Differential coordination of stomatal conductance,mesophyll conductance,and leaf hydraulic conductance in response to changing light across species

Stomatal conductance (g_s) and mesophyll conductance (g_m) represent major constraints to photosynthetic rate (A), and these traits are expected to coordinate with leaf hydraulic conductance (K_leaf) across species, under both steady‐state and dynamic conditions. However, empirical information about their coordination is scarce. In this study, K_leaf, gas exchange, stomatal kinetics, and leaf anatomy in 10 species including ferns, gymnosperms, and angiosperms were investigated to elucidate the correlation of H₂O and CO₂ diffusion inside leaves under varying light conditions. Gas exchange, K_leaf, and anatomical traits varied widely across species. Under light‐saturated conditions, the A, g_s, g_m, and K_leaf were strongly correlated across species. However, the response patterns of A, g_s, g_m, and K_leaf to varying light intensities were highly species dependent. Moreover, stomatal opening upon light exposure of dark‐adapted leaves in the studied ferns and gymnosperms was generally faster than in the angiosperms; however, stomatal closing in light‐adapted leaves after darkening was faster in angiosperms. The present results show that there is a large variability in the coordination of leaf hydraulic and gas exchange parameters across terrestrial plant species, as well as in their responses to changing light.     

Acclimation to humidity modifies the link between leaf size and the density of veins and stomata

The coordination of veins and stomata during leaf acclimation to sun and shade can be facilitated by differential epidermal cell expansion so large leaves with low vein and stomatal densities grow in shade, effectively balancing liquid‐ and vapour‐phase conductances. As the difference in vapour pressure between leaf and atmosphere (VPD) determines transpiration at any given stomatal density, we predict that plants grown under high VPD will modify the balance between veins and stomata to accommodate greater maximum transpiration. Thus, we examined the developmental responses of these traits to contrasting VPD in a woody angiosperm (Toona ciliata M. Roem.) and tested whether the relationship between them was altered. High VPD leaves were one‐third the size of low VPD leaves with only marginally greater vein and stomatal density. Transpirational homeostasis was thus maintained by reducing stomatal conductance. VPD acclimation changed leaf size by modifying cell number. Hence, plasticity in vein and stomatal density appears to be generated by plasticity in cell size rather than cell number. Thus, VPD affects cell number and leaf size without changing the relationship between liquid‐ and vapour‐phase conductances. This results in inefficient acclimation to VPD as stomata remain partially closed under high VPD.     

Relations between stomatal closure, leaf turgor and xylem vulnerability in eight tropical dry forest trees

This study examined the linkage between xylem vulnerability, stomatal response to leaf water potential (Ψ_L), and loss of leaf turgor in eight species of seasonally dry tropical forest trees. In order to maximize the potential variation in these traits species that exhibit a range of leaf habits and phenologies were selected. It was found that in all species stomatal conductance was responsive to Ψ_L over a narrow range of water potentials, and that Ψ_L inducing 50% stomatal closure was correlated with both the Ψ_L inducing a 20% loss of xylem hydraulic conductivity and leaf water potential at turgor loss in all species. In contrast, there was no correlation between the water potential causing a 50% loss of conductivity in the stem xylem, and the water potential at stomatal closure (Ψ_SC) amongst species. It was concluded that although both leaf and xylem characters are correlated with the response of stomata to Ψ_L, there is considerable flexibility in this linkage. The range of responses is discussed in terms of the differing leaf‐loss strategies exhibited by these species.     

The 'hydrology' of leaves: co-ordination of structure and function in temperate woody species

The hydraulic conductance of the leaf lamina (K_lamina) substantially constrains whole‐plant water transport, but little is known of its association with leaf structure and function. K_lamina was measured for sun and shade leaves of six woody temperate species growing in moist soil, and tested for correlation with the prevailing leaf irradiance, and with 22 other leaf traits. K_lamina varied from 7.40 × 10^?5 kg m^?2 s^?1 MPa^?1 for Acer saccharum shade leaves to 2.89 × 10^?4 kg m^?2 s^?1 MPa^?1 for Vitis labrusca sun leaves. Tree sun leaves had 15–67% higher K_lamina than shade leaves. K_lamina was co‐ordinated with traits associated with high water flux, including leaf irradiance, petiole hydraulic conductance, guard cell length, and stomatal pore area per lamina area. K_lamina was also co‐ordinated with lamina thickness, water storage capacitance, 1/mesophyll water transfer resistance, and, in five of the six species, with lamina perimeter/area. However, for the six species, K_lamina was independent of inter‐related leaf traits including leaf dry mass per area, density, modulus of elasticity, osmotic potential, and cuticular conductance. K_lamina was thus co‐ordinated with structural and functional traits relating to liquid‐phase water transport and to maximum rates of gas exchange, but independent of other traits relating to drought tolerance and to aspects of carbon economy.     

