Stomatal and environmental control of transpiration in a lowland tropical forest tree

Stomatal control of crown transpiration was studied in Anacardium excelsum, a large-leaved, emergent canopy species common in the moist forests of Central and northern South America. A construction crane equipped with a gondola was used to gain access to the uppermost level in the crown of a 35-m-tall individual. Stomatal conductance at the single leaf scale, and transpiration and total vapour phase conductance (stomatal and boundary layer) at the branch scale were measured simultaneously using the independent techniques of porometry and stem heat balance, respectively. This permitted the sensitivity of transpiration to a marginal change in stomatal conductance to be evaluated using a dimensionless coupling coefficient (1-ω) ranging from zero to 1, with 1 representing maximal stomatal control of transpiration. Average stomatal conductance varied from 0.09 mol m^?2 s^?1 during the dry season to 0.3 mol m^?2 s^?1 during the wet season. Since boundary layer conductance was relatively low (0.4 mol m^?2 s^?1), 1-ω ranged from 0.46 during the dry season to only 0.25 during the wet season. A pronounced stomatal response to humidity was observed, which strongly limited transpiration as evaporative demand increased. The stomatal response to humidity was apparent only when the leaf surface was used as the reference point for measurement of external vapour pressure. Average transpiration was predicted to be nearly the same during the dry and wet seasons despite a 1 kPa difference in the prevailing leaf-to-air vapour pressure difference. The patterns of stomatal behaviour and transpiration observed were consistent with recent proposals that stomatal responses to humidity are based on sensing the transpiration rate itself.     

Stomatal response to vapour pressure deficit and the effect of plant water stress

Abstract The dynamic response of stomata to changes in atmospheric humidity was investigated in Fragaria × ananassa Duch., Picea engelmannii Parry, and Pseudotsuga menziesii (Mirb.) Franco; and the effect of water stress on this response was determined in Pseudotsuga menziesii. The plants were rotated through three regimes of ambient temperature and vapour pressure deficit: 35°C–3. 5kPa, 35°C–0. 5 kPa, and 20°C–1. 5kPa. Branch and leaflet conductance were measured with a steady-state porometer, first at ambient vapour pressure deficit and then at one of four treatment conditions achieved by increasing or decreasing vapour pressure within the porometer cuvette. All three species showed similar stomatal response: enhanced conductance at low vapour pressure deficit and depressed conductance at high vapour pressure deficit. Engelmann spruce was more sensitive than Douglas fir and strawberry. Plant water status significantly altered stomatal response to vapour pressure deficit. The relationship of conductance of xylem water potential was linear under ambient conditions but became curvilinear when conductance was measured above and below ambient vapour pressure deficit. Between ?0. 5 MPa and ?2. 0 MPa xylem water potential, the stomata were sensitive to vapour pressure deficit, but below ? 2. 0 MPa, the sensitivity decreased.     

Stomatal responses to changes in vapour pressure difference between leaf and air

We observed that stomata of Gossypium hirsutum, Glycine max and Xanthium strumarium respond to a change in vapour pressure difference between leaf and air at ambient partial pressures of CO₂ and below the CO₂ compensation point. Our report is at variance with a recent report (J. A. Bunce 1997, Plant, Cell and Environment 20, pp. 131–135) that stomatal sensitivity of leaves to a change in vapour pressure difference between leaf and air was eliminated when gas exchange measurements were made at near-zero carbon dioxide partial pressures (0–5 Pa).     

Survey and synthesis of intra- and interspecific variation in stomatal sensitivity to vapour pressure deficit

Responses of stomatal conductance (g_s) to increasing vapour pressure deficit (D) generally follow an exponential decrease described equally well by several empirical functions. However, the magnitude of the decrease – the stomatal sensitivity – varies considerably both within and between species. Here we analysed data from a variety of sources employing both porometric and sap flux estimates of g_s to evaluate the hypothesis that stomatal sensitivity is proportional to the magnitude of g_s at low D ( ≤ 1 kPa). To test this relationship we used the function g_s = g_sref–m· lnD where m is the stomatal sensitivity and g_sref = g_s at D = 1 kPa. Regardless of species or methodology, m was highly correlated with g_sref (average r² = 0·75) with a slope of approximately 0·6. We demonstrate that this empirical slope is consistent with the theoretical slope derived from a simple hydraulic model that assumes stomatal regulation of leaf water potential. The theoretical slope is robust to deviations from underlying assumptions and variation in model parameters. The relationships within and among species are close to theoretical predictions, regardless of whether the analysis is based on porometric measurements of g_s in relation to leaf-surface D (D_s), or on sap flux-based stomatal conductance of whole trees (G_Si), or stand-level stomatal conductance (G_S) in relation to D. Thus, individuals, species, and stands with high stomatal conductance at low D show a greater sensitivity to D, as required by the role of stomata in regulating leaf water potential.     

