Steady-state stomatal responses of C3 and C4 species to blue light fraction: Interactions with CO2 concentration

Blue light induced stomatal opening has been studied by applying a short pulse (~5 to 60 s) of blue light to a background of saturating photosynthetic red photons, but little is known about steady-state stomatal responses. Here we report stomatal responses to blue light at high and low CO₂ concentrations. Steady-state stomatal conductance (g_s) of C₃ plants increased asymptotically with increasing blue light to a maximum at 20% blue (120 μmol m⁻² s⁻¹). This response was consistent from 200 to 800 μmol mol⁻¹ atmospheric CO₂ (C_a). In contrast, blue light induced only a transient stomatal opening (~5 min) in C₄ species above a C_a of 400 μmol mol⁻¹. Steady-state g_s of C₄ plants generally decreased with increasing blue intensity. The net photosynthetic rate of all species decreased above 20% blue because blue photons have lower quantum yield (moles carbon fixed per mole photons absorbed) than red photons. Our findings indicate that photosynthesis, rather than a blue light signal, plays a dominant role in stomatal regulation in C₄ species. Additionally, we found that blue light affected only stomata on the illuminated side of the leaf. Contrary to widely held belief, the blue light-induced stomatal opening minimally enhanced photosynthesis and consistently decreased water use efficiency.     

Oscillations in stomatal conductance of hybrid poplar leaves in the light and dark

Cycling of stomatal conductance in three hybrid poplar ( Populus sp.) cultivars was observed under a variety of conditions. Illumination of plants kept previously in the dark induced very large oscillations with a period of about 40 min and large oscillations with a shorter period (< 10 min) were superimposed on the longer cycles. During these oscillations, large changes in conductance could occur very rapidly (1.0 cm s⁻¹ in 3 min). Plants in constant light also displayed both long and short term cycles in conductance, but these were smaller in amplitude than those induced by sudden illumination. Stomatal oscillations were also observed in darkness and after darkening of previously illuminated plants. These oscillations had shorter (< 30 min) and less regular periods than those observed in the light. Such cycling in the dark is rare. Cycling of the two leaf surfaces was sometimes in synchrony in the light, and more so after a perturbation. Little synchrony between the two surfaces was observed in the dark. Stomatal movements of different leaves on a plant were usually relatively independent. Transient stomatal opening occurred following leaf excision in the light or dark, and often after sudden darkening of intact leaves. Also, stomata of intact leaves sometimes transiently closed following illumination.     

Abaxial and adaxial stomata from Pima cotton (Gossypium barbadense L.) differ in their pigment content and sensitivity to light quality

Sensitivity to light quality and pigment composition were analysed and compared in abaxial and adaxial stomata of Gossypium barbadense L. (Pima cotton). In most plants, abaxial (lower) stomatal conductances are higher than adaxial (upper) ones, and stomatal opening is more sensitive to blue light than to red. In greenhouse-grown Pima cotton, abaxial stomatal conductances were two to three times higher than adaxial ones. In contrast, adaxial stomatal conductances were 1·5 to two times higher than abaxial ones in leaves from growth chamber-grown plants. To establish whether light quality was a factor in the regulation of the relationship between abaxial and adaxial stomatal conductances, growth-chamber-grown plants were exposed to solar radiation outdoors and to increased red light in the growth chamber. In both cases, the ratios of adaxial to abaxial stomatal conductance reverted to those typical of greenhouse plants. We investigated the hypothesis that adaxial stomata are more sensitive to blue light and abaxial stomata are more sensitive to red light. Measurements of stomatal apertures in mechanically isolated epidermal peels from growth chamber and greenhouse plants showed that adaxial stomata opened more under blue light than under red light, while abaxial stomata had the opposite response. Using HPLC, we quantified the chlorophylls and carotenoids extracted from isolated adaxial and abaxial guard cells. All pigments analysed were more abundant in the adaxial than in the abaxial guard cells. Antheraxanthin and β-carotene contents were 2·3 times higher in adaxial than in abaxial guard cells, comparing with ad/ab ratios of 1·5–1·9 for the other pigments. We conclude that adaxial and abaxial stomata from Pima cotton have a differential sensitivity to light quality and their distinct responses are correlated with different pigment content.     

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.     

