Stomatal Density and Bio-water Saving

Bio-water saving is to increase water use efficiency of crops or crop yield per unit of water input. Plant water use efficiency is determined by photosynthesis and transpiration, for both of which stomata are crucial. Stomata are pores on leaf epidermis for both water and carbon dioxide fluxes that are controlled by two major factors: stomatal behavior and density. Stomatal behavior has been the focus of intensive research, while less attention has been paid to stomatal density. Recently, a number of genes controlling stomatal development have been identified. This review summarizes the recent progress on the genes regulating stomatal density, and discusses the role of stomatal density in plant water use efficiency and the possibility to increase plant water use efficiency, hence bio-water saving by genetically manipulating stomatal density.     

Gas exchange and water‐use efficiency in plant canopies

In this review, I first address the basics of gas exchange, water‐use efficiency and carbon isotope discrimination in C₃ plant canopies. I then present a case study of water‐use efficiency in northern Australian tree species. In general, C₃ plants face a trade‐off whereby increasing stomatal conductance for a given set of conditions will result in a higher CO₂ assimilation rate, but a lower photosynthetic water‐use efficiency. A common garden experiment suggested that tree species which are able to establish and grow in drier parts of northern Australia have a capacity to use water rapidly when it is available through high stomatal conductance, but that they do so at the expense of low water‐use efficiency. This may explain why community‐level carbon isotope discrimination does not decrease as steeply with decreasing rainfall on the North Australian Tropical Transect as has been observed on some other precipitation gradients. Next, I discuss changes in water‐use efficiency that take place during leaf expansion in C₃ plant leaves. Leaf phenology has recently been recognised as a significant driver of canopy gas exchange in evergreen forest canopies, and leaf expansion involves changes in both photosynthetic capacity and water‐use efficiency. Following this, I discuss the role of woody tissue respiration in canopy gas exchange and how photosynthetic refixation of respired CO₂ can increase whole‐plant water‐use efficiency. Finally, I discuss the role of water‐use efficiency in driving terrestrial plant responses to global change, especially the rising concentration of atmospheric CO₂. In coming decades, increases in plant water‐use efficiency caused by rising CO₂ are likely to partially mitigate impacts on plants of drought stress caused by global warming.     

Stomatal characteristics of ferns and angiosperms and their responses to changing light intensity at different habitats

气孔是植物与大气环境进行气体交换的重要通道, 在调控植物碳水平衡方面发挥着重要作用。为探讨生境和植物类型对气孔形态特征的影响以及气孔对光强变化的响应格局在不同植物间和不同生境条件下的变异, 选取开阔生境和林下生境的5种蕨类植物和4种被子植物, 测定了它们的气孔形态特征和气孔导度对光强变化的响应。此外, 还收集了8篇文献中开阔和林下生境的45种蕨类植物和70种被子植物的气孔密度和气孔长度数据, 以增大样本量从而更好地探讨不同生境条件下蕨类和被子植物气孔密度及长度的变异格局, 并通过分析生境和植物类型对气孔形态特征的影响来推测生境和植物类型对气孔响应行为的可能影响。实验结果表明, 与林下植物相比, 开阔环境下的植物气孔密度更大, 气孔长度更小, 气孔对光强降低的响应更敏感; 但植物类型对气孔形态特征的影响以及对气孔响应光强的敏感程度的影响均不显著。对文献数据的分析表明, 生境和植物类型对气孔形态特征均有显著影响。考虑到气孔响应快慢与气孔形态特征密切相关, 与蕨类植物相比, 被子植物小而密的气孔可能为其更快地响应环境变化提供了基础。研究表明生境和植物类型对气孔响应行为均有显著影响。     