Stomatal responses to changes in vapor pressure deficit reflect tissue‐specific differences in hydraulic conductance

The vapor pressure deficit (D) of the atmosphere can negatively affect plant growth as plants reduce stomatal conductance to water vapor (g_wv) in response to increasing D, limiting the ability of plants to assimilate carbon. The sensitivity of g_wv to changes in D varies among species and has been correlated with the hydraulic conductance of leaves (K_leaf), but the hydraulic conductance of other tissues has also been implicated in plant responses to changing D. Among the 19 grass species, we found that K_leaf was correlated with the hydraulic conductance of large longitudinal veins (K_lv, r² = 0.81), but was not related to K_root (r² = 0.01). Stomatal sensitivity to D was correlated with K_leaf relative to total leaf area (r² = 0.50), and did not differ between C₃ and C₄ species. Transpiration (E) increased in response to D, but 8 of the 19 plants showed a decline in E at high D, indicative of an ‘apparent feedforward’ response. For these individuals, E began to decline at lower values of D in plants with low K_root (r² = 0.72). These results show the significance of both leaf and root hydraulic conductance as drivers of plant responses to evaporative demand.     

Stomatal sensitivity to vapour pressure deficit of temperate and tropical evergreen rainforest trees of Australia

Although rainforests of eastern Australia grow in regions of high precipitation, there is a shift from a summer dry season in the temperate south to a winter dry season in the tropical north. Therefore, rainforest trees of eastern Australia provide an opportunity to investigate stomatal sensitivity of mesic trees to vapour pressure deficit (VPD) along a gradient in seasonality of precipitation. Eight rainforest canopy tree species were selected to cover the latitudinal range of rainforests in eastern Australia. Seedlings of these species were grown for a year in glasshouses under ambient conditions or at low VPD and water vapour exchange of leaves was measured during summer. Tropical species, which experience summer-dominant precipitation, showed higher stomatal sensitivities to VPD than temperate species, which experience winter-dominant precipitation. Growing plants under a low VPD increased stomatal sensitivity to increasing VPD in most species. The high stomatal sensitivity to VPD of the tropical species is consistent with the infrequent water stress experienced during their growing season and suggests a high susceptibility to water deficits. In contrast, temperate species may use other mechanisms to maintain photosynthesis under the relatively drier conditions of the temperate growing season.     

Stomatal responses to vapour pressure deficit are regulated by high speed gene expression in angiosperms

Plants dynamically regulate water use by the movement of stomata on the surface of leaves. Stomatal responses to changes in vapour pressure deficit (VPD) are the principal regulator of daytime transpiration and water use efficiency in land plants. In angiosperms, stomatal responses to VPD appear to be regulated by the phytohormone abscisic acid (ABA), yet the origin of this ABA is controversial. After a 20 min exposure of plants, from three diverse angiosperm species, to a doubling in VPD, stomata closed, foliar ABA levels increased and the expression of the gene encoding the key, rate‐limiting carotenoid cleavage enzyme (9‐cis‐epoxycarotenoid dioxygenase, NCED) in the ABA biosynthetic pathway was significantly up‐regulated. The NCED gene was the only gene in the ABA biosynthetic pathway to be up‐regulated over the short time scale corresponding to the response of stomata. The closure of stomata and rapid increase in foliar ABA levels could not be explained by the release of ABA from internal stores in the leaf or the hydrolysis of the conjugate ABA‐glucose ester. These results implicate an extremely rapid de novo biosynthesis of ABA, mediated by a single gene, as the means by which angiosperm stomata respond to natural changes in VPD.     

