Gas exchange responses of two Eucalyptus species to salinity and waterlogging

The effects of salinity and waterlogging on stomatal conductance, net photosynthesis and transpiration of 3-month-old Eucalyptus camaldulensis Dehnh. and Eucalyptus lesouefii Maiden seedlings were studied under greenhouse conditions. Under non-saline conditions, waterlogging induced stomatal closure in both species. However, the stomata of E. camaldulensis reopened after five weeks, when adventitious roots were produced. Relative to that of controls, height growth of waterlogged seedlings was greater in E. camaldulensis than in E. lesouefii, as were rates of photosynthesis and transpiration. In a freely drained medium, high salinity reduced rates of seedling height growth and photosynthesis, relative to those in controls, less in E. lesouefii than in E. camaldulensis. In both species, height growth, stomatal conductance and photosynthetic rate were lowest under conditions of saline waterlogging.     

Photosynthetic capacity and foliar nitrogen distribution in Eucalyptus nitens is altered by high-intensity thinning

The changes in photosynthetic rates, light environment and foliar nutrient concentrations following thinning were examined in an 8-year-old Eucalyptus nitens (Deane and Maiden) Maiden plantation. The objectives of the study were to: (1) determine the extent to which maximum photosynthetic rates (Amax) of E. nitens are affected by stand thinning; (2) relate the spatial pattern of Amax within the crown to the changes in light environment caused by thinning; and (3) establish if the responses of Amax to thinning are driven by changes in area-based foliar nitrogen (Na) or phosphorus (Pa) concentrations. Photosynthetic rates measured under saturating light availability increased throughout the crown after thinning. The greatest increases were observed in the lower and middle crown zones. Photosynthetic rate was positively related to foliar N concentration. Thinning increased Na and Pa because of a significant decrease in specific leaf area (SLA) after thinning. Changes in photosynthetic rates, SLA and foliar nutrient distributions with thinning were well correlated with changes in incident solar irradiance throughout the tree crown.     

保水剂对巨尾桉盆栽苗木生长的影响

为评价保水剂在桉树生长中的作用,研究了2种保水剂用量和4种水分供应条件下,巨尾桉苗木的生长量、净光合速率、蒸腾速率和叶绿素含量的变化情况.结果表明,随着田间持水量增加,除了叶绿素含量差别不大,苗木的高增长量、地径增长量、净光合速率均显著上升;施用保水剂的效果与田间持水量60％和80％处理的作用基本一致,保水剂在桉树人工...     

Vertical gradients in photosynthetic light response within an old-growth Douglas-fir and western hemlock canopy

We examined needle-level light response of photosynthesis across a vertical light gradient within 45-55-m-tall western hemlock (Tsuga heterophylla (Raf.) Sarg.) and Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees growing in a 400-500-year-old mixed species stand. We determined: (1) whether light-saturated photosynthetic rates, light compensation points, and respiration rates varied from the upper to the lower canopy, and (2) if light-saturated photosynthetic rates, light compensation points, and respiration rates varied between Douglas-fir and western hemlock. Over a 25-m gradient from the canopy top to the lower canopy, mean light-saturated photosynthetic rates, light compensation points, and respiration rates declined in overstory Douglas-fir and western hemlock needles, paralleling a 65% decline in the mean daily photosynthetic photon flux density (PPFD). At the canopy top, increasing light-saturated photosynthetic rates relative to lower canopy needles increased carbon uptake at high PPFD. In the lower canopy, reduced respiration rates relative to upper canopy needles increased carbon uptake at low PPFD by reducing the light compensation point. At all canopy positions, western hemlock had lower mean light-saturated photosynthetic rates, light compensation points and respiration rates than Douglas-fir. As a result, western hemlock had higher net photosynthetic rates at low PPFD, but lower net photosynthetic rates at high PPFD compared with Douglas-fir.     

