Leaf morphology of 40 evergreen and deciduous broadleaved subtropical tree species and relationships to functional ecophysiological traits

We explored potential of morphological and anatomical leaf traits for predicting ecophysiological key functions in subtropical trees. We asked whether the ecophysiological parameters stomatal conductance and xylem cavitation vulnerability could be predicted from microscopy leaf traits. We investigated 21 deciduous and 19 evergreen subtropical tree species, using individuals of the same age and from the same environment in the Biodiversity‐Ecosystem Functioning experiment at Jiangxi (BEF‐China). Information‐theoretic linear model selection was used to identify the best combination of morphological and anatomical predictors for ecophysiological functions. Leaf anatomy and morphology strongly depended on leaf habit. Evergreen species tended to have thicker leaves, thicker spongy and palisade mesophyll, more palisade mesophyll layers and a thicker subepidermis. Over 50% of all evergreen species had leaves with multi‐layered palisade parenchyma, while only one deciduous species (Koelreuteria bipinnata) had this. Interactions with leaf habit were also included in best multi‐predictor models for stomatal conductance (g_s) and xylem cavitation vulnerability. In addition, maximum g_s was positively related to log ratio of palisade to spongy mesophyll thickness. Vapour pressure deficit (vpd) for maximum g_s increased with the log ratio of palisade to spongy mesophyll thickness in species having leaves with papillae. In contrast, maximum specific hydraulic conductivity and xylem pressure at which 50% loss of maximum specific xylem hydraulic conductivity occurred (Ψ₅₀) were best predicted by leaf habit and density of spongy parenchyma. Evergreen species had lower Ψ₅₀ values and lower maximum xylem hydraulic conductivities. As hydraulic leaf and wood characteristics were reflected in structural leaf traits, there is high potential for identifying further linkages between morphological and anatomical leaf traits and ecophysiological responses.     

Transpiration and stomatal control: a cross-species study of leaf traits in 39 evergreen and deciduous broadleaved subtropical tree species

Key message

Using an extensive dataset for 39 subtropical broadleaved tree species, we found traits of the leaf economics spectrum to be linked to mean stomatal conductance but not to stomatal regulation.

Abstract

The aim of our study was to establish links between stomatal control and functional leaf traits. We hypothesized that mean and maximum stomatal conductance (g _s) varies with the traits described by the leaf economics spectrum, such as specific leaf area and leaf dry matter content, and that high g _s values correspond to species with tender leaves and high photosynthetic capacity. In addition, we hypothesized that species with leaves of low stomata density have more limited stomatal closure than those with high stomata density. In order to account for confounding site condition effects, we made use of a common garden situation in which 39 deciduous and evergreen species of the same age were grown in a biodiversity ecosystem functioning experiment in Jiangxi (China). Daily courses of g _s were measured with porometry, and the species-specific g _s~vpd relationships were modeled. Our results show that mean stomatal conductance can be predicted from leaf traits that represent the leaf economics spectrum, with a positive relationship being related to leaf nitrogen content and a negative relationship with the leaf carbon: nitrogen ratio. In contrast, parameters of stomatal control were related to traits unassociated with the leaf economics spectrum. The maximum of the conductance~vpd curve was positively related to leaf carbon content and vein length. The vpd at the point of inflection of the conductance~vpd curve was lower for species with higher stomata density and higher for species with a high leaf carbon content. Overall, stomata size and density as well as vein length were more effective at explaining stomatal regulation than traits used in the leaf economics spectrum.     