不同生境条件下蕨类和被子植物的气孔形态特征及其对光强变化的响应

气孔是植物与大气环境进行气体交换的重要通道, 在调控植物碳水平衡方面发挥着重要作用。为探讨生境和植物类型对气孔形态特征的影响以及气孔对光强变化的响应格局在不同植物间和不同生境条件下的变异, 选取开阔生境和林下生境的5种蕨类植物和4种被子植物, 测定了它们的气孔形态特征和气孔导度对光强变化的响应。此外, 还收集了8篇文献中开阔和林下生境的45种蕨类植物和70种被子植物的气孔密度和气孔长度数据, 以增大样本量从而更好地探讨不同生境条件下蕨类和被子植物气孔密度及长度的变异格局, 并通过分析生境和植物类型对气孔形态特征的影响来推测生境和植物类型对气孔响应行为的可能影响。实验结果表明, 与林下植物相比, 开阔环境下的植物气孔密度更大, 气孔长度更小, 气孔对光强降低的响应更敏感; 但植物类型对气孔形态特征的影响以及对气孔响应光强的敏感程度的影响均不显著。对文献数据的分析表明, 生境和植物类型对气孔形态特征均有显著影响。考虑到气孔响应快慢与气孔形态特征密切相关, 与蕨类植物相比, 被子植物小而密的气孔可能为其更快地响应环境变化提供了基础。研究表明生境和植物类型对气孔响应行为均有显著影响。     

基于遥感图像处理技术胡杨叶气孔密度的估算及其生态意义

拟利用遥感图像处理技术--面向对象分类,计算胡杨叶片气孔密度,采用面向对象分类的专业软件eCognition对气孔图像进行多尺度分割,将生成的分类图像导入ArcGIS中计算气孔密度,最后用R语言编写代码进行批处理。研究结果显示:该方法用于计算叶片气孔的密度精度高;18个样点胡杨气孔密度存在着较大的差异,从76.7 个/mm²到139.4 个/mm²不等,其平均密度为105 个/mm²;随着干旱胁迫加强,气孔密度表现下降上升再下降的趋势。     

不同尺度上植物叶气孔导度对升高CO2的响应

植物叶气孔导度对大气CO2浓度升高的响应可表现在以下几个层面：在叶水平上,叶气孔导度和气孔密度下降;在植物个体水平上,单位叶面积蒸腾下降,植株的水分利用率升高;在生态系统水平上,蒸散降低,土壤泾流和土壤水分含量增加;在全球尺度上,扩大了温室气体的增温效应,同时也降低了全球降雨量增加的趋势。正是因为植物叶气孔导度的变化会影响全球水循环,所以它在全球变化中起着非常重要的作用。但目前的研究结果还不能外推到更大的尺度上去。     

Optimal stomatal behavior theory for simulating stomatal conductance

Among the most critical processes in simulating terrestrial ecosystem performance is the regulatory role of stomata in carbon and water cycles. Compared with field measurements, the changes in stomatal slope caused by the biophysical environment provide a simple but effective synthetic framework for studying climate-related carbon and water cycling, due to its sensitivity to CO₂, vapor pressure deficit, and photosynthesis. It is also crucial in understanding the effects of climate change on photosynthesis and water use efficiency. Endeavored by numerous scholastic efforts, stomatal conductance models have been improved based on experimental, semi-experimental, and mechanical processes. However, the underlying biological mechanisms and the dynamics of key parameters in these models remain unexplored, especially regarding the changes in stomatal slope. By improving the understanding of the stomata’s regulatory role, we reduced the uncertainty of stomatal conductance simulation. We then synthesized the recent developments and lessons in optimal stomatal behavior theory to simulate stomatal conductance and included an introduction to widely used stomatal conductance models and parameters, the main factors influencing stomatal slopes, and applications of the mechanical stomatal conductance models in different ecosystems. Based on our literature review, we proposed that future research is needed on the optimal stomatal behavior theory and its applications in simulating stomatal conductance.     