Parameterization of the CO2 and H2O gas exchange of several temperate deciduous broad-leaved trees at the leaf scale considering seasonal changes

A combined model to simulate CO₂ and H₂O gas exchange at the leaf scale was parameterized using data obtained from in situ leaf‐scale observations of diurnal and seasonal changes in the CO₂ and H₂O gas exchange of four temperate deciduous broad‐leaved trees using a porometric method. The model consists of a Ball et al. type stomatal conductance submodel [Ball, Woodrow & Berry, pp. 221–224 in Progress in Photosynthesis Research (ed. I. Biggins), Martinus‐Nijhoff Publishers, Dordrecht, The Netherlands, 1987] and a Farquhar et al. type biochemical submodel of photosynthesis (Farquhar, von Caemmerer & Berry, Planta 149, 78–90, 1980). In these submodels, several parameters were optimized for each tree species as representative of the quantitative characteristics related to gas exchange. The results show that the seasonal physiological changes of V_cmax25 in the biochemical model of photosynthesis should be used to estimate the long‐term CO₂ gas exchange. For R_d25 in the biochemical model of photosynthesis and m in the Ball et al. type stomatal conductance model, the difference should be counted during the leaf expansion period.     

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.     

胡杨木质部水分传导对盐胁迫的响应与适应北大核心CSCD

解析植物木质部导水率对逆境的响应和适应对促进植物抗逆性机理研究和受损植被恢复具有重要意义。该文以荒漠河岸林建群种胡杨(Populus euphratica)为研究对象,系统分析了胡杨幼株根、茎、叶水分传输通道对不同浓度盐胁迫的响应和适应。结果表明:(1)胡杨幼株根系对盐胁迫的敏感性高于茎和叶,盐胁迫下根系生长和根尖数显著受到抑制,根木质部易于发生栓塞,导水率明显降低。(2)胡杨幼株茎木质部导水率对盐胁迫的响应依盐浓度而定,轻度(0.05 mol·L–1 Na Cl)和中度(0.15 mol·L–1 Na Cl)盐胁迫下,胡杨可以通过协调导管输水的有效性和安全性来调节木质部的导水率,维持植物正常生长;重度(0.30 mol·L–1 Na Cl)盐胁迫下,胡杨茎木质部导管输水有效性和安全性均明显降低,木质部导水率显著下降,并伴随叶片气孔导度的显著降低,从而严重抑制了胡杨的光合和生长。     

Differential leaf expansion can enable hydraulic acclimation to sun and shade

Although leaf size is one of the most responsive plant traits to environmental change, the functional benefits of large versus small leaves remain unclear. We hypothesized that modification of leaf size within species resulting from differences in irradiance can allow leaves to acclimate to different photosynthetic or evaporative conditions while maintaining an efficient balance between hydraulic supply (vein density) and evaporative demand. To test this, we compared the function and anatomy of leaf hydraulic systems in the leaves of a woody angiosperm (Toona ciliata M. Roem.) grown under high and low irradiance in controlled conditions. Our results confirm that in this species, differential leaf expansion regulates the density of veins and stomata such that leaf hydraulic conductance and stomatal conductance remain proportional. A broader sample of field-grown tree species suggested that differences in leaf venation and stomatal traits induced by sun and shade were not regulated by leaf size in all cases. Our results, however, suggest that leaf size plasticity can provide an efficient way for plants to acclimate hydraulic and stomatal conductances to the contrasting evaporative conditions of sun and shade.     

