subjected to higher wind speeds and air saturation deficits than the protected sides, leaves throughout the crown had similar minimum leaf water potential (Ψ_L). The two species were able to maintain homeostasis in minimum Ψ_L using different physiological mechanisms. Berberis microphylla avoided a decrease in the minimum Ψ_L in the exposed side of the crown by reducing water loss by stomatal control, loss of cell turgor and low epidermal conductance. Colliguaja integerrima increased leaf water transport efficiency to maintain transpiration rates without increasing the driving force for water loss in the wind-exposed crown side. Leaf physiological changes within the crown help to prevent the decrease of minimum Ψ_L and thus contribute to the maintenance of homeostasis, assuring the hydraulic integrity of the plant under unfavorable conditions. The responses of leaf traits that contribute to mechanical resistance (leaf mass per area and thickness) differed from those of large physiological traits by exhibiting low phenotypic plasticity. The results of this study help us to understand the unique properties of shrubs which have different hydraulic architecture compared to trees.     

Heteroblastic development and the optimal partitioning of traits among contrasting environments in Acacia implexa

Background and Aims: Optimal partitioning theory (OPT) predicts plants will allocatebiomass to organs where resources are limiting. Studies of OPTfocus on root, stem and leaf mass ratios where roots and stemsare often further sub-divided into organs such as fine roots/taproots or branches/main stem. Leaves, however, are rarely sub-dividedinto different organs. Heteroblastic species develop juvenileand adult foliage and provide an opportunity of sub-dividingleaf mass ratio into distinct organs. Acacia implexa (Mimosaceae)is a heteroblastic species that develops compound (juvenile),transitional and phyllode (adult) leaves that differ dramaticallyin form and function. The aims of the present study were togrow A. implexa to examine patterns of plastic development ofwhole-plant and leaf traits under the OPT framework. Methods: Plants were grown in a glasshouse under contrasting nutrient,light and water environments in a full factorial design. Allocationto whole-plant and leaf-level traits was measured and analysedwith multivariate statistics. Key Results: Whole-plant traits strongly followed patterns predicted by OPT.Leaf-level traits showed a more complex pattern in responseto experimental treatments. Compound leaves on low nutrientplants had significantly lower specific leaf area (SLA) andwere retained for longer as quantified by a significantly greatercompound leaf mass ratio after 120 d. There was no significantdifference in SLA of compound leaves in the light treatment,yet transitional SLA was significantly higher under the lowlight treatment. The timing of heteroblastic shift from compoundto transitional leaves was significantly delayed only in thelow light treatment. Therefore, plants in the light treatmentresponded at the whole-plant level by adjusting allocation toproductive compound leaves and at the leaf-level by adjustingSLA. There were no significant SLA differences in the watertreatment despite strong trends at the whole-plant level. Conclusion: Explicitly sub-dividing leaves into different types providedgreater insights into OPT.     

植物叶片水力与经济性状权衡关系的研究进展

叶片既是植物光合产物形成的主要场所, 又是整株植物的水力瓶颈、应对灾难性水力失调的安全阀门, 是植物碳水耦合权衡的重要器官。叶经济型谱反映了叶片经济性状“投资-收益”的权衡, 为验证植物进化过程中形成的物种对策提供了适用的理论框架。叶片水力性状变化会影响叶片经济性状及植物存活和生长。因此, 探索植物叶片水力与经济性状的权衡关系, 对建立植物碳-水耦合模型、揭示植物水-碳投资机理、扩展植物性状型谱等均有重要意义。该文首先综述了叶片水力性状、经济性状及两者之间的权衡关系, 分析了叶片导水率与水力脆弱性、失膨点水势、水容、安全阈值等水力性状以及与叶片的形态、结构和气体交换功能性状之间的关系。然后, 从叶片形态、解剖和叶脉网络结构以及气孔功能方面探讨了叶片水力性状与经济性状的调节机制。最后, 提出今后应加强三方面的研究: (1)探索建立植物根-茎-叶水力输导系统的碳-氮-水资源的整株经济型谱, 以揭示植物功能结构耦合、高效固碳用水的生理生态学机制; (2)探索叶片水力安全、水力效率和固碳效率之间的普适性权衡关系, 以深入理解抗旱植物叶片构建的生物物理结构与生理代谢的关系; (3)探索个体水平碳水代谢关系、水分运输与生长速率的耦合, 为代谢推演理论和植物群落尺度预测提供基础。     

Sensitivity of growth of roots versus leaves to water stress: biophysical analysis and relation to water transport