The response of stomatal conductance to seasonal drought in tropical forests

Stomata regulate CO₂ uptake for photosynthesis and water loss through transpiration. The approaches used to represent stomatal conductance (g_s) in models vary. In particular, current understanding of drivers of the variation in a key parameter in those models, the slope parameter (i.e. a measure of intrinsic plant water‐use‐efficiency), is still limited, particularly in the tropics. Here we collected diurnal measurements of leaf gas exchange and leaf water potential (Ψ_leaf), and a suite of plant traits from the upper canopy of 15 tropical trees in two contrasting Panamanian forests throughout the dry season of the 2016 El Niño. The plant traits included wood density, leaf‐mass‐per‐area (LMA), leaf carboxylation capacity (V_c,max), leaf water content, the degree of isohydry, and predawn Ψ_leaf. We first investigated how the choice of four commonly used leaf‐level g_s models with and without the inclusion of Ψ_leaf as an additional predictor variable influence the ability to predict g_s, and then explored the abiotic (i.e. month, site‐month interaction) and biotic (i.e. tree‐species‐specific characteristics) drivers of slope parameter variation. Our results show that the inclusion of Ψ_leaf did not improve model performance and that the models that represent the response of g_s to vapor pressure deficit performed better than corresponding models that respond to relative humidity. Within each g_s model, we found large variation in the slope parameter, and this variation was attributable to the biotic driver, rather than abiotic drivers. We further investigated potential relationships between the slope parameter and the six available plant traits mentioned above, and found that only one trait, LMA, had a significant correlation with the slope parameter (R² = 0.66, n = 15), highlighting a potential path towards improved model parameterization. This study advances understanding of g_s dynamics over seasonal drought, and identifies a practical, trait‐based approach to improve modeling of carbon and water exchange in tropical forests.     

Distinct responses to ozone of abaxial and adaxial stomata in three Euramerican poplar genotypes

Ozone induces stomatal sluggishness, which impacts photosynthesis and transpiration. Stomatal responses to variation of environmental parameters are slowed and reduced by ozone and may be linked to difference of ozone sensitivity. Here we determine the ozone effects on stomatal conductance of each leaf surface. Potential causes of this sluggish movement, such as ultrastructural or ionic fluxes modification, were studied independently on both leaf surfaces of three Euramerican poplar genotypes differing in ozone sensitivity and in stomatal behaviour. The element contents in guard cells were linked to the gene expression of ion channels and transporters involved in stomatal movements, directly in microdissected stomata. In response to ozone, we found a decrease in the stomatal conductance of the leaf adaxial surface correlated with high calcium content in guard cells compared with a slight decrease on the abaxial surface. No ultrastructural modifications of stomata were shown except an increase in the number of mitochondria. The expression of vacuolar H⁺/Ca²⁺‐antiports (CAX1 and CAX3 homologs), β‐carbonic anhydrases (βCA1 and βCA4) and proton H⁺‐ATPase (AHA11) genes was strongly decreased under ozone treatment. The sensitive genotype characterized by constitutive slow stomatal response was also characterized by constitutive low expression of genes encoding vacuolar H⁺/Ca²⁺‐antiports.     

Genetic variation of stomatal conductance, blue light sensitivity and zeaxanthin content in guard cells of Pima cotton (Gossypium barbadense)

Pima S‐6 ( Gossypium barbadense L.) is a modern line with high stomatal conductance, while B368 is a primitive cotton with low conductance. The blue light sensitivity of adaxial guard cells, probed as the blue light‐dependent enhancement of the red light‐induced chlorophyll a fluorescence quenching, was investigated in these two cotton lines with contrasting stomatal conductance. Adaxial guard cells isolated from Pima S‐6 cotton plants had a significantly higher carotenoid content and a higher blue light sensitivity than those isolated from B368 plants. In a growth chamber‐grown F₂ population of a cross between these two lines, adaxial stomatal conductances of individual plants segregated over a range exceeding the average conductances of the parents. Carotenoid content and the blue light sensitivity of adaxial guard cells also segregated. The concentrations of xanthophylls and β‐carotene in the adaxial guard cells were poorly correlated with the blue light response, except for zeaxanthin. The co‐segregation of stomatal conductance and blue light sensitivity suggested that the stomatal response to blue light may play a role in the regulation of stomatal conductance in the intact leaf. Zeaxanthin content and blue light sensitivity also co‐segregated, suggesting that both parameters are under genetic control. The co‐segregation of zeaxanthin content, blue light sensitivity and stomatal conductance provides further evidence for a role of zeaxanthin in the blue light photoreception of guard cells.     