Higher rates of leaf gas exchange are associated with higher leaf hydrodynamic pressure gradients

Steady-state leaf gas-exchange parameters and leaf hydraulic conductance were measured on 10 vascular plant species, grown under high light and well-watered conditions, in order to test for evidence of a departure from hydraulic homeostasis within leaves as hydraulic conductance varied across species. The plants ranged from herbaceous crop plants to mature forest trees. Across species, under standardized environmental conditions (saturating light, well watered), mean steady-state stomatal conductance to water vapour (g(w)) was highly correlated with mean rate of CO2 assimilation (A) and mean leaf hydraulic conductance normalized to leaf area (k(leaf)). The relationship between A and g(w) was well described by a power function, while that between A and k(leaf) was highly linear. Non-linearity in the relationship between g(w) and k(leaf) contributed to an increase in the hydrodynamic (transpiration-induced) water potential drawdown across the leaf (delta psi(leaf)) as k(leaf) increased across species, although across the 10 species the total increase in delta psi(leaf) was slightly more than twofold for an almost 30-fold increase in g(w). Higher rates of leaf gas exchange were therefore associated with higher k(leaf) and higher leaf hydrodynamic pressure gradients. A mechanistic model incorporating the stomatal hydromechanical feedback loop is used to predict the relationship between delta psi(leaf) and k(leaf), and to explore the coordination of stomatal and leaf hydraulic properties in supporting higher rates of leaf gas exchange.     

胡杨木质部水分传导对盐胁迫的响应与适应

解析植物木质部导水率对逆境的响应和适应对促进植物抗逆性机理研究和受损植被恢复具有重要意义。该文以荒漠河岸林建群种胡杨(Populus euphratica)为研究对象, 系统分析了胡杨幼株根、茎、叶水分传输通道对不同浓度盐胁迫的响应和适应。结果表明: (1)胡杨幼株根系对盐胁迫的敏感性高于茎和叶, 盐胁迫下根系生长和根尖数显著受到抑制, 根木质部易于发生栓塞, 导水率明显降低。(2)胡杨幼株茎木质部导水率对盐胁迫的响应依盐浓度而定, 轻度(0.05 mol·L^-1 NaCl)和中度(0.15 mol·L^-1 NaCl)盐胁迫下, 胡杨可以通过协调导管输水的有效性和安全性来调节木质部的导水率, 维持植物正常生长; 重度(0.30 mol·L^-1 NaCl)盐胁迫下, 胡杨茎木质部导管输水有效性和安全性均明显降低, 木质部导水率显著下降, 并伴随叶片气孔导度的显著降低, 从而严重抑制了胡杨的光合和生长。     

Limitation of transpiration by hydraulic conductance and xylem cavitation in Betula occidentalis

The extent to which stomatal conductance (g_s) was capable of responding to reduced hydraulic conductance (k)and preventing cavitation-inducing xylem pressures was evaluated in the small riparian tree, Betula occidentalis Hook. We decreased k by inducing xylem cavitation in shoots using an air-injection technique. From 1 to 18 d after shoot injection we measured midday transpiration rate (E), g_s, and xylem pressure (Ψ_p-xylem) on individual leaves of the crown. We then harvested the shoot and made direct measurements of k from the trunk (2–3 cm diameter) to the distal tip of the petioles of the same leaves measured for E and g_s. The k measurement was expressed per unit leaf area (k_l, leaf-specific conductance). Leaves measured within 2 d of shoot injection showed reduced g_s and E relative to non-injected controls, and both parameters were strongly correlated with k_l At this time, there was no difference in leaf Ψ_p-xylem between injected shoots and controls, and leaf Ψ_p-xylem was not significantly different from the highest cavitation-inducing pressure (Ψ_p-cav) in the branch xylem (-1.43 ± 0.029 MPa, n=8). Leaves measured 7–18 d after shoots were injected exhibited a partial return of g_s and E values to the control range. This was associated with a decrease in leaf Ψ_p-xylem below Ψ_p-cav and loss of foliage. The results suggest the stomata were incapable of long-term regulation of E below control values and that reversion to higher E caused dieback via cavitation.     

The influence of soil temperature and water content on belowground hydraulic conductance and leaf gas exchange in mature trees of three boreal species

Understanding stomatal regulation is fundamental to predicting the impact of changing environmental conditions on vegetation. However, the influence of soil temperature (ST) and soil water content (SWC) on canopy conductance (g_s) through changes in belowground hydraulic conductance (k_bg) remains poorly understood, because k_bg has seldom been measured in field conditions. Our aim was to (a) examine the dependence of k_bg on ST and SWC, (b) examine the dependence of g_s on k_bg and (c) test a recent stomatal optimization model according to which g_s and soil-to-leaf hydraulic conductance are strongly coupled. We estimated k_bg from continuous sap flow and xylem diameter measurements in three boreal species. k_bg increased strongly with increasing ST when ST was below +8°C, and typically increased with increasing SWC when ST was not limiting. g_s was correlated with k_bg in all three species, and modelled and measured g_s were well correlated in Pinus sylvestris (a model comparison was only possible for this species). These results imply an important role for k_bg in mediating linkages between the soil environment and leaf gas exchange. In particular, our finding that ST strongly influences k_bg in mature trees may help us to better understand tree behaviour in cold environments.     