Water transport is an integral part of the process of growth by cell expansion and accounts for most of the increase in cell volume characterizing growth. Under water deficiency, growth is readily inhibited and growth of roots is favoured over that of leaves. The mechanisms underlying this differential response are examined in terms of Lockhart's equations and water transport. For roots, when water potential (psi) is suddenly reduced, osmotic adjustment occurs rapidly to allow partial turgor recovery and re-establishment of psi gradient for water uptake, and the loosening ability of the cell wall increases as indicated by a rapid decline in yield-threshold turgor. These adjustments permit roots to resume growth under low psi. In contrast, in leaves under reductions in psi of similar magnitude, osmotic adjustment occurs slowly and wall loosening ability either does not increase substantially or actually decreases, leading to marked growth inhibition. The growth region of both roots and leaves are hydraulically isolated from the vascular system. This isolation protects the root from low psi in the mature xylem and facilitates the continued growth into new moist soil volume. Simulations with a leaky cable model that includes a sink term for growth water uptake show that growth zone psi is barely affected by soil water removal through transpiration. On the other hand, hydraulic isolation dictates that psi of the leaf growth region would be low and subjected to further reduction by high evaporative demand. Thus, a combination of transport and changes in growth parameters is proposed as the mechanism co-ordinating the growth of the two organs under conditions of soil moisture depletion. The model simulation also showed that roots behave as reversibly leaky cable in water uptake. Some field data on root water extraction and vertical profiles of psi in shoots are viewed as manifestations of these basic phenomena. Also discussed is the trade-off between high xylem conductance and strong osmotic adjustment.     

Trade-offs between plant leaf hydraulic and economic traits

Leaf is the most important organ for carbon-water coupling of a plant because it is the primary medium for photosynthesis. It also acts as the hydraulic bottleneck and safety valve against hydraulic catastrophic dysfunctions. The leaf economics spectrum, which reflects the balance between investments and returns of leaf economic traits, provides a useful framework for examining species strategies as shaped by their evolutionary history. Changes in leaf hydraulic traits will influence leaf economic traits as well as plant survival and growth. Exploring trade-offs between leaf hydraulic and economic traits is thus of significance for modeling carbon-water relations, understanding the mechanisms of water/carbon investments, and extending the leaf economic spectrum. In this review, we first examined the trade-offs between leaf hydraulic and economic traits. Specially, we analyzed the relationships between leaf hydraulic conductivity and hydraulic vulnerability, water potential at the turgor loss point, water capacitance, safety margin, and leaf morphological, structural and functional traits. We then discussed potential mechanisms regulating leaf hydraulic and economic traits from leaf morphology, anatomy, venation, and stomatal functions. Finally, we proposed future research to: (1) develop an integrated whole-plant economics spectrum, including carbon-nitrogen-water resources and root-stem-leaf hydraulic transport system that will help revealing ecophysiological mechanisms of plant structure-functional coupling, carbon sequestration and water use; (2) explore a generalized trade-offs among leaf hydraulic safety, hydraulic efficiency and carbon fixation efficiency to advance our understanding of the relationships between biophysical structure and physiological metabolism in plant leaf construction under drought stress; and (3) explore the carbon-water metabolic relationship and coupling of water transport and growth rate for the metabolic theory and predictions at community scale.     

散孔材与环孔材树种枝干、叶水力学特性的比较研究

为揭示散孔材与环孔材树种树木水分生理特性的差异,选取了常见的3种散孔材落叶树种(毛白杨、法国梧桐和樱花)和3种环孔材落叶树种(刺槐、合欢和白蜡),研究了其枝干与叶水力学性质的差异及其协调性。结果表明:3种环孔材树种枝干横截面积基础上的最大比导水率(Ks-max)大于3种散孔材树种,但其木质部对空穴化的脆弱性(P50branch)高于散孔材树种,6种树木枝干的水分传输能力和抵抗空穴化能力之间存在一种相互制约的权衡关系。3种散孔材与3种环孔材树种的叶最大水力导度(Kl-max)和水力脆弱性(P50leaf)并无显著差异;对于3种散孔材树种,叶的水力脆弱性要高于枝干,但对3种环孔材树种而言,枝干的水力脆弱性要高于叶。6种树木枝干和叶的水力学性质(Kmax、P50)之间并无相关关系。这些结果表明:散孔材与环孔材树种的枝干水力学特性有明显差异,但叶水力学特性无差异;枝干与叶水力学性质之间是相互独立的。     

Could rapid diameter changes be facilitated by a variable hydraulic conductance?