最优气孔行为理论和气孔导度模拟

气孔调节功能是陆地生态系统碳-水耦合过程中最重要的环节。与即时的气孔导度测量相比, 气孔导度斜率能有效地反映气孔导度对CO₂浓度、饱和水汽压亏缺和光合作用的敏感性, 包含了环境因子对光合作用和临界水分利用效率等的综合影响, 为研究全球变化下陆地生态系统碳-水耦合关系提供了一个简明且综合的框架。气孔导度模型从经验模型、半经验模型发展到机理模型, 经过很多学者的改进, 但是模型参数的生物学意义和变化规律还不明确。鉴于气孔导度斜率方面研究的重要性和研究的不足, 为了加强对气孔导度调节规律的认识, 并减少气孔导度模拟的不确定性, 该文主要综述了长期以来国内外关于最优气孔行为理论和气孔导度模拟方面的研究成果, 其中包括广泛使用的气孔导度模型及参数意义, 讨论影响气孔导度斜率的主要因素以及气孔导度机理模型的应用, 并对最优气孔行为理论和气孔导度模拟方面的研究做了简单展望。     

Correlated evolution of stem and leaf hydraulic traits in Pereskia (Cactaceae)

Recent studies have demonstrated significant correlations between stem and leaf hydraulic properties when comparing across species within ecological communities. This implies that these traits are co-evolving, but there have been few studies addressing plant water relations within an explicitly evolutionary framework. This study tests for correlated evolution among a suite of plant water-use traits and environmental parameters in seven species of Pereskia (Cactaceae), using phylogenetically independent contrasts. There were significant evolutionary correlations between leaf-specific xylem hydraulic conductivity, Huber Value, leaf stomatal pore index, leaf venation density and leaf size, but none of these traits appeared to be correlated with environmental water availability; only two water relations traits - mid-day leaf water potentials and photosynthetic water use efficiency - correlated with estimates of moisture regime. In Pereskia, it appears that many stem and leaf hydraulic properties thought to be critical to whole-plant water use have not evolved in response to habitat shifts in water availability. This may be because of the extremely conservative stomatal behavior and particular rooting strategy demonstrated by all Pereskia species investigated. These results highlight the need for a lineage-based approach to understand the relative roles of functional traits in ecological adaptation.     

Evidence of changing intrinsic water‐use efficiency under rising atmospheric CO2 concentrations in Boreal Fennoscandia from subfossil leaves and tree ring δ13C ratios

Investigating the many internal feedbacks within the climate system is a vital component of the effort to quantify the full effects of future anthropogenic climate change. The stomatal apertures of plants tend to close and decrease in number under elevated CO₂ concentrations, increasing water‐use efficiency (WUE) and reducing canopy evapotranspiration. Experimental and modelling studies reveal huge variations in these changes such that the warming associated with reduced evapotranspiration (known as physiological forcing) is neither well understood or constrained. Palaeo‐observations of changes in stomatal response and plant WUE under rising CO₂ might be used to better understand the processes underlying the physiological forcing feedback and to link measured changes in plant WUE to a specific physiological change in stomata. Here we use time series of tree ring (Pinus sylvestris L.) δ¹³C and subfossil leaf (Betula nana L.) measurements of stomatal density and geometry to derive records of changes in intrinsic water‐use efficiency (iWUE) and maximum stomatal conductance in the Boreal zone of northern Finland and Sweden. We investigate the rate of change in both proxies, over the recent past. The independent lines of evidence from these two different Boreal species indicate increased iWUE and reduced maximum stomatal conductance of similar magnitude from preindustrial times (ca. ad 1850) to around ad 1970. After this maximum stomatal conductance continues to decrease to ad 2000 in B. nana but iWUE in P. sylvestris reaches a plateau. We suggest that northern boreal P. sylvestris might have reached a threshold in its ability to increase WUE as CO₂ rises.     