Morphological and physiological reactions of young deciduous trees (Quercus robur L., Q. petraea [Matt.] Liebl., Fagus sylvatica L.) to waterlogging

One-year-old seedlings of Quercus robur L., Q. petraea (Matt.) Liebl. and Fagus sylvatica L. were cultivated in lysimeters and subjected to waterlogging for 17 weeks, interrupted by a five-week drainage period during summer. The growth of Q. robur was less affected by waterlogging than that of Q. petraea and Fagus. Waterlogging resulted in the formation of adventitious roots in Q. robur and Q. petraea, but not in Fagus. In contrast to Fagus, Q. robur and, to a lesser extent, Q. petraea were able to generate roots even below the water table. The hydraulic conductance of the excised root systems, the stomatal conductance and, in Fagus, the leaf water potential and the leaf-mass related hydraulic conductance were decreased by waterlogging. The decrease in the hydraulic conductance was largest in Fagus, and smallest in Q. robur. The roots of Fagus responded to anaerobic conditions with an increase in ethanol concentration. The measurements of nitrate reductase activities in roots and leaves provided no indications of a persistent contribution of NO₃ ⁻ metabolism to the alleviation of waterlogging-induced stress. It is concluded that Q. robur and, to a lesser extent, Q. petraea can tolerate waterlogging periods better than Fagus due to a different pattern of root formation, and to a better adjustment of leaf biomass production to the hydraulic conductivity of the root system. This revised version was published online in June 2006 with corrections to the Cover Date.     

A leaf gas exchange model that accounts for intra-canopy variability by considering leaf nitrogen content and local acclimation to radiation in grapevine (Vitis vinifera L.)

Understanding the distribution of gas exchange within a plant is a prerequisite for scaling up from leaves to canopies. We evaluated whether leaf traits were reliable predictors of the effects of leaf ageing and leaf irradiance on leaf photosynthetic capacity (V(cmax) , J(max) ) in field-grown vines (Vitis vinifera L). Simultaneously, we measured gas exchange, leaf mass per area (LMA) and nitrogen content (N(m) ) of leaves at different positions within the canopy and at different phenological stages. Daily mean leaf irradiance cumulated over 10 d (PPFD(10) ) was obtained by 3D modelling of the canopy structure. N(m) decreased over the season in parallel to leaf ageing while LMA was mainly affected by leaf position. PPFD(10) explained 66, 28 and 73% of the variation of LMA, N(m) and nitrogen content per area (N(a) ), respectively. Nitrogen content per unit area (N(a) = LMA × N(m) ) was the best predictor of the intra-canopy variability of leaf photosynthetic capacity. Finally, we developed a classical photosynthesis-stomatal conductance submodel and by introducing N(a) as an input, the model accurately simulated the daily pattern of gas exchange for leaves at different positions in the canopy and at different phenological stages during the season.     

Neither xylem collapse,cavitation, or changing leaf conductance drive stomatal closure in wheat

Identifying the drivers of stomatal closure and leaf damage during stress in grasses is a critical prerequisite for understanding crop resilience. Here, we investigated whether changes in stomatal conductance (g_s) during dehydration were associated with changes in leaf hydraulic conductance (K_leaf), xylem cavitation, xylem collapse, and leaf cell turgor in wheat (Triticum aestivum). During soil dehydration, the decline of g_s was concomitant with declining K_leaf under mild water stress. This early decline of leaf hydraulic conductance was not driven by cavitation, as the first cavitation events in leaf and stem were detected well after K_leaf had declined. Xylem vessel deformation could only account for <5% of the observed decline in leaf hydraulic conductance during dehydration. Thus, we concluded that changes in the hydraulic conductance of tissues outside the xylem were responsible for the majority of K_leaf decline during leaf dehydration in wheat. However, the contribution of leaf resistance to whole plant resistance was less than other tissues (<35% of whole plant resistance), and this proportion remained constant as plants dehydrated, indicating that K_leaf decline during water stress was not a major driver of stomatal closure.     