Leaf Gas Exchange in Canopy Species of a Venezuelan Cloud Forest

Tropical cloud forests are considered humid ecosystems with frequent cloud cover down to the ground surface. However, seasonal variation in precipitation may induce short-term water stress. For canopy leaves, this water stress may also be a consequence of large atmospheric vapor pressure deficits. The objective of this work was to study five canopy cloud forest species to determine if there are restrictions to leaf gas exchange as a consequence of seasonality in precipitation and to daily water deficit due to air evaporative demand mainly during maximum incoming radiation hours. Seasonal daily courses of microclimatic variables (air temperature, relative humidity, photosynthetic photon flux density) and plant responses (leaf water potential, stomatal conductance, CO₂ assimilation rates, leaf nitrogen concentration) were measured at 2400 m asl in Monterrey, an intermontane valley of the Venezuelan Andes. A gradient in terms of responses to water stress conditions was observed between the species, with Clusia multiflora (a 46% reduction in stomatal conductance between seasons) as the most affected and Miconia resimoides (increased stomatal conductance) responding more favorably to slight water stress conditions. If we consider the limitations of water stress and/or light conditions on CO₂ assimilation we may arrange the species into those in which water stress conditions have a greater impact on leaf carbon gain, those where light conditions are determinant and one in which both water stress and light conditions may affect leaf carbon assimilation.     

叶脉结构与功能及其对叶片经济谱的影响

叶脉由贯穿于叶肉内部的维管组织及其外围机械组织构成,多样化的脉序及网络结构使叶脉系统发生变异和功能分化。该文综述了叶脉系统结构与功能的最新研究进展。通过聚焦叶脉分级系统的结构与功能及其在叶片经济谱(LES)中的重要性,解释叶脉性状与其它叶片功能性状之间的关系及机制。不同等级叶脉在机械支撑与水分运输方面存在功能分化,其中1–3级粗脉在维持叶片形状和叶表面积以及物理支撑方面发挥重要作用,有利于维持叶片最大受光面积;4级及以上细脉具有水分调节功能,它们与气孔相互协调,影响叶片水分运输、蒸腾散热和光合作用速率。叶片生长过程与叶脉发育的动态变化模式决定叶脉密度,并影响叶脉密度与叶片大小之间的关系:叶面积与粗脉密度呈显著负相关,与粗脉直径呈显著正相关,而与细脉密度无关。与叶脉性状相关的叶片经济谱框架模型预测,叶脉密度较高的叶片寿命短、比叶重较小,叶片最大碳同化速率、代谢速率以及资源获取策略潜力较高。     

热岛效应下亚热带城市植被叶气孔权衡特征及其与叶功能性状的关系

叶片气孔不仅是植物平衡光合-蒸腾关系的重要门户,也是影响大气碳循环与水循环的关键结构。分析热岛效应下福州市乔木、灌木、草本3种生活型和常绿、落叶2种叶习性植物的气孔性状间的差异及其与其他叶功能性状间的权衡关系有助于探究不同类型植物在热环境下的适应策略。以福州市区的自然和半自然植被为研究对象,测定441个植物样本的气孔特征、化学计量特征和形态特征,结果表明：（1）3种生活型、2种叶习性植物的气孔长度（SL）、气孔密度（SD）差异显著（P<0.05）,潜在气孔导度指数（PCI）不存在显著差异（P>0.05）。草本的SL高于灌木和乔木,乔木的SD最高,灌木次之,草本最低;落叶植物的SL高于常绿植物,SD低于常绿植物。（2）SL与SD间的权衡关系稳定存在于3种生活型和2种叶习性植物中,且随着不同生活型和落叶习性植物的生态策略而呈现各异的权衡特征,即当SL一定时,乔木的SD最大,灌木的SD最小,常绿植物的SD大于落叶植物。（3）气孔性状和叶片形态、化学计量特征紧密联系,SL与比叶面积（SLA）正相关（P<0.01）,与叶面积（LA）负相关（P<0.01）;SD与叶氮含量（LNC）、叶磷含量（LPC）、SLA负相关（P<0.01）,与LA正相关（P<0.01）;PCI与LNC、SLA负相关（P<0.01）,与叶厚度（LT）正相关（P<0.05）。（4）复杂的环境是气孔性状变异的重要驱动因素,SL、PCI均与年均温（MAT）负相关（P<0.05）。     