Regulation Mechanisms of Stomatal Oscillation

Stomata function as the gates between the plant and the atmospheric environment. Stomatal movement, including stomatal opening and closing, controls CO2 absorption as the raw material for photosynthesis and water loss through transpiration. How to reduce water loss and maintain enough CO2 absorption has been an interesting research topic for some time. Simple stomatal opening may elevate CO2 absorption, but, in the meantime, promote the water loss, whereas simple closing of stomatal pores may reduce both water loss and CO2 absorption, resulting in impairment of plant photosynthesis. Both processes are not economical to the plant. As a special rhythmic stomatal movement that usually occurs at smaller stomatal apertures, stomatal oscillation can keep CO2 absorption at a sufficient level and reduce water loss at the same time, suggesting a potential improvement in water use efficiency. Stomatal oscillation is usually found after a sudden change in one environmental factor in relatively constant environments. Many environmental stimuli can induce stomatal oscillation. It appears that, at the physiological level, feedback controls are involved in stomatal oscillation. At the cellular level, possibly two different patterns exist： （i） a quicker responsive pattern; and （ii） a slower response. Both involve water potential changes and water channel regulation, but the mechanisms of regulation of the two patterns are different. Some evidence suggests that the regulation of water channels may play a vital and primary role in stomatal oscillation. The present review summarizes studies on stomatal oscillation and concludes with some discussion regarding the mechanisms of regulation of stomatal oscillation.     

Allometric co-variation of xylem and stomata across diverse woody seedlings

Leaf stomatal density is known to co-vary with leaf vein density. However, the functional underpinning of this relation, and how it scales to whole-plant water transport anatomy, is still unresolved. We hypothesized that the balance of water exchange between the vapour phase (in stomata) and liquid phase (in vessels) depends on the consistent scaling between the summed stomatal areas and xylem cross-sectional areas, both at the whole-plant and single-leaf level. This predicted size co-variation should be driven by the co-variation of numbers of stomata and terminal vessels. We examined the relationships of stomatal traits and xylem anatomical traits from the entire plant to individual leaves across seedlings of 53 European woody angiosperm species. There was strong and convergent scaling between total stomatal area and stem xylem area per plant and between leaf total stomatal area and midvein xylem area per leaf across all the species, irrespective of variation in leaf habit, growth-form or relative growth rate. Moreover, strong scaling was found between stomatal number and terminal vessel number, whereas not in their respective average areas. Our findings have broad implications for integrating xylem architecture and stomatal distribution and deepen our understanding of the design rules of plants' water transport network.     

Specialized stomatal humidity responses underpin ecological diversity in C3 bromeliads

The Neotropical Bromeliaceae display an extraordinary level of ecological variety, with species differing widely in habit, photosynthetic pathway and growth form. Divergences in stomatal structure and function, hitherto understudied in treatments of bromeliad evolutionary physiology, could have been critical to the generation of variety in ecophysiological strategies among the bromeliads. Because humidity is a key factor in bromeliad niches, we focussed on stomatal responses to vapour pressure deficit (VPD). We measured the sensitivity of stomatal conductance and assimilation rate to VPD in eight C₃ bromeliad species of contrasting growth forms and ecophysiological strategies and parameterised the kinetics of stomatal responses to a step change in VPD. Notably, three tank‐epiphyte species displayed low conductance, high sensitivity and fast kinetics relative to the lithophytes, while three xeromorphic terrestrial species showed high conductance and sensitivity but slow stomatal kinetics. An apparent feedforward response of transpiration to VPD occurred in the tank epiphytes, while water‐use efficiency was differentially impacted by stomatal closure depending on photosynthetic responses. Differences in stomatal responses to VPD between species of different ecophysiological strategies are closely linked to modifications of stomatal morphology, which we argue has been a pivotal component of the evolution of high diversity in this important plant family.     

Stomatal function,density and pattern,and CO2 assimilation in Arabidopsis thaliana tmm1 and sdd1‐1 mutants

