Gas exchange and photosynthetic acclimation over subambient to elevated CO2 in a C3–C4 grassland

Atmospheric CO₂ (C_a) has risen dramatically since preglacial times and is projected to double in the next century. As part of a 4‐year study, we examined leaf gas exchange and photosynthetic acclimation in C₃ and C₄ plants using unique chambers that maintained a continuous C_a gradient from 200 to 550 µmol mol^?1 in a natural grassland. Our goals were to characterize linear, nonlinear and threshold responses to increasing C_a from past to future C_a levels. Photosynthesis (A), stomatal conductance (g_s), leaf water‐use efficiency (A/g_s) and leaf N content were measured in three common species: Bothriochloa ischaemum, a C₄ perennial grass, Bromus japonicus, a C₃ annual grass, and Solanum dimidiatum, a C₃ perennial forb. Assimilation responses to internal CO₂ concentrations (A/C_i curves) and photosynthetically active radiation (A/PAR curves) were also assessed, and acclimation parameters estimated from these data. Photosynthesis increased linearly with C_a in all species (P < 0.05). S. dimidiatum and B. ischaemum had greater carboxylation rates for Rubisco and PEP carboxylase, respectively, at subambient than superambient C_a (P < 0.05). To our knowledge, this is the first published evidence of A up‐regulation at subambient C_a in the field. No species showed down‐regulation at superambient C_a. Stomatal conductance generally showed curvilinear decreases with C_a in the perennial species (P < 0.05), with steeper declines over subambient C_a than superambient, suggesting that plant water relations have already changed significantly with past C_a increases. Resource‐use efficiency (A/g_s and A/leaf N) in all species increased linearly with C_a. As both C₃ and C₄ plants had significant responses in A, g_s, A/g_s and A/leaf N to C_a enrichment, future C_a increases in this grassland may not favour C₃ species as much as originally thought. Non‐linear responses and acclimation to low C_a should be incorporated into mechanistic models to better predict the effects of past and present rising C_a on grassland ecosystems.     

A branch bag technique for simultaneous CO2 enrichment and assimilation measurements on beech (Fagus sylvatica L.)

A cheap CO₂ enrichment system was designed to perform continuous gas exchange measurements of branches of mature European beech trees (Fagus sylvatica L.). Branches were grown at ambient (350 cm³ m^-3) and elevated CO₂ (700cm³ m^-3) during the whole 1992 leafy period. Leaks resulting from airtightness defaults in the system appeared to be low enough to measure accurately net CO₂ assimilation and transpiration rates during the day. However, the CO₂ exchange rates during the night (respiration) were too low to allow accurate measurements. Elevated CO₂ had a great effect on the net assimilation rate of branches via its influence on both the C₃ photosynthetic pathway and the shade-tolerance of beech trees (85% increase). The A/Ca curves showed no acclimation effect to high CO₂, both control and enriched branches increasing their net assimilation in the same way. The decrease of net assimilation rates in mature leaves was similar for both control and enriched branches. The pattern of daily transpiration rates remained the same for both control and enriched branches, hence we can assume that there was no visible CO₂ effect on stomata.     

CO_2增长对杉木中龄林针叶光合生理生态的影响

通过对 1 7a生杉木人工林小枝的活体测定 ,研究了大气 CO2 增长对杉木中龄林净光合、呼吸、气孔导度和水分利用效率等生理生态特性的影响。结果表明 ,在 CO2 浓度为 4 5μmol/ L左右时 ,杉木针叶净光合速率比正常大气 CO2 下提高 1倍以上 ,气孔导度和蒸腾速率有不同程度的降低 ,水分利用效率提高约 1～ 2倍 ;同时使光补偿点降低 ,饱和点和光抑制点提高 ,光量子效率提高 4 0 %～ 2 95%且阳枝大于阴枝 ,针叶暗呼吸降低 