尾巨桉与杉木、香樟耗水特征分析

在广西黄冕林场采用Granier热扩散技术方法对人工纯林桉树、杉木和香樟的液流密度、液流通量、单株日均耗水量及耗水量与主要环境因子的关系进行了研究.结果表明:3个树种树干液流密度均与光合有效辐射有一致的变化规律和单峰曲线.综合分析显示桉树在3个树种中耗水能力最强,虽然香樟液流密度比杉木大,但因为边材面积最小,所以表现出耗水量最小,单株日均耗水量仅有桉树的0.5倍左右,而杉木虽然边材面积比桉树大,但耗水量并不高,仅为桉树的0.7倍左右;通过偏相关分析发现3个树种的液流通量均与光合有效辐射、水气压亏缺和空气温度有极显著的正相关关系,杉木和香樟液流通量与空气相对湿度有极显著的负相关关系,而桉树表现不显著;通过多元线性分析发现3个树种的液流通量均与光合有效辐射、空气温度和空气相对湿度有很强的线性相关关系.3个不同树种的耗水研究结果可为本地区造林树种选择的优劣性提供参考.     

Increasing the sink:source balance enhances photosynthetic rate of 1-year-old balsam fir foliage by increasing allocation of mineral nutrients

To determine the impact of altering the sink:source balance on gas exchange rates in 1-year-old foliage of balsam fir (Abies balsamea (L.) Mill.), seedlings were either debudded before the growing season began or left intact. To assess alternative explanations for the observed photosynthetic response, additional seedlings were variously root pruned, shaded, drought stressed or deprived of fertilizer in combination with debudding. Foliar gas exchange rates and carbohydrate concentrations, xylem water potential and current-year growth were measured on three occasions spanning the shoot extension period. In addition, growth, foliar concentrations of chlorophyll, nitrogen, phosphorus and potassium, chlorophyll fluorescence and photosynthetic rate at different CO2 concentrations were determined 1 month after shoot extension ceased. Debudded seedlings had higher photosynthetic rates than budded seedlings, and the difference increased as the growing season progressed. We observed no evidence that debudding increased photosynthetic rate by improving water relations or by reducing foliar carbohydrate concentrations. Debudding increased foliar concentrations of nitrogen, phosphorus, potassium and chlorophyll, as well as quantum yield and photochemical quenching as determined by chlorophyll fluorescence measurements. Therefore, we conclude that debudding increased the photosynthetic rate by increasing the allocation of nutrients to 1-year-old foliage, thereby enhancing the amount or activity, or both, of photosynthetic enzymes, as well as increasing chlorophyll concentration.     

五种彩叶植物光合日变化研究

采用美国生产的LCI便携式植物光合测定仪测定自然生长的红龙草、黄金榕、变叶木、红背桂和鹅掌柴等5种彩叶植物的净光合速率（Pn）、气孔导度（Gs）、细胞间隙CO2浓度（Gi）、蒸腾速率（Tr）和气孔限制值（Ls）等光合特征值的日变化。结果表明：红龙草、黄金榕和变叶木的Pn、Gs和Tr日变化曲线呈“单峰”型，其Pn、Gs和0在10：00达到高峰；红背桂和鹅掌柴的Pn、Gs和Tr日变化曲线呈“双峰”型，其Pn、Gs和Tr分别在10：00和16：00达到高峰，且第1个高峰值大于第2个高峰值。5种彩叶植物的Ci日变化均先降后升，红龙草、黄金榕和变叶木的Ci在14：00出现低谷，红背桂和鹅掌柴的Ci在10：00出现低谷。5种彩叶植物的Ls日变化均先升后降，红龙草和变叶木的Ls在14：00迭最高值，黄金榕的Ls在12：00达最高值，红背桂和鹅掌柴的Ls在10：00达最高值。     

红瑞木光合特性研究

应用LI-6400便携式光合测定系统对红瑞木光合作用的日变化规律进行研究,利用SPSS统计分析软件,采用相关分析及多元逐步回归分析得出红瑞木春季、夏季、秋季净光合速率与生理生态因子间的关系。结果表明:春季光合速率呈单峰曲线,影响光合速率的主要因子为气孔导度、胞间CO2浓度和温度;夏季光合速率呈单峰曲线,在16:30左右又出现一个小高峰,影响光合速率的主要因子为蒸腾速率、光合有效辐射和温度;秋季净光合速率呈双峰曲线,影响光合速率的主要因子为光合有效辐射、蒸腾速率、胞间CO2浓度和大气水气压亏缺;夏季净光合速率最大。     