Comparing model predictions and experimental data for the response of stomatal conductance and guard cell turgor to manipulations of cuticular conductance, leaf-to-air vapour pressure difference and temperature: feedback mechanisms are able to account for all observations

Stomata respond to increasing leaf-to-air vapour pressure difference (LAVPD) ( D ) by closing. The mechanism by which this occurs is debated. A role for feedback and peristomatal transpiration has been proposed. In this paper, we apply a recent mechanistic model of stomatal behaviour, and compare model and experimental data for the influence of increasing D on stomatal conductance.
We manipulated cuticular conductance ( g _c) by three independent methods. First, we increased g _c by using a solvent mixture applied to both leaf surfaces prior to determining stomatal responses to D ; second, we increased g _c by increasing leaf temperature at constant D ; and third, we coated a small area of leaf with a light oil to decrease g _c. In all three experiments, experimental data and model outputs showed very close agreement.
We conclude, from the close agreement between model and experimental data and the fact that manipulations of g _c, and hence cuticular transpiration, influenced g _s in ways consistent with a feedback mechanism, that feedback is central in determining stomatal responses to D .     

Asymmetric responses of adaxial and abaxial stomata to elevated CO2: impacts on the control of gas exchange by leaves

The response of adaxial and abaxial stomatal conductance in Rumex obtusifolius to growth at elevated atmospheric concentrations of CO₂ (250 μmol mol^?1 above ambient) was investigated over two growing seasons. The conductance of both the adaxial and abaxial leaf surfaces was found to be reduced by elevated concentrations of CO₂. Elevated CO₂ caused a much greater reduction in conductance for the adaxial surface than for the abaxial surface. The absence of effects upon stomatal density indicated that the reductions were probably the result of changes in stomatal aperture. Partitioning of gas exchange between the leaf surfaces revealed that increased concentrations of CO₂ caused increased rates of photosynthesis only via the abaxial surface. Additionally, leaf thickness was found to increase during growth at elevated concentrations of CO₂. The tendency for these amphistomatous leaves to develop a distribution of conductance approaching that of hypostomatous leaves clearly reduced their maximum photosynthetic potential. This conclusion was supported by measurements of stomatal limitation, which showed greater values for the adaxial surfaces, and greater values at elevated CO₂. This reduction in photosynthesis may in part be caused by higher diffusive limitations imposed because of increased leaf thickness. In an uncoupled canopy, asymmetrical stomatal responses of the kind identified here may appreciably reduce transpiration. Species which show symmetrical responses are less likely to show reduced transpirational rates, and a redistribution of water loss between species may occur. The implications of asymmetrical stomatal responses for photosynthesis and canopy transpiration are discussed.     

Isolation of genes predominantly expressed in guard cells and epidermal cells of Nicotiana glauca

Guard cells are specialized and metabolically active cells which arise during the differentiation of the epidermis. Using Nicotiana glauca epidermal peels as a source of purified guard cells, we have constructed a cDNA library from guard cell RNA. In order to isolate genes that are predominantly expressed in guard cells, we performed a differential screen of this library, comparing the hybridization of a radiolabeled cDNA probe synthesized from guard cell RNA to that from a mesophyll cell cDNA probe. Sixteen clones were isolated based on their greater level of hybridization with the guard cell probe. Of these, eight had high homology to lipid transfer protein (LTP), two were similar to glycine-rich protein (GRP), and one displayed high homology to proline-rich proteins from Arabidopsis thaliana (AtPRP2, AtPRP4) and from potato guard cells (GPP). Northern analysis confirmed that one or more NgLTP genes, NgGRP1, and NgGPP1 are all differentially expressed, with highest levels in guard cells, and low or undetectable levels in mesophyll cells and in roots. In addition, all are induced to some degree in drought-stressed guard cells. NgLTP and NgGRP1 expression was localized by in situ hybridization to the guard cells and pavement cells in the epidermis. NgGRP1 expression was also detected in cells of the vasculature. Genomic Southern analysis indicated that LTP is encoded by a family of highly similar genes in N. glauca. This work has identified members of a subset of epidermis- and guard cell-predominant genes, whose protein products are likely to contribute to the unique properties acquired by guard cells and pavement cells during differentiation.     