Adequate radial water transport between elastic bark tissue and xylem is crucial in trees, because it smoothens abrupt changes in xylem water potential, greatly reducing the likelihood of suffering dangerous levels of embolism. The radial hydraulic conductance involved is generally thought to be constant. Evidence collected about variable root and leaf hydraulic conductance led us to speculate that radial hydraulic conductance in stem/branches might also be variable and possibly modulated by putative aquaporins. We therefore correlated diameter changes in walnut (Juglans regia L.) with changes in water potential, altered by perfusion of twig samples with D-mannitol solutions having different osmotic potentials. Temperature and cycloheximide (CHX; a protein synthesis inhibitor) treatments were performed. The temperature response and diameter change inhibition found in CHX-treated twigs underpinned our hypothesis that radial hydraulic conductance is variable and likely mediated by a putative aquaporin abundance and/or activity. Our data demonstrate that radial water transport in stem/branches can take two routes in parallel: an apoplastic and a cell-to-cell route. The contribution of either route depends on the hydraulic demand and is closely linked to a boost of putative aquaporins, causing radial conductance to be variable. This variability should be considered when interpreting and modelling diameter changes.     

Stomatal and hydraulic conductance in growing sugarcane: stomatal adjustment to water transport capacity*

Abstract Stomatal conductance per unit leaf area in well-irrigated field- and greenhouse-grown sugarcane increased with leaf area up to 0.2 m² plant ¹, then declined so that maximum transpiration per plant tended to saturate rather than increase linearly with further increase in leaf area. Conductance to liquid water transport exhibited parallel changes with plant size. This coordiantion of vapour phase and liquid phase conductances resulted in a balance between water loss and water transport capacity, maintaining leaf water status remarkably constant over a wide range of plant size and growing conditions. The changes in stomatal conductance were not related to plant or leaf age. Partial defoliation caused rapid increases in stomatal conductance, to re-establish the original relationship with remaining leaf area. Similarly, pruning of roots caused rapid reductions in stomatal conductance, which maintained or improved leaf water status. These results suggest that sugarcane stomata adjusted to the ratio of total hydraulic conductance to total transpiring leaf area. This could be mediated by root metabolites in the transpiration stream, whose delivery per unit leaf area would be a function of the relative magnitudes of root system size, transpiration rate and leaf area.     

Leaf hydraulic conductivity and stomatal responses to humidity in amphistomatous leaves

The response of stomata to changes in humidity for a single surface of an amphistomatous leaf was investigated in Xanthium strumarium and Vicia faba using gas exchange and direct observation of stomatal apertures. The stomatal response to humidity for a given surface was found to be the same whether or not the humidity for the opposite surface was changed concurrently. Stomata on the surface for which humidity was constant showed no response to changes in humidity for the opposite surface. Despite large changes in epidermal turgor on the surface for which humidity was changed, there was no change in epidermal turgor for the surface with constant humidity. Measurements of transpiration and epidermal turgor as functions of the mole fraction gradient of water between leaf and air were used to calculate a value for leaf hydraulic resistance. The results suggest that in these species, the mechanism for the stomatal response to humidity resides in the epidermis or the mesophyll very close to the epidermis, and that most of the hydraulic resistance of the leaf occurs between the xylem and the evaporating sites.     

The ecological significance of long-distance water transport: short-term regulation, long-term acclimation and the hydraulic costs of stature across plant life forms

Plant hydraulic conductance, namely the rate of water flow inside plants per unit time and unit pressure difference, varies largely from plant to plant and under different environmental conditions. Herein the main factors affecting: (a) the scaling between whole‐plant hydraulic conductance and leaf area; (b) the relationship between gas exchange at the leaf level and leaf‐specific xylem hydraulic conductance; (c) the short‐term physiological regulation of plant hydraulic conductance under conditions of ample soil water, and (d) the long‐term structural acclimation of xylem hydraulic conductance to changes in environmental conditions are reviewed. It is shown that plant hydraulic conductance is a highly plastic character that varies as a result of multiple processes acting at several time scales. Across species ranging from coniferous and broad‐leaved trees to shrubs, crop and herbaceous species, and desert subshrubs, hydraulic conductance scaled linearly with leaf area, as expected from first principles. Despite considerable convergence in the scaling of hydraulic properties, significant differences were apparent across life forms that underlie their different abilities to conduct gas exchange at the leaf level. A simple model of carbon allocation between leaves and support tissues explained the observed patterns and correctly predicted the inverse relationships with plant height. Therefore, stature appears as a fundamental factor affecting gas exchange across plant life forms. Both short‐term physiological regulation and long‐term structural acclimation can change the levels of hydraulic conductance significantly. Based on a meta‐analysis of the existing literature, any change in environmental parameters that increases the availability of resources (either above‐ or below‐ground) results in the long‐term acclimation of a less efficient (per unit leaf area) hydraulic system.