• Stomata modulate the exchange of water and CO₂ between plant and atmosphere. Although stomatal density is known to affect CO₂ diffusion into the leaf and thus photosynthetic rate, the effect of stomatal density and patterning on CO₂ assimilation is not fully understood.
• We used wild types Col‐0 and C24 and stomatal mutants sdd1‐1 and tmm1 of Arabidopsis thaliana, differing in stomatal density and pattern, to study the effects of these variations on both stomatal and mesophyll conductance and CO₂ assimilation rate. Anatomical parameters of stomata, leaf temperature and carbon isotope discrimination were also assessed.
• Our results indicate that increased stomatal density enhanced stomatal conductance in sdd1‐1 plants, with no effect on photosynthesis, due to both unchanged photosynthetic capacity and decreased mesophyll conductance. Clustering (abnormal patterning formed by clusters of two or more stomata) and a highly unequal distribution of stomata between the adaxial and abaxial leaf sides in tmm1 mutants also had no effect on photosynthesis.
• Except at very high stomatal densities, stomatal conductance and water loss were proportional to stomatal density. Stomatal formation in clusters reduced stomatal dynamics and their operational range as well as the efficiency of CO₂ transport.

     

Relating Stomatal Conductance to Leaf Functional Traits

Leaf functional traits are important because they reflect physiological functions, such as transpiration and carbon assimilation. In particular, morphological leaf traits have the potential to summarize plants strategies in terms of water use efficiency, growth pattern and nutrient use. The leaf economics spectrum (LES) is a recognized framework in functional plant ecology and reflects a gradient of increasing specific leaf area (SLA), leaf nitrogen, phosphorus and cation content, and decreasing leaf dry matter content (LDMC) and carbon nitrogen ratio (CN). The LES describes different strategies ranging from that of short-lived leaves with high photosynthetic capacity per leaf mass to long-lived leaves with low mass-based carbon assimilation rates. However, traits that are not included in the LES might provide additional information on the species'' physiology, such as those related to stomatal control. Protocols are presented for a wide range of leaf functional traits, including traits of the LES, but also traits that are independent of the LES. In particular, a new method is introduced that relates the plants’ regulatory behavior in stomatal conductance to vapor pressure deficit. The resulting parameters of stomatal regulation can then be compared to the LES and other plant functional traits. The results show that functional leaf traits of the LES were also valid predictors for the parameters of stomatal regulation. For example, leaf carbon concentration was positively related to the vapor pressure deficit (vpd) at the point of inflection and the maximum of the conductance-vpd curve. However, traits that are not included in the LES added information in explaining parameters of stomatal control: the vpd at the point of inflection of the conductance-vpd curve was lower for species with higher stomatal density and higher stomatal index. Overall, stomata and vein traits were more powerful predictors for explaining stomatal regulation than traits used in the LES.     

遮阴和干旱对白桦幼苗光诱导的气孔动力学影响

以黄土丘陵区演替早期种白桦的幼龄实生苗为材料,采用全光照和遮阴(光照为全光照的30%)以及正常水分(田间持水量的75%～80%)和干旱(田间持水量的40%～45%)处理,研究了遮阴和干旱下白桦幼苗光诱导的气孔动力学参数、气孔解剖特征、整株植物生长和水分利用的变化。结果表明: 遮阴使气孔开放过程的滞后时间和响应时间显著增加0.8和1.8倍,气孔开放和关闭过程的响应速度显著降低82.2%和65.0%,气孔开放和关闭过程的响应幅度显著降低43.3%和56.9%;干旱使气孔开放过程的响应幅度显著降低43.9%、气孔关闭过程的响应速度显著降低33.0%;二者仅对气孔开放过程的滞后时间存在交互作用。白桦幼苗气孔关闭过程的响应速度与气孔密度和气孔指数呈显著正相关,其他气孔动力学参数与气孔解剖结构间无显著相关性。白桦气孔关闭过程中的响应速度与整株生物量、耗水量呈显著正相关,所有气孔动力学参数和整株水分利用效率间无显著相关性。遮阴和干旱对白桦光诱导的气孔动力学参数的影响与其对气孔解剖结构的影响有关,动态光下的气孔动力学参数可在一定程度上阐释白桦不同生境下生长的差异。     