紫茎泽兰对模拟酸雨胁迫的生理反应

对pH值为3．0、4．0和5．0的模拟酸雨对紫茎泽兰光合等生理特性影响的研究结果表明，随着模拟酸雨pH值的降低，紫茎泽兰叶片中丙二醛及可溶性糖的含量均较对照（pH5．6）显著升高，但并未表现出明显的受害症状；叶片中叶绿体色素含量缓慢下降（pH5．0～4．0）或显著增加（pH3．0），光饱和点和补偿点均较对照有所降低；净光合速率（Pn）除PAR低于200μmol／m^2·s时pH5．0的处理略低于对照外，其他处理均明显高于对照。在模拟酸雨的胁迫下，紫茎泽兰生长良好，叶片中仍能维持较高的叶绿体色素含量和净光舍速率，表明紫茎泽兰对酸雨具有较强的耐受性。     

干旱对6种野生花卉光合色素含量与气体交换的影响

为给野生耐旱花卉在城市绿化中的应用提供理论依据,在温室内用盆栽法测定了燕山地区6种野生耐旱花卉苗木干旱胁迫进程中的对照、干旱胁迫5d、暂时萎蔫和永久萎蔫4点的光合色素含量与气体交换变化。结果表明:在土壤干旱进程的对照、干旱5d、暂时萎蔫和永久萎蔫各时间的总叶绿素、类胡萝卜素和叶绿素a与叶绿素b比值,除栓翅卫矛、孩儿拳头和木半夏各自的总叶绿素含量没有明显差别外,其余各自的总叶绿素含量、类胡萝卜素含量和叶绿素a与叶绿素b比值等指标的差异均显著或极显著;随土壤含水量降低,6种苗木蒸腾速率、净光合速率和气孔导度总体呈下降趋势,在永久萎蔫点三项指标降低幅度排序:蒸腾速率为木半夏>蚂蚱腿子>南蛇藤>孩儿拳头>叶底珠>栓翅卫矛;净光合速率为孩儿拳头>叶底珠>蚂蚱腿子>南蛇藤>木半夏>栓翅卫矛;气孔导度为蚂蚱腿子>叶底珠>孩儿拳头>南蛇藤>木半夏>栓翅卫矛;不同苗木的光合色素含量和三个指标对干旱胁迫的反应不同预示各苗木具有不同适应干旱的机理。     

Increased photosynthesis following partial defoliation of field-grown Eucalyptus globulus seedlings is not caused by increased leaf nitrogen

Increased photosynthetic rates following partial defoliation may arise from changes in leaf biochemistry, water relations or nutrient status. Twelve-month-old field-grown Eucalyptus globulus Labill. seedlings were pruned from below to reduce the green crown depth by 50 (D50) or 70% (D70). Photosynthetic responses to light and CO2 concentration were examined before and one, three and five weeks after partial defoliation. One week after defoliation, photosynthetic rates were greater in seedlings in the D50 (21 micromol m(-2) s(-1)) and D70 (23 micromol m(-2) s(-1)) treatments than in control seedlings (15 micromol m(-2) s(-1)); however, there was little difference in photosynthetic rates between partially defoliated seedlings and control seedlings after 5 weeks. An analysis of the sensitivity of photosynthesis to biochemical parameters revealed that the transient increase in photosynthetic rate in response to partial defoliation was largely a function of the maximum rate of carboxylation (85-87%) and the maximum rate of RuBP regeneration (55-60%) rather than stomatal conductance (12-13%). Nitrogen increased in leaves following partial defoliation (increases of 0.6 and 1.2 g m(-2) for D50 and D70, respectively), but was accumulated in a non-photosynthetic form (i.e., there was no increase in nitrogen concentration of Rubisco or chlorophyll). Increased photosynthetic rates immediately following partial defoliation were primarily a result of increased activity rather than amount of photosynthetic machinery. There was no evidence that phosphorus was responsible for the increase in photosynthetic rates after partial defoliation.     