The stomatal response to evaporative demand persists at night in Ricinus communis plants with high nocturnal conductance

Evidence is building that stomatal conductance to water vapour (g(s)) can be quite high in the dark in some species. However, it is unclear whether nocturnal opening reflects a mechanistic limitation (i.e. an inability to close at night) or an adaptive response (i.e. promoting water loss for reasons unrelated to carbon gain). Further, it is unclear if stomatal responses to leaf-air vapour pressure difference (D) persist in the dark. We investigated nocturnal stomatal behaviour in castor bean (Ricinus communis L.) by measuring gas exchange and stomatal responses to D in the light and in the dark. Results were compared among eight growth environments [two levels for each of three treatment variables: air saturation deficit (D(a)), light and water availability]. In most plants, stomata remained open and sensitive to D at night. g(s) was typically lower at night than in the day, whereas leaf osmotic pressure (Pi) was higher at night. In well-watered plants grown at low D(a), stomata were less sensitive to D in the dark than in the light, but the reverse was found for plants grown at high D(a). Stomata of droughted plants were less sensitive to D in the dark than in the light regardless of growth D(a). Drought also reduced g(s) and elevated Pi in both the light and the dark, but had variable effects on stomatal sensitivity to D. These results are interpreted with the aid of models of stomatal conductance.     

The apparent feedforward response of stomata to air vapour pressure deficit: information revealed by different experimental procedures with two rainforest trees

A decrease in steady-state leaf transpiration rate with increased vapour pressure difference between leaf and air, which is reversible and independent of leaf water status, is evidence for feedforward control of stomatal aperture (Cowan 1977). A recent survey of gas exchange data by Monteith (1995), covering 52 sets of measurements on 16 species, reported that evidence for feedforward control was rare and usually reliant on a single point. We conducted gas exchange experiments on an additional 13 species and observed an apparent feedforward response in only two. However, the response was not reversible and depended upon experimental procedure. In view of this we discuss the appropriate use of the term ‘feedforward’.     

Leaf stomatal responses to vapour pressure deficit under current and CO2-enriched atmosphere explained by the economics of gas exchange

Using the economics of gas exchange, early studies derived an expression of stomatal conductance ( g ) assuming that water cost per unit carbon is constant as the daily loss of water in transpiration ( f _e) is minimized for a given gain in photosynthesis ( f _c). Other studies reached identical results, yet assumed different forms for the underlying functions and defined the daily cost parameter as carbon cost per unit water. We demonstrated that the solution can be recovered when optimization is formulated at time scales commensurate with the response time of g to environmental stimuli. The optimization theory produced three emergent gas exchange responses that are consistent with observed behaviour: (1) the sensitivity of g to vapour pressure deficit ( D ) is similar to that obtained from a previous synthesis of more than 40 species showing g to scale as 1 −  m  log( D ), where m   ∈  [0.5,0.6], (2) the theory is consistent with the onset of an apparent 'feed-forward' mechanism in g , and (3) the emergent non-linear relationship between the ratio of intercellular to atmospheric [CO₂] ( c _i/ c _a) and D agrees with the results available on this response. We extended the theory to diagnosing experimental results on the sensitivity of g to D under varying c _a.     

Gender specific patterns of carbon uptake and water use in a dominant riparian tree species exposed to a warming climate

Air temperatures in the arid western United States are predicted to increase over the next century. These increases will likely impact the distribution of plant species, particularly dioecious species that show a spatial segregation of the sexes across broad resource gradients. On the basis of spatial segregation patterns, we hypothesized that temperature increases will have a greater negative impact on female plants compared with co‐occurring male plants of dioecious species. This hypothesis was tested by examining the whole‐plant carbon and water relations of 10‐year‐old female (n = 18) and male (n = 13) Acer negundo Sarg. trees grown in a common garden in Salt Lake City, UT. The trees were established from cuttings collected where the growing season temperature averaged about 6.5 °C cooler than at the common garden. During May and June, stem sap flux (J_s) was similar between genders, but averaged 25% higher in males during the warmer months of July and August. Daytime canopy stomatal conductance (g_s) per unit leaf area was 12% higher in females in May : June, but was 11% higher in males in July : August. We combined measurements of sap flux–scaled transpiration with measurements of tree allometry and δ¹³C of leaf soluble sugars to estimate whole‐tree carbon assimilation (A_tree) and water use efficiency (WUE) (A_tree : E_tree). A_tree was similar between genders until late August when A_tree was 32% higher in male trees. A_tree : E_tree was on average 7% higher in females than in males during the growing season. Patterns of J_s, g_s, A_tree and A_tree : E_tree in the present study were in contrast to those previously reported for A. negundo genders under native growing season temperatures. Results suggest that the spatial segregation of the sexes could shift under global warming such that female plants lose their dominance in high‐resource habitats, and males increase their dominance in relatively lower‐resource habitats.     