Gas Exchange and Epidermal Characteristics of Miscanthus Populations in Taiwan Varying with Habitats and Nitrogen Application

Seventeen clones of C₄ grass Miscanthus spp. collected from different climatic regions and elevations of Taiwan were transplanted in pots. 15–16 months after collection the plants received 0, 1, and 2 g of nitrogen fertiliser (N₀, N₁, and N₂, respectively) per pot. All the measurements were done 10–12 d after N application. The relationships between net photosynthetic rate (P _N) and photon flux density (PFD) showed a saturated curve, with PFD saturation at about 1 000 µmol m^–2 s^–1. The ranges of PFD saturated P _N (P _sat) for all the tested clones with N₀, N₁, and N₂ were 8–16, 11–18, and 12–21 µmol m^–2 s^–1, respectively. The clones from southern Taiwan, a tropical region, showed the highest P _sat, followed by the clones from northern Taiwan, a subtropical region, while those from mountainous area showed the lowest P _sat. The clones collected from southern Taiwan showed the highest frequency of stomata on the adaxial surface, and those collected from the high mountainous area showed the lowest frequency. Also the adaxial surface of leaves from the higher mountainous area had more wax deposited than the leaves from the lowland. Thus the low P _sat in mountain clones is limited by both stomatal and non-stomatal factors. Further, the lower leaf conductance and different epidermal characteristics of mountain clones might prevent excessive loss of heat through transpiration and provide production against ultraviolet-B radiation.     

北京山区油松和元宝槭冠层气孔导度特征及其环境响应

叶片气孔是植物进行水汽交换的通道, 影响着植物的蒸腾和光合作用。然而叶片气孔行为受环境条件和树种类型的影响, 不同树种冠层气孔导度对环境因子响应的差异性, 以及在生长季不同时期叶片气孔对冠层蒸腾的调节作用是否会发生改变, 仍不清楚。该研究目的是通过探究各环境因子对不同树种冠层气孔导度的相对贡献率以及叶片气孔对冠层蒸腾的调节作用, 为深入了解植物水分利用状况和山区森林经营提供参考依据。于2018年生长季以北京八达岭国家森林公园内的58年生油松(Pinus tabuliformis)和39年生元宝槭(Acer truncatum)为研究对象, 利用热扩散技术对其树干液流进行连续监测, 并同步监测环境因子。利用彭曼公式计算冠层气孔导度(G_s)。主要结果: (1)油松和元宝槭日间G_s在日、月时间尺度上存在明显差异。5-7月油松和元宝槭日动态G_s均随饱和水汽压差(VPD)和太阳辐射(GR)的增加呈上升趋势, 上升持续时间比8月和9月长; 在月尺度上, 随着VPD、GR的降低和土壤湿度(VWC)的升高, G_s从5月到9月整体上升。(2)利用增强回归树法分析得到VWC和VPD对G_s的贡献率最大, 其次是GR、气温和风速。VWC和VPD对油松G_s的贡献率分别为66.4%和17.4%, 对元宝槭G_s的贡献率分别为54.8%和21.0%。(3)油松和元宝槭的dG_s/dlnVPD值与参考冠层气孔导度之间的斜率均显著高于0.6, 气孔调节作用相对较强。综上所述, 气孔对环境因子的响应在树种以及生长季不同时期之间存在差异, 为防止水分过度散失, 两树种在不同土壤水分条件下均通过严格的气孔调节控制蒸腾量。     

Sensitivity of infrared water vapor analyzers to oxygen concentration and errors in stomatal conductance

Use of infrared analyzers to measure water vapor concentrations in photosynthesis systems is becoming common. It is known that sensitivity of infrared carbon dioxide and water vapor analyzers is affected by the oxygen concentration in the background gas, particularly for absolute analyzers, but the potential for large errors in estimates of stomatal conductance due to effects of oxygen concentration on the sensitivity of infrared water vapor analyzers is not widely recognized. This work tested three types of infrared water vapor analyzers for changes in sensitivity of infrared water vapor analyzers depending on the oxygen content of the background gas. It was found that changing from either 0 or 2% to 21% oxygen in nitrogen decreased the sensitivity to water vapor for all three types of infrared water vapor analyzers by about 4%. The change in sensitivity was linear with oxygen mole fraction. The resulting error in calculated stomatal conductance would depend strongly on the leaf to air vapor pressure difference and leaf temperature, and also on whether leaf temperature was directly measured or calculated from energy balance. Examples of measurements of gas exchange on soybean leaves under glasshouse conditions indicated that changing from 21% to 2% oxygen produced an artifactual apparent increase in stomatal conductance which averaged about 30%. Similar errors occurred for `conductances' of wet filter paper. Such errors could affect inferences about the carbon dioxide dependence of the sensitivity of photosynthesis to oxygen. This revised version was published online in June 2006 with corrections to the Cover Date.     