Responses of transpiration and photosynthesis to reversible changes in photosynthetic foliage area in western red cedar (Thuja plicata) seedlings

Experiments were conducted on 1-year-old western red cedar (Thuja plicata Donn.) seedlings to determine the response of illuminated foliage to reversible changes in total photosynthetic foliage area (L(A)). Reductions in L(A) were brought about by either shading the lower foliage or by reducing the ambient CO2 concentration (c(a)) of the air surrounding the lower part of the seedling. In the latter case, the vapor pressure was also changed so that transpiration rates (E) could be manipulated independently of photosynthetic rates (A). We hypothesized that following such treatments, short-term compensatory changes would occur in stomatal conductance (g(s)) and A of the remaining foliage. These changes would occur in response to hydraulic signals generated by changes in the water potential gradient rather than changes in the distribution of sources and sinks of carbon within the seedling. When a portion of the foliage was shaded, there was an immediate reduction in whole-seedling E and a concomitant increase in g(s), A and E in the remaining illuminated foliage. However, the intercellular CO2 concentration did not change. These compensatory effects were fully reversed after the shade was removed. When the lower foliage A was reduced to < 0 micromol m-2 s-1, by shading or lowering c(a), and E was either unchanged or increased (by adjusting the vapor pressure deficit), there was no significant increase in g(s) and A in the remaining foliage. We conclude that compensatory responses in illuminated foliage occur only when reductions in L(A) are accompanied by a reduction in whole-plant E. The relationship between the reduction in whole-seedling E and the increase in A is highly linear (r2 = 0.68) and confirms our hypothesis of the strong regulation of g(s) by hydraulic signals generated within the seedling. We suggest that the mechanism of the compensatory effects is a combination of both increased CO2 supply, resulting from increased g(s), and a response of the rate of carboxylation, possibly related to the activity of Rubisco.     

Morphological and kinetic parameters of the absorption of nitrogen forms for selection of Eucalyptus clones

Eucalyptus clones are selected according to productivity,wood quality,rooting capacity,and resistance to drought,frost and diseases.However,kinetic and morphological parameters that determine the absorption efficiency of nutrients such as nitrate(NO_3~-) and ammonium(NH_4~+)are often not considered in breeding programs.The objective of this study was to evaluate the morphological,physiological and kinetic parameters of nitrogen uptake by clones of Eucalyptus saligna(32,864) and Eucalyptus grandis(GPC23).Morphological parameters in shoot and root systems,biomass and N concentrations in different organs,photosynthetic pigment concentrations,parameters of chlorophyll a fluorescence and photosynthetic rates were evaluated.Kinetic parameters,maximum absorption velocity(V_(max)),Michaelis-Menten constant(K_m),minimum concentration(C_(min)) and influx(I) were calculated for NO_3~-and NH_4~+ in the two clones.E.granais clone was more efficient in the uptake of NO_3~-and NH_4~+,and showed lower K_m and C_(min)values,allowing for the absorption of nitrogen at low concentrations due to the high affinity of the absorption sites of clone roots to NO_3~-and NH_4~+.Higher root lengths,area and volume helped the E.grandis clone in absorption efficiency and consequently,resulted in higher root and shoot biomass.The E.saligna clone had higher K_m and Cmin for NO_3~-and NH_4~+,indicating adaptation to environments with higher N availability.The results of NO_3~-and NH_4~+ kinetic parameters indicate that they can be used in Eucalyptus clone selection and breeding programs as they can predict the ability of clones to absorb NO_3~-and NH_4~+ at different concentrations.     

Climatic factors controlling the productivity of Norway spruce: A model-based analysis

The process-based growth model, BIOMASS, was modified to incorporate low-temperature effects on photosynthetic production in Norway spruce (Picea abies) stands growing in northern Sweden. The low-temperature features incorporated in BIOMASS made it possible to simulate and estimate the reduction in photosynthetic rates caused by boreal conditions. The following four simulation-scenarios were used: (i) ‘potential' photosynthesis without boreal restrictions; (ii) reduction caused by a frozen soil; (iii) reduction caused by incomplete recovery of photosynthetic capacity during spring as a result of damage caused by low winter temperatures; and (iv) reduction as an effect of frost-induced autumn decline. Annual photosynthetic production (or gross primary production (GPP)) was simulated for three calendar years, 1990–1992, for stands with low (control) and high (irrigated and fertilized) nutrient availability. The reduction of ‘potential' GPP, caused by the low-temperature effects, ranged from 35–44% for control (C) and from 34–42% for irrigated-fertilised (IL) stands, respectively. The most pronounced loss of ‘potential' GPP originated from reduced photosynthetic capacity, in spring and early summer, which led to losses of 21–28% for C and 19–26% for IL stands. The variation between years differed mainly as an effect of differences in spring temperatures, which resulted in different rates of recovery of photosynthetic capacity. Reductions caused by frozen soil and low photosynthetic capacity during winter were similar in C and IL stands (12–13%), as were the losses resulting from severe autumn frosts (3–4%). It is concluded that, unless the effects of frozen soils and reduced photosynthetic capacity during spring and early summer are considered, large errors (ca. 40%) will be introduced into estimates of the annual photosynthetic production of boreal conifer forests.     