一个气孔对环境因子响应的机理性教学模型

本文在已知的气孔运动机理的基础上，对现有的教学模型加以改进，建立了除ＶＰＤ、ＣＯ２和土壤水势的影响外，还能模拟光照和温度等因子对气孔影响的数学模型。在不同组合的环境因子下模拟得到的结果与公认的实验观测结果基本一致。它可以作为研究气孔整体行为的理论框架，也可用于检验多种关于气孔运动的假设。     

华南荷木林冠层气孔导度对水汽压亏缺的响应

冠层气孔导度(Gs)是量化气孔在冠层尺度水平上表现的参数,能够表征森林冠层表面水汽和能量交换的动态.本研究利用Granier树干液流测定系统,连续监测华南地区荷木林的树干液流,通过尺度转换和扩展获得冠层蒸腾速率,结合微气象观测值,以Pen-man-Monteith公式计算了Gs,并比较不同土壤水分条件下Gs对水汽压亏缺的响应.结果显示,Gs与气孔气体交换方法实测的叶片气孔导度(gs)日变化相似,单位转换数值大小与实测gs数量级一致.Gs对水汽压亏缺的响应在干季和湿季有明显差别:(1)在土壤水分充足的湿季(土壤含水量θ ＞33％),Gs对水汽压亏缺的响应更敏感(偏相关系数-0.316),而在干季(θ＜23％)则对光合有效辐射的响应更敏感(偏相关系数0.885).(2)荷木林冠层-大气脱耦联系数(Ω)在湿季接近l,干季则较湿季小,说明湿季叶片的界面层较厚,水汽压亏缺对Gs影响较小,而光合有效辐射是控制Gs的主要环境因子.     

Leaf- and stand-level responses of a forested mesocosm to independent manipulations of temperature and vapor pressure deficit

Alterations in temperature (T) and vapor pressure deficit (VPD) strongly influence gas exchange, but because VPD is highly influenced by T, the effects of these two factors are difficult to separate. Here, the concomitant effects of T and VPD on CO(2) uptake, stomatal conductance, and transpiration at leaf- and canopy-levels were examined for a stand of trees (Populus deltoides) enclosed within large mesocosms. T and VPD were independently altered to yield a factorial combination of treatments of low (24 degrees C) or high (30 degrees C) T and low (0.75) or high (1.75 kPa) VPD. Traditional leaf-level gas exchange measurements were compared with whole-canopy exchange to verify typical scaling methods. Elevated T resulted in an average 40% and 14% increase in midday leaf-level and canopy-level net CO(2) uptake, respectively. Other physiological responses to elevated T and VPD were similar at both scales, but the magnitude of change was usually less pronounced at the canopy-level. Surprisingly, only minimal interactions between T and VPD were found to influence responses of CO(2) uptake and stomatal conductance at either level.     

Stomatal sensitivity to vapour pressure difference over a subambient to elevated CO2 gradient in a C3/C4 grassland