Responses of stomatal conductance to light, humidity and temperature in winter wheat and barley grown at three concentrations of carbon dioxide in the field

Responses of leaf stomatal conductance to light, humidity and temperature were characterized for winter wheat and barely grown at ambient (about 350 μmol mol^?1 in the daytime), ambient + 175 and ambient + 350 μmol mol^?1 concentrations of carbon dioxide in open‐topped chambers in field plots over a three year period. Stomatal responses to environment were determined by direct manipulation of single environmental factors, and those results were compared with responses derived from natural day to day variation in mid‐day stomatal conductance. The purpose of these experiments was to determine the magnitude of reduction in stomatal conductance at elevated [CO₂], and to assess whether the relative response of conductance to elevated [CO₂] was constant across light, humidity and temperature conditions. The results indicated that light, humidity and temperature all significantly affected the relative decrease in stomatal conductance at elevated [CO₂]. The relative decrease in conductance with elevated [CO₂] was greater at low light, low water vapour pressure difference, and high temperature in both species. For measurements made at saturating light near mid‐day, the ratio of mid‐day stomatal conductances at doubled [CO₂] to that at ambient [CO₂] ranged from 0.42 to 0.86, with a mean of 0.66 in barley, and from 0.33 to 0.80, with a mean of 0.56 in wheat. Day‐to‐day variation in the relative effect of elevated [CO₂] on conductance was correlated with the relative stimulation of [CO₂] assimilation rate and with temperature. Some limitations of multiple linear regression, multiplicative, and ‘Ball–Berry' models as summaries of the data are discussed. In barley, a better fit to the models occurred in individual years than for the combined data, and in wheat a better fit to the models occurred when data from near the end of the season were removed.     

A model explaining genotypic and ontogenetic variation of leaf photosynthetic rate in rice (Oryza sativa) based on leaf nitrogen content and stomatal conductance

BACKGROUNDS AND AIMS: Identification of physiological traits associated with leaf photosynthetic rate (Pn) is important for improving potential productivity of rice (Oryza sativa). The objectives of this study were to develop a model which can explain genotypic variation and ontogenetic change of Pn in rice under optimal conditions as a function of leaf nitrogen content per unit area (N) and stomatal conductance (g(s)), and to quantify the effects of interaction between N and g(s) on the variation of Pn. METHODS: Pn, N and g(s) were measured at different developmental stages for the topmost fully expanded leaves in ten rice genotypes with diverse backgrounds grown in pots (2002) and in the field (2001 and 2002). A model of Pn that accounts for carboxylation and CO diffusion processes, and assumes that the ratio of internal conductance to g(s) is constant, was constructed, and its goodness of fit was examined. KEY RESULTS: Considerable genotypic differences in Pn were evident for rice throughout development in both the pot and field experiments. The genotypic variation of Pn was correlated with that of g(s) at a given stage, and the change of Pn with plant development was closely related to the change of N. The variation of g(s) among genotypes was independent of that of N. The model explained well the variation in Pn of the ten genotypes grown under different conditions at different developmental stages. Conclusions The response of Pn to increased N differs with g(s), and the increase in Pn of genotypes with low g(s) is smaller than that of genotypes with high g(s). Therefore, simultaneous improvements of these two traits are essential for an effective breeding of rice genotypes with increased Pn.     

The role of calcium in ABA-induced gene expression and stomatal movements

There is much interest in the transduction pathways by which abscisic acid (ABA) regulates stomatal movements (ABA-turgor signalling) and by which this phytohormone regulates the pattern of gene expression in plant cells (ABA-nuclear signalling). A number of second messengers have been identified in both the ABA-turgor and ABA-nuclear signalling pathways. A major challenge is to understand the architecture of ABA-signalling pathways and to determine how the ABA signal is coupled to the appropriate response. We have investigated whether separate Ca2+-dependent and -independent ABA-signalling pathways are present in guard cells. Our data suggest that increases in [Ca2+]i are a common component of the guard cell ABA-turgor and ABA-nuclear signalling pathways. The effects of Ca2+ antagonists on ABA-induced stomatal closure and the ABA-responsive CDeT6-19 gene promoter suggest that Ca2+ is involved in both ABA-turgor signalling and ABA-nuclear signalling in guard cells. However, the sensitivity of these pathways to alterations in the external calcium concentration differ, suggesting that the ABA-nuclear and ABA-turgor signalling pathways are not completely convergent. Our data suggest that whilst Ca2+-independent signalling elements are present in the guard cell, they do not form a completely separate Ca2+-independent ABA-signalling pathway.     