Light availability and photosynthesis of Pseudotsuga menziesii seedlings grown in the open and in the forest understory

The light environment, photosynthetic dynamics and steady-state net photosynthetic rates of lateral branch shoots of Pseudotsuga menziesii var. glauca (Beissn.) Franco seedlings growing in the open and in the forest understory were investigated in situ. Mean incident photosynthetic photon flux density (PPFD) was 702.5 micro mol m(-2) s(-1) on open-grown branches and 52.0 micro mol m(-2) s(-1) on understory-grown branches. Mean daily durations of PPFD greater than 500, 200, and 50 micro mol m(-2) s(-1) were 8.5, 31.5, and 270.3 min, respectively, on understory-grown branches, and 559.1, 700.7, and 803.3 min, respectively, on open-grown branches. Sunflecks accounted for 32.4% of total daily photosynthetically active radiation incident on understory branches. Following 10 min at a PPFD of 50 micro mol m(-2) s(-1), the induction time required for net photosysnthesis to reach 50 and 90% of steady-state rates was shorter at a PPFD of 200 than at a PPFD of 500 micro mol m(-2) s(-1) and shorter in understory-grown branches than in open-grown branches. On a leaf area basis, dark respiration rates of understory-grown branches were lower and net photosynthetic rates were higher than those of open-grown branches exposed to low PPFD. However, at high PPFDs, understory-grown branches had lower photosynthetic rates than open-grown branches. When measurements were expressed on a leaf dry mass basis, there was no difference in dark respiration rates between understory branches and open-grown branches, but net photosynthetic rates of understory branches were equal to or higher than those of open-grown branches at all PPFDs.     

Influence of water stress on the physiology and growth of red spruce seedlings

Two-year-old, container-grown red spruce (Picea rubens Sarg.) seedlings from a New Hampshire seed source were exposed to 10 or 11 drying cycles in which the seedlings were not watered until their midday (1400 h) xylem water potentials averaged -1.57 MPa. Control seedlings were kept well watered to maintain midday water potentials of about -0.73 MPa. After the final drying cycle, the water-stressed seedlings were rehydrated and osmotic potentials were determined by pressure-volume analysis. Gas exchange at ambient CO(2) concentration (338 ppm) and at an elevated CO(2) concentration (838 ppm) was measured on both groups of plants as they slowly dried down. No osmotic adjustment or photosynthetic acclimation occurred as a result of the water-stress treatment and both groups of seedlings maintained photosynthesis to water potentials as low as -3.0 MPa. Twenty-four hours after rehydration, the water-stressed seedlings had photosynthetic rates as high as the control seedlings. Estimated stomatal limitation to photosynthesis was approximately 30% down to water potentials of -1.4 MPa, but increased steadily as water potentials decreased further. At ambient CO(2) concentrations (338 ppm) and water potentials averaging -2.45 MPa, photosynthetic rates of water-stressed seedlings were 15% those of well-watered seedlings, whereas when the same water-stressed seedlings were measured in the presence of an elevated concentration of CO(2) (838 ppm) their photosynthetic rates were 73% those of well-watered seedlings measured at an ambient CO(2) concentration (338 ppm).     

Photosynthesis and water relations of savanna tree species differing in leaf phenology

Godmania macrocarpa Hemsley, a deciduous tree characteristic of fire protected areas of the savanna region of central Venezuela, was more drought tolerant, allocated more N to leaves and had consistently higher photosynthetic rates than Curatella americana L., a ubiquitous species growing successfully within the grasslands of tropical American savannas. Godmania macrocarpa maintained higher leaf conductance and photosynthesized at higher xylem water tensions than C. americana. As the dry season progressed, G. macrocarpa was more affected by water stress than C. americana, which may explain why G. macrocarpa shed its leaves before forming new leaves. For both species, leaf sap osmolality was strongly correlated with, but not completely accounted for by, soluble sugars. Integrated water-use efficiency, as measured by delta(13)C, was similar for both species, but young leaves were more efficient than old leaves. Water-use efficiency of adult leaves was similar in both species as a result of higher photosynthetic rates in G. macrocarpa and lower leaf conductances in C. americana. Compared to G. macrocarpa, instantaneous photosynthetic N-use efficiency was higher in C. americana despite its lower maximum photosynthetic rates. The absence of G. macrocarpa trees from open grasslands, despite their high productive capacity, is possibly the result of unfavorable conditions for germination, poor survival of seedlings, and lack of resistance against fire.     