In the present study the response of stomatal conductance (g_s) to increasing leaf‐to‐air vapour pressure difference (D) in early season C₃ (Bromus japonicus) and late season C₄ (Bothriochloa ischaemum) grasses grown in the field across a range of CO₂ (200–550 µmol mol^?1) was examined. Stomatal sensitivity to D was calculated as the slope of the response of g_s to the natural log of externally manipulated D (dg_s/dlnD). Increasing D and CO₂ significantly reduced g_s in both species. Increasing CO₂ caused a significant decrease in stomatal sensitivity to D in Br. japonicus, but not in Bo. ischaemum. The decrease in stomatal sensitivity to D at high CO₂ for Br. japonicus fit theoretical expectations of a hydraulic model of stomatal regulation, in which g_s varies to maintain constant transpiration and leaf water potential. The weaker stomatal sensitivity to D in Bo. ischaemum suggested that stomatal regulation of leaf water potential was poor in this species, or that non‐hydraulic signals influenced guard cell behaviour. Photosynthesis (A) declined with increasing D in both species, but analyses of the ratio of intercellular to atmospheric CO₂ (C_i/C_a) suggested that stomatal limitation of A occurred only in Br. japonicus. Rising CO₂ had the greatest effect on g_s and A in Br. japonicus at low D. In contrast, the strength of stomatal and photosynthetic responses to CO₂ were not affected by D in Bo. ischaemum. Carbon and water dynamics in this grassland are dominated by a seasonal transition from C₃ to C₄ photosynthesis. Interspecific variation in the response of g_s to D therefore has implications for predicting seasonal ecosystem responses to CO₂.     

The apparent feed-forward response to vapour pressure deficit of stomata in droughted, field-grown Eucalyptus globulus Labill

This study tested a multiplicative model of stomatal response to environment for drought‐affected trees of Eucalyptus globulus Labill. growing in southern Australia. The model incorporates a feed‐forward response to vapour pressure deficit of ambient air (δe_a) and performed well if evaluated using reduced major axis regression and log‐transformed data. There was strong evidence from gas‐exchange data, leaf water potentials and sapflow measurements of the feed‐forward response by stomata to leaf‐to‐air vapour pressure deficit (δe_l). The response of stomata to δe_l was irreversible. Stomatal conductance and the rate of net photosynthesis were highly correlated and declined, together with the rate of transpiration, throughout the afternoon as δe_a increased despite increasing leaf water potentials. The concentration of CO₂ inside leaves (c_i) increased as stomatal conductance declined indicating increasing non‐stomatal limitations to photosynthesis. The stomatal response to δe_l of E. globulus in the field is best described as an ‘apparent feed‐forward response’ that probably results from both slowly reversible depression of net photosynthesis and abscisic acid accumulation in guard cells. We suggest that the stomatal response to c_i may strengthen the link between photosynthetic capacity and stomatal conductance during leaf drying as a result of either drought or large δ e_l.     

Effects of water vapour pressure deficit and stomatal conductance on photosynthesis,internal pressurization and convective flow in three emergent wetland plants

Internal pressurization and convective gas flow in emergent wetland plants is a function of the water vapour pressure deficit (WPD) and stomatal conductance (G _s) separating the external atmosphere from the internal aerenchyma. We have compared the effects of WPD and G _s under a range of light intensities on static pressures and convective flows in Phragmites australis, Typha orientalis and Baumea articulata. The capacity of the three species to generate flows per unit leaf area differed, being greatest in P. australisand lowest in B. articulata. In all three species, decreasing light intensity from full sunlight (2200 mol m^–2 s^–1 photosynthetically active photon flux density (PPFD)) to < 200 and < 10 mol m^–2 s^–1PPFD caused immediate decreases in photosynthetic assimilation, followed by more gradual decreases in transpiration and G _s. However, internal pressures and flows in the two low light intensities remained similar to values recorded in full sunlight. WPD was more significantly related to pressures and flows in P. australis and T. orientalis than G _s. In B. articulata, pressures increased at low G _s values but flow rates were unaffected, as predicted by earlier models describing pore size effects on pressures and flows. The data suggest that emergent macrophytes can maintain significant internal convection even at low light intensities, and this may be beneficial for nocturnal aeration, particularly in arid climates where the atmospheric humidity at night is low.     