Optimization of leaf morphology in relation to leaf water status: A theory

The leaf economic traits such as leaf area, maximum carbon assimilation rate, and venation are all correlated and related to water availability. Furthermore, leaves are often broad and large in humid areas and narrower in arid/semiarid and hot and cold areas. We use optimization theory to explain these patterns. We have created a constrained optimization leaf model linking leaf shape to vein structure that is integrated into coupled transpiration and carbon assimilation processes. The model maximizes net leaf carbon gain (NPP_leaf) over the loss of xylem water potential. Modeled relations between leaf traits are consistent with empirically observed patterns. As the results of the leaf shape–venation relation, our model further predicts that a broadleaf has overall higher NPP_leaf compared to a narrowleaf. In addition, a broadleaf has a lower stomatal resistance compared to a narrowleaf under the same level of constraint. With the same leaf area, a broadleaf will have, on average, larger conduits and lower total leaf xylem resistance and thus be more efficient in water transportation but less resistant to cavitation. By linking venation structure to leaf shape and using water potential as the constraint, our model provides a physical explanation for the general pattern of the covariance of leaf traits through the safety–efficiency trade‐off of leaf hydraulic design.     

Does transpiration control stomatal responses to water vapour pressure deficit?

Three types of observations were used to test the hypothesis that the response of stomatal conductance to a change in vapour pressure deficit is controlled by whole-leaf transpiration rate or by feedback from leaf water potential. Varying the leaf water potential of a measured leaf by controlling the transpiration rate of other leaves on the plant did not affect the response of stomatal conductance to vapour pressure deficit in Glycine max. In three species, stomatal sensitivity to vapour pressure deficit was eliminated when measurements were made at near-zero carbon dioxide concentrations, despite the much higher transpiration rates of leaves at low carbon dioxide. In Abutilon theophrasti, increasing vapour pressure deficit sometimes resulted in both decreased stomatal conductance and a lower transpiration rate even though the response of assimilation rate to the calculated substomatal carbon dioxide concentration indicated that there was no ‘patchy’ stomatal closure at high vapour pressure deficit in this case. These results are not consistent with stomatal closure at high vapour pressure deficit caused by increased whole-leaf transpiration rate or by lower leaf water potential. The lack of response of conductance to vapour pressure deficit in carbon dioxide-free air suggests that abscisic acid may mediate the response.     

植物功能性状权衡关系的研究进展

植物功能性状权衡关系反映了植物在资源获取与分配中采取的不同策略, 是近年来生态学研究的一个热点问题。该综述从研究范围、叶性状、器官和植物类群4个方面入手, 简要介绍植物功能性状关系研究在近10余年是如何在叶经济谱(LES)的基础上逐渐扩展和深入的。1)相关研究拓展到全球更多极端环境与特殊气候地区, 发现在不同的气候环境条件下, 植物叶片功能性状关系相对稳定, 植物种内的功能性状关系已被证实与LES相似; 2)功能性状网络从最初的6个经济性状扩展到叶片的分解、燃烧和水力等性状, 发现叶片的分解速率和可燃性均与叶片形态性状、养分含量等显著相关, 但叶片水力性状与经济性状的关系则取决于所研究的物种及生存环境的水分条件; 3)研究对象从植物叶片拓展到了根、茎、花、种子及植株整体, 叶片的比叶质量与茎的木质密度、种子大小相耦合, 但叶片形态性状与根和花的相关性状却无显著相关关系, 证明这些器官可能是独立进化的; 4) LES可以很好地解释特殊维管植物的生存适应策略: 入侵植物具有较高的资源利用效率和更快的相对生长速率, 在LES中处于“低投入-快速回报”的一端; 食虫植物的叶片特化为捕食器官, 光合作用及生长速率相对较低, 居于LES “高投入-缓慢回报”的另一端, 此外, 无论是最古老的种子植物苏铁属(Cycas)植物, 或是蕨类和变水植物(苔藓和地衣), 其功能性状关系都与LES大致相同。该文梳理了功能性状关系研究的进展脉络, 提出了一些建议, 期望为未来植物功能性状关系研究的选题和发展提供一些参考。     

What do we know about parasitic plants and the leaf economic spectrum?

对于寄生植物和叶经济谱,我们了解多少？ 叶经济谱(leaf economic spectrum, LES)对维管植物中的关键性叶片性状间的相关性进行了量化,并且将这些性状的大量变异信息提炼为一根单轴。叶经济谱的显著优势是其近乎完美的普适性,且已在诸多领域得到了广泛的研究。然而,对于寄生植物与叶经济谱框架间关系的研究仍相对缺乏。由于叶经济谱的部分驱动力是碳获取中的生理性权衡,因此从理论上而言,寄生植物中的异养性(其取代了叶片的一些基本功能)可能导致对叶经济谱的偏离。利用从TRY数据库中获得的全局叶片性状数据,本研究评价了TRY数据库对寄生植物的整体代表性情况,然后将寄生植物叶片性状的叶经济谱组与非寄生的对应植物进行了比较。尽管寄生植物有着独特的生理特征,但它们并未显著偏离叶经济谱,不过还是有一些例子可以表明存在着叶经济谱上位置和性状间的差异。尤为重要的是,TRY数据库未能很好地代表寄生植物,因而就此得出的任何结论都还是不成熟的。     

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.     

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.     

Stomatal dynamics are limited by leaf hydraulics in ferns and conifers: results from simultaneous measurements of liquid and vapour fluxes in leaves

Stomatal responsiveness to vapour pressure deficit (VPD) results in continuous regulation of daytime gas‐exchange directly influencing leaf water status and carbon gain. Current models can reasonably predict steady‐state stomatal conductance (g_s) to changes in VPD but the g_s dynamics between steady‐states are poorly known. Here, we used a diverse sample of conifers and ferns to show that leaf hydraulic architecture, in particular leaf capacitance, has a major role in determining the g_s response time to perturbations in VPD. By using simultaneous measurements of liquid and vapour fluxes into and out of leaves, the in situ fluctuations in leaf water balance were calculated and appeared to be closely tracked by changes in g_s thus supporting a passive model of stomatal control. Indeed, good agreement was found between observed and predicted g_s when using a hydropassive model based on hydraulic traits. We contend that a simple passive hydraulic control of stomata in response to changes in leaf water status provides for efficient stomatal responses to VPD in ferns and conifers, leading to closure rates as fast or faster than those seen in most angiosperms.     

濒危种观光木小枝生物量分配与功能性状的纬度变异规律及其影响因素

为了探讨观光木当年生枝条生物量的分配规律和叶片功能性状的变化规律及其影响因素,该研究以濒危物种观光木(Michelia odora)为对象,测定了广西地区5个不同纬度上观光木当年生小枝及叶片功能性状.结果表明:(1)随着纬度的增加小枝总重和总叶重总体呈异速生长关系,其生物量更多趋于对小枝的构建.(2)观光木叶功能性状呈...     

Plant functional traits shape growth rate for xerophytic shrubs

• Trade-offs exist for xerophytic shrubs between functional traits, involving in water loss and assimilate accumulation, can contribute to its survival and growth rate regulation in arid environments. However, growth analysis based on plant functional traits has been focused on the study of herbs and woody species. It is still unclear how the functional traits of xerophytic shrubs regulate their growth rate.
• In this study, we selectedeight xerophytic shrubs as samples to analyze the regulation process of the functional traits of shrubs on growth rate. Plants were cultivated for three years, and three harvests (every one year) were carried out. Factors explaining between-species differences in relative growth rate (RGR) varied, depending on whether different ages were considered.
• The results showed that RGR was positively correlated with net assimilation rate, but there was a significant negative correlation with leaf area ration (LAR), specific leaf area (SLA), and leaf biomass ratio in the age 1. However, in the age 2, RGR showed a significant positive correlation with the morphological traits (i.e., leaf area ration and specific leaf area), but not with physiological traits (i.e., net assimilation rate) and leaf biomass allocation.
• Our results suggested that the fluctuation of environmental factors affects the regulation path of the plant functional traits on RGR of xerophytic shrubs. However, the analysis of causality model showed that no matter in which age, net assimilation rate and leaf area ration principally drive the variation in RGR among xerophytic shrubs.

     

Gas Exchange,Water Loss and Biomass Production in Rice and Wild Oryza Species in Well-Watered and Water-Limiting Growth Conditions

A major reason for the drought susceptibility of cultivated rice is the inability to regulate water loss as effectively as other cereals. We report studies of growth and gas exchange under well-watered and water-limiting conditions on selected species of Oryza and seek to relate this physiological information to the ecology of the genus. The immediate ancestor of the Oryza sativa cultigen is native to swamps and marshes, and wet habitats are typical for the genus as a whole. However, the ecological range of some species does extend to habitats that dry out seasonally. In all species studied growth was reduced by water deficit. In all species studied leaves had small absolute water contents and began to roll at a relative water content above 90%. However, there were species differences in leaf rolling and the response of stomatal conductance to an increase in vapour pressure deficit. Following re-watering, there were persistent reductions in stomatal conductance in most of the species tested, but the assimilation rate was not reduced in all of these cases. Where there was a persistant reduction in assimilation rate, there was also a reduction in carboxylation efficiency. It was a frequent observation that plants had a stomatal conductance greater than expected for their carbon assimilation rate; that is in the range where substantial changes in conductance have little effect upon photosynthesis. It is suggested that a reason for this may be the cooling of leaves which have a small thermal capacity in environments which often combine high temperature, humidity and irradiance. Large conductances combined with small water contents may be no disadvantage in the natural habitats of Oryza, but provide some reasons for the poor regulation of water loss in cultivated rice. Although there were significant differences in gas exchange amongst species the advantages that were observed over O. sativa were not of a magnitude likely to justify wide hybridisation. This implies that improvement in the drought resistance of rice is more likely to come from increasing water acquisition than from decreasing water loss.     

柯-青冈常绿阔叶林优势树种叶片性状变异及适应策略

厘清叶片功能性状的变异及性状间的权衡关系,对揭示植物对环境变化的响应及适应策略具有重要意义。以中亚热带柯(Lithocarpus glaber)-青冈(Cyclobalanopsis glauca)常绿阔叶林为对象,测量了1 hm²固定监测样地内6个优势树种的叶面积(LA)、比叶面积(SLA)、干物质含量(LDMC)、叶片厚度(LT)、叶片碳(LC)、叶片氮(LN)、叶片磷(LP)含量和叶片碳氮比(LC ∶ LN)8个功能性状,采用多重比较、主成分分析(PCA)法分析了群落叶片功能性状的生活型、种内和种间变异及性状间关系。结果表明：(1)群落内叶片功能性状种内、种间差异显著,变异中等(CV： 0.02-0.59),其结构性状的可塑性较化学性状保守,变异格局符合"性状空间变异分割假说";针叶树种的LA、SLA显著低于阔叶树种,常绿树种的LC和LDMC最小,而落叶树种的SLA、LN和LP最大以及LT和LC ∶ LN最小。(2)群落叶片功能性状变异主要来源于生活型和种间变异,种内变异亦有显著贡献;生活型对多数性状的贡献率较大,其对LDMC、LC ∶ LN的贡献分别高达93.11%和91.76%;种间变异(LDMC除外)对结构性状的贡献率均高于化学性状;种内变异对LP的贡献率(23.66%)较种间变异高。(3)叶片性状之间多表现出显著相关关系,针叶树和阔叶树在PCA排序图中聚集于不同区域,叶经济型谱在柯-青冈群落中具有适用性。不同树种通过叶片结构、化学性状之间的权衡策略来适应环境变化,从而实现群落物种共存。结果可为理解森林群落物种的环境适应策略、预测群落动态变化和制定植被恢复措施提供科学依据。     

叶脉网络功能性状及其生态学意义

叶脉网络结构是叶脉系统在叶片里的分布和排列样式。早期叶脉网络结构研究主要集中在其分类学意义上; 近年来叶脉网络功能性状及其在植物水分利用上的意义已成为植物生态学研究的热点。该文介绍了叶脉网络功能性状的指标体系(包括叶脉密度、叶脉直径、叶脉之间的距离、叶脉闭合度等), 综述了叶脉网络功能性状与叶脉系统功能(包括水分、养分和光合产物等物质运输、机械支撑和虫害防御等)的关系, 叶脉网络功能性状与叶片其他功能性状(包括比叶重、叶寿命、光合速率、叶片大小、气孔密度等)的协同变异和权衡关系, 以及叶脉网络功能性状随环境因子(包括水分、温度、光照等)的变化规律等方面的最新研究进展。此外, 叶脉网络功能性状的研究成果也被应用于古环境重建、城市交通规划、流域规划及全球变化研究中。由于叶脉网络功能性状是环境因子与系统发育共同作用的结果, 未来开展分子—叶片—植物—生态系统等多尺度的叶脉网络功能性状研究, 理清叶脉网络功能性状与气孔失水—茎干导水—根系吸水等植物水分利用的关系, 将为预测植物及生态系统对全球变化的响应提供新的启示。     

了哥王叶功能性状特征及其对土壤因子的响应

为探究了哥王Wikstroemiaindica的叶功能性状特征及其影响因素,在海岛植被调查的基础上对了哥王叶片进行取样并测定其功能性状指标,利用变异系数法和Pearson相关性分析探讨叶功能性状之间的差异与联系,运用冗余分析研究了哥王叶功能性状对土壤因子的响应。结果表明,了哥王的叶功能性状变异系数介于9.76%～23.73%,其中叶体积变异幅度最大(23.73%),叶干物质含量变异幅度最小(9.76%),整体上了哥王叶功能性状保持相对稳定。了哥王各项叶功能性状之间具有一定的相关性,联系较密切。了哥王叶功能性状主要受土壤中有机质、全氮、碱解氮的影响,土壤中有机质、全氮、碱解氮的含量与比叶面积呈正比,与叶厚度、叶体积成反比。了哥王的叶片可以通过一定的性状变异和组合来适应外部环境的变化,以较好地适应海岛恶劣的环境。该研究结果可为了哥王野生种质资源的保护、利用以及人工栽培提供参考。     

Impact of drought on productivity and water use efficiency in 29 genotypes of Populus deltoides x Populus nigra

We examined the relationships among productivity, water use efficiency (WUE) and drought tolerance in 29 genotypes of Populus x euramericana (Populus deltoides x Populus nigra), and investigated whether some leaf traits could be used as predictors for productivity, WUE and drought tolerance. At Orléans, France, drought was induced on one field plot by withholding water, while a second plot remained irrigated and was used as a control. Recorded variables included stem traits (e.g. biomass) and leaf structural (e.g. leaf area) and functional traits [e.g. intrinsic water use efficiency (Wi) and carbon isotope discrimination (Delta)]. Productivity and Delta displayed large genotypic variability and were not correlated. Delta scaled negatively with Wi and positively with stomatal conductance under moderate drought, suggesting that the diversity for Delta was mainly driven by stomatal conductance. Most of the productive genotypes displayed a low level of drought tolerance (i.e. a large reduction of biomass), while the less productive genotypes presented a large range of drought tolerance. The ability to increase WUE in response to water deficit was necessary but not sufficient to explain the genotypic diversity of drought tolerance.     

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.