Coordination and trade‐offs between leaf and stem hydraulic traits and stomatal regulation along a spectrum of isohydry to anisohydry

The degree of plant iso/anisohydry, a widely used framework for classifying species‐specific hydraulic strategies, integrates multiple components of the whole‐plant hydraulic pathway. However, little is known about how it associates with coordination of functional and structural traits within and across different organs. We examined stem and leaf hydraulic capacitance and conductivity/conductance, stem xylem anatomical features, stomatal regulation of daily minimum leaf and stem water potential (Ψ), and the kinetics of stomatal responses to vapour pressure deficit (VPD) in six diverse woody species differing markedly in their degree of iso/anisohydry. At the stem level, more anisohydric species had higher wood density and lower native capacitance and conductivity. Like stems, leaves of more anisohydric species had lower hydraulic conductance; however, unlike stems, their leaves had higher native capacitance at their daily minimum values of leaf Ψ. Moreover, rates of VPD‐induced stomatal closure were related to intrinsic rather than native leaf capacitance and were not associated with species' degree of iso/anisohydry. Our results suggest a trade‐off between hydraulic storage and efficiency in the leaf, but a coordination between hydraulic storage and efficiency in the stem along a spectrum of plant iso/anisohydry.     

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.     

Impacts of leaf age and heat stress duration on photosynthetic gas exchange and foliar nonstructural carbohydrates in Coffea arabica

Given future climate predictions of increased temperature, and frequency and intensity of heat waves in the tropics, suitable habitat to grow ecologically, economically, and socially valuable Coffea arabica is severely threatened. We investigated how leaf age and heat stress duration impact recovery from heat stress in C. arabica. Treated plants were heated in a growth chamber at 49°C for 45 or 90 min. Physiological recovery was monitored in situ using gas exchange, chlorophyll fluorescence (the ratio of variable to maximum fluorescence, F_V/F_M), and leaf nonstructural carbohydrate (NSC) on mature and expanding leaves before and 2, 15, 25, and 50 days after treatment. Regardless of leaf age, the 90‐min treatment resulted in greater F_V/F_M reduction 2 days after treatment and slower recovery than the 45‐min treatment. In both treatments, photosynthesis of expanding leaves recovered more slowly than in mature leaves. Stomatal conductance (g_s) decreased in expanding leaves but did not change in mature leaves. These responses led to reduced intrinsic water‐use efficiency with increasing heat stress duration in both age classes. Based on a leaf energy balance model, aftereffects of heat stress would be exacerbated by increases in leaf temperature at low g_s under full sunlight where C. arabica is often grown, but also under partial sunlight. Starch and total NSC content of the 45‐min group significantly decreased 2 days after treatment and then accumulated 15 and 25 days after treatment coinciding with recovery of photosynthesis and F_V/F_M. In contrast, sucrose of the 90‐min group accumulated at day 2 suggesting that phloem transport was inhibited. Both treatment group responses contrasted with control plant total NSC and starch, which declined with time associated with subsequent flower and fruit production. No treated plants produced flowers or fruits, suggesting that short duration heat stress can lead to crop failure.     

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。     

Leaf responses to drought stress in Mediterranean accessions of Solanum lycopersicum: anatomical adaptations in relation to gas exchange parameters

In a previous study, important acclimation to water stress was observed in the Ramellet tomato cultivar (TR) from the Balearic Islands, related to an increase in the water‐use efficiency through modifications in both stomatal (g_s) and mesophyll conductances (g_m). In the present work, the comparison of physiological and morphological traits between TR accessions grown with and without water stress confirmed that variability in the photosynthetic capacity was mostly explained by differences in the diffusion of CO₂ through stomata and leaf mesophyll. Maximization of g_m under both treatments was mainly achieved through adjustments in the mesophyll thickness and porosity and the surface area of chloroplasts exposed to intercellular airspace (S_c). In addition, the lower g_m/S_c ratio for a given porosity in drought‐acclimated plants suggests that the decrease in g_m was due to an increased cell wall thickness. Stomatal conductance was also affected by drought‐associated changes in the morphological properties of stomata, in an accession and treatment‐dependent manner. The results confirm the presence of advantageous physiological traits in the response to drought stress in Mediterranean accessions of tomato, and relate them to particular changes in the leaf anatomical properties, suggesting specific adaptive processes operating at the leaf anatomical level.