Stomata of trees growing in CO2-enriched air show reduced sensitivity to vapour pressure deficit and drought

Stomatal conductance ( g _s) and photosynthetic rate ( A ) were measured in young beech ( Fagus sylvatica ), chestnut ( Castanea sativa ) and oak ( Quercus robur ) growing in ambient or CO₂-enriched air. In oak, g _s was consistently reduced in elevated CO₂. However, in beech and chestnut, the stomata of trees growing in elevated CO₂ failed to close normally in response to increased leaf-to-air vapour pressure deficit (LAVPD). Consequently, while g _s was reduced in elevated CO₂ on days with low LAVPD, on warm sunny days (with correspondingly high LAVPD) g _s was unchanged or even slightly higher in elevated CO₂. Furthermore, during drought, g _s of beech and chestnut was unresponsive to [CO₂], over a wide range of ambient LAVPD, whereas in oak g _s was reduced by an average of 50% in elevated CO₂. Stimulation of A by elevated CO₂ in beech and chestnut was restricted to days with high irradiance, and was greatest in beech during drought. Hence, most of the additional carbon gain in elevated CO₂ was made at the expense of water economy, at precisely those times (drought, high evaporative demand) when water conservation was most important. Such effects could have serious consequences for drought tolerance, growth and, ultimately, survival as atmospheric [CO₂] increases.     

Stomatal response to humidity in a sugarcane field: simultaneous porometric and micrometeorological measurements*

Abstract. Gas exchange data obtained with wellventilated leaf cuvettes provide clear evidence of a stomatal response to leaf-air vapour pressure difference (V). In contrast, remotely sensed leaf temperatures with specific assumptions regarding canopy boundary layer characteristics, have been interpreted to mean that stomata do not respond to V. We address this apparent discrepancy in a sugarcane field by simultaneous application of a single-leaf, porometric technique and a whole-canopy, Bowen ratioenergy balance technique. These methods indicated significant stomatal response to V in well-irrigated sugarcane. Stomatal responses to V in the field were obscured by strong covariance of major environmental parameters so that opening responses to light and closing responses to V tended to offset each other. Low boundary layer conductance significantly uncoupled V at the leaf surface (V_s) from V determined in the bulk atmosphere (V_a). This reduced the range of the stimulus, V_s, thereby reducing the range of the stomatal response, without indicating low stomatal sensitivity to V. Stomatal responses to V_a may be smaller than expected from V response curves in cuvettes, since V_s rather than the conventionally measured V_a is analogous to V in a well-stirred cuvette. Recently published conclusions that remotely sensed canopy temperatures are inconsistent with stomatal response to V may be based on erroneous estimates of canopy boundary layer conductance and thus of V_s, use of air saturation deficit rather than V to express evaporative demand, and investigation at higher levels of evaporative demand than those eliciting maximal stomatal response.     

High temperature acclimation of C4 photosynthesis is linked to changes in photosynthetic biochemistry

With average global temperatures predicted to increase over the next century, it is important to understand the extent and mechanisms of C4 photosynthetic acclimation to modest increases in growth temperature. To this end, we compared the photosynthetic responses of two C4 grasses (Panicum coloratum and Cenchrus ciliaris) and one C4 dicot (Flaveria bidentis) to growth at moderate (25/20 degrees C, day/night) or high (35/30 degrees C, day/night) temperatures. In all three C4 species, CO2 assimilation rates (A) underwent significant thermal acclimation, such that when compared at growth temperatures, A increased less than what would be expected given the strong response of A to short-term changes in leaf temperature. Thermal photosynthetic acclimation was further manifested by an increase in the temperature optima of A, and a decrease in leaf nitrogen content and leaf mass per area in the high- relative to the moderate-temperature-grown plants. Reduced photosynthetic capacity at the higher growth temperature was underpinned by selective changes in photosynthetic components. Plants grown at the higher temperature had lower amounts of ribulose-1,5-bisphosphate carboxylase/oxygenase and cytochrome f and activity of carbonic anhydrase. The activities of photosystem II (PSII) and phosphoenolpyruvate carboxylase were not affected by growth temperature. Chlorophyll fluorescence measurements of F. bidentis showed a corresponding decrease in the quantum yield of PSII (phi(PSII)) and an increase in non-photochemical quenching (phi(NPQ)). It is concluded that through these biochemical changes, C4 plants maintain the balance between the various photosynthetic components at each growth temperature, despite the differing temperature dependence of each process. As such, at higher temperatures photosynthetic nitrogen use efficiency increases more than A. Our results suggest C4 plants will show only modest changes in photosynthetic rates in response to changes in growth temperature, such as those expected within or between seasons, or the warming anticipated as a result of global climate change.