High yields in advanced lines of Pima cotton are associated with higher stomatal conductance, reduced leaf area and lower leaf temperature

Advanced lines of Pima cotton ( Gossypium barbadense L.) bred for higher yield potential and heat resistance have higher stomata conductance and smaller leaf areas than those of obsolete lines. In controlled experiments, five commercial lines of Pima cotton having increasing lint yield and heat resistance showed a gradient of increasing stomatal conductance and decreasing leaf size. In field experiments, heat-sensitive, low yield Pima lines showed a lower stomatal conductance than high yielding, advanced lines. This indicates that selection for high yield potential and heat resistance has imposed a selection pressure for higher stomatal conductance and smaller leaf areas. The higher stomatal conductance and smaller leaf area in the advanced lines resulted in a lower leaf temperature in both controlled environments and in the field. The largest leaf temperature differences between obsolete and advanced lines were observed in the afternoon. These differences coincided with the largest differences in stomatal conductance and the highest air temperatures. Measurements of stomatal conductance and leaf temperature in field-grown progeny from a cross between the advanced line, Pima S-6. and the obsolete line, Pima 32, showed that genetically determined differences in stomatal conductance resulted in corresponding differences in leaf temperature. None of the altered physiological traits were selected for in the breeding program, indicating that selection for the desired agronomic traits imposed selection pressures on the altered physiological traits. The increases in stomatal conductance and decreases in leaf area could represent an integrated response to selection pressures on enhanced evaporative cooling, ensuing from selection for heat resistance.     

Modelling the soil-plant-atmosphere continuum in a Quercus–Acer stand at Harvard Forest: the regulation of stomatal conductance by light, nitrogen and soil/plant hydraulic properties

Our objective is to describe a multi-layer model of C₃-canopy processes that effectively simulates hourly CO₂ and latent energy (LE) fluxes in a mixed deciduous Quercus-Acer (oak–maple) stand in central Massachusetts, USA. The key hypothesis governing the biological component of the model is that stomatal conductance (g_s) is varied so that daily carbon uptake per unit of foliar nitrogen is maximized within the limitations of canopy water availability. The hydraulic system is modelled as an analogue to simple electrical circuits in parallel, including a separate soil hydraulic resistance, plant resistance and plant capacitance for each canopy layer. Stomatal opening is initially controlled to conserve plant water stores and delay the onset of water stress. Stomatal closure at a threshold minimum leaf water potential prevents xylem cavitation and controls the maximum rate of water flux through the hydraulic system. We show a strong correlation between predicted hourly CO₂ exchange rate (r²= 0.86) and LE (r²= 0.87) with independent whole-forest measurements made by the eddy correlation method during the summer of 1992. Our theoretical derivation shows that observed relationships between CO₂ assimilation and LE flux can be explained on the basis of stomatal behaviour optimizing carbon gain, and provides an explicit link between canopy structure, soil properties, atmospheric conditions and stomatal conductance.     

三种植被恢复树种的冠层气孔导度特征及其对环境因子的敏感性

应用Granier热消散探针,长期监测华南地区荷木、大叶相思和柠檬桉林不同径级样树的树干液流,结合同步观测的气象数据,求算冠层气孔导度(g_c),并分析其对环境因子的响应方式及敏感性.结果表明: 不同季节荷木林日间平均g_c显著高于大叶相思和柠檬桉(P<0.05)（除3月外）.在干季和湿季,g_c与光合有效辐射(PAR)呈现对数正相关关系(P<0.001),湿季g_c对PAR响应比干季更敏感.g_c与水汽压亏缺(VPD)在干湿季均呈现对数负相关关系(P<0.001),同样在湿季表现出更高的敏感性.湿季g_c与VPD的偏相关系数高于干季,VPD对气孔行为的调控作用在湿季更为明显.随着土壤含水量的降低,g_c对VPD的敏感性下降,荷木和柠檬桉林下降的幅度大于大叶相思林,荷木和柠檬桉林下降的幅度相当.通过综合分析g_c对环境因子(PAR和VPD)的敏感性及其对土壤含水量变化的响应规律,发现乡土树种荷木作为植被恢复树种比外来引种的大叶相思和柠檬桉更为适宜.