遮光对野鸦椿幼苗生长和光合生理特性的影响

为了解不同光照条件下野鸦椿幼苗的生长发育状况和光合生理特性,探讨野鸦椿幼苗生长发育的最佳光照条件.采用遮光方式模拟不同光照强度(全光照,遮光30％、60％和90％),研究野鸦椿幼苗的生长、生物量、光合色素含量、生理指标和光合日变化等光合特性的响应规律.结果表明:随着遮光程度的增加,野鸦椿幼苗的生长量均高于对照,以遮光6...     

Needle longevity, photosynthetic rate and nitrogen concentration of eight spruce taxa planted in northern Japan

Growth characteristics of Picea glehnii Masters, P. jezoensis (Sieb. et Zucc) Carr., P. jezoensis var. hondoensis (Mayr) Rehder and P. shirasawae Hayashi from Japan, P. abies (L.) Karst. from Europe and P. glauca Voss, P. mariana Britt., Sterns and Pogg. and P. rubens Sarg. from North America were compared. The trees were grown in similar conditions at the Tomakomai Experimental Forest of Hokkaido University in northern Japan. Tree growth, needle biomass, longevity, photosynthetic rate, nitrogen concentration and specific leaf area (SLA) were measured, and photosynthetic nitrogen-use efficiency was calculated. Picea jezoensis, P. jezoensis var. hondoensis, P. abies and P. glauca had high growth rates, high photosynthetic rates in young needles, high needle nitrogen concentrations and short needle life spans. In contrast, P. glehnii, P. shirasawae, P. mariana and P. rubens had low growth and photosynthetic rates, low needle nitrogen concentrations, long needle life spans and maintained a high photosynthetic nitrogen-use efficiency in older needles. Examination of relationships between several growth parameters of the eight taxa revealed positive correlations between SLA and mass-based photosynthetic rate and between SLA and mass-based nitrogen concentration, whereas mass-based photosynthetic rate and mass-based nitrogen concentration were negatively correlated with needle longevity. The species differed greatly in growth characteristics despite being grown in similar conditions.     

Seasonal trends in photosynthetic parameters and stomatal conductance of blue oak (Quercus douglasii) under prolonged summer drought and high temperature

Understanding seasonal changes in photosynthetic parameters and stomatal conductance is crucial for modeling long-term carbon uptake and energy fluxes of ecosystems. Gas exchange measurements of CO2 and light response curves on blue oak leaves (Quercus douglasii H. & A.) were conducted weekly throughout the growing season to study the seasonality of photosynthetic capacity (Vcmax) and Ball-Berry slope (m) under prolonged summer drought and high temperature. A leaf photosynthetic model was used to determine Vcmax. There was a pronounced seasonal pattern in Vcmax. The maximum value of Vcmax, 127 micromol m(-2) s(-1), was reached shortly after leaf expansion in early summer, when air temperature was moderate and soil water availability was high. Thereafter, Vcmax declined as the soil water profile became depleted and the trees experienced extreme air temperatures, exceeding 40 degrees C. The decline in Vcmax was gradual in midsummer, however, despite extremely low predawn leaf water potentials (Psipd, approximately -4.0 MPa). Overall, temporal changes in Vcmax were well correlated with changes in leaf nitrogen content. During spring leaf development, high rates of leaf dark respiration (Rd, 5-6 micromol m(-2) s(-1)) were observed. Once a leaf reached maturity, Rd remained low, around 0.5 micromol m(-2) s(-1). In contrast to the strong seasonality of Vcmax, m and marginal water cost per unit carbon gain (partial partial differential E/ partial partial differential A) were relatively constant over the season, even when leaf Psipd dropped to -6.8 MPa. The constancy of partial partial differential E/ partial partial differential A suggests that stomata behaved optimally under severe water-stress conditions. We discuss the implications of our findings in the context of modeling carbon and water vapor exchange between ecosystems and the atmosphere.