不同生境下入侵种假臭草叶片的气孔特征

【背景】探讨入侵种假臭草不同生境下气孔的变化规律,揭示假臭草种群在不同生境下所采取的生长对策及适应机制,可为入侵生物的防治提供参考。【方法】采取光学显微镜系统观察桉树林、木薯地、弃耕地、公路边4种生境下假臭草叶片的气孔特征。【结果】光照和土壤肥、水条件对假臭草叶片的气孔孔径（横轴方向和纵轴方向）、单个气孔器面积、气孔器总面积、气孔密度及气孔指数的影响显著。低光照及肥沃、湿润土壤生境与高光照及贫瘠、干旱土壤相比,假臭草的气孔孔径（横轴方向和纵轴方向）、单个气孔器面积、气孔器总面积较大,气孔密度及气孔指数较小。【结论与意义】假臭草叶片气孔特征表现可塑性,说明其对异质环境具有一定的生态适应能力。     

气孔参数与大气CO2浓度的相关性及其影响因素

通常认为气孔参数(气孔密度和气孔指数)和大气CO2浓度有负相关关系,但不是每种植物的气孔参数都与CO2浓度的变化有负相关关系,气孔参数对大气CO2浓度的显著反应也只在一定的CO2浓度范围内发生。大气CO2浓度是影响气孔参数变化的主要因素,同时温度、水分的供应和光照条件等其它环境因素也影响气孔参数。CO2浓度和光照条件主要影响气孔发生,而其它环境因素主要影响叶片表皮细胞的大小。气孔指数部分消除了表皮细胞大小带来的影响,用气孔指数指示大气CO2浓度比用气孔密度指示更为可靠。     

Reconciling the optimal and empirical approaches to modelling stomatal conductance

Models of vegetation function are widely used to predict the effects of climate change on carbon, water and nutrient cycles of terrestrial ecosystems, and their feedbacks to climate. Stomatal conductance, the process that governs plant water use and carbon uptake, is fundamental to such models. In this paper, we reconcile two long‐standing theories of stomatal conductance. The empirical approach, which is most commonly used in vegetation models, is phenomenological, based on experimental observations of stomatal behaviour in response to environmental conditions. The optimal approach is based on the theoretical argument that stomata should act to minimize the amount of water used per unit carbon gained. We reconcile these two approaches by showing that the theory of optimal stomatal conductance can be used to derive a model of stomatal conductance that is closely analogous to the empirical models. Consequently, we obtain a unified stomatal model which has a similar form to existing empirical models, but which now provides a theoretical interpretation for model parameter values. The key model parameter, g₁, is predicted to increase with growth temperature and with the marginal water cost of carbon gain. The new model is fitted to a range of datasets ranging from tropical to boreal trees. The parameter g₁ is shown to vary with growth temperature, as predicted, and also with plant functional type. The model is shown to correctly capture responses of stomatal conductance to changing atmospheric CO₂, and thus can be used to test for stomatal acclimation to elevated CO₂. The reconciliation of the optimal and empirical approaches to modelling stomatal conductance is important for global change biology because it provides a simple theoretical framework for analyzing, and simulating, the coupling between carbon and water cycles under environmental change.     

Regulation of Transpiration to Improve Crop Water Use

Decreasing fresh water supplies and increasing agricultural drought threaten sustainable worldwide crop production. Consequently, there is a global priority to develop crops with higher water use efficiency (WUE): biomass production or yield per unit of water used. Water use efficiency varies substantially among species and genotypes within a species, and a major effort is now underway to identify the genetic determinants of WUE. Today, it is known that genotypes in primary gene pools exhibit allelic variation for WUE through mechanisms that regulate transpiration, which is the conductance of water through stomata, the cuticle, and the boundary layer. Because of the differential diffusion properties of water and carbon dioxide (CO₂) through these pathways, it is feasible that WUE could be improved by decreasing transpiration without a concomitant reduction in CO₂ uptake. Since CO₂ uptake and transpirational water loss occur predominantly through stomatal pores, it is not surprising that genes involved in stomatal development and stomatal opening/closing impact WUE. Furthermore, loss- and gain-of-function genetic screens have identified genes that regulate transpiration and WUE by yet undetermined mechanisms. This review will discuss the genetic determinants that regulate transpiration and WUE in the context of the modern agricultural goal of improving WUE while sustaining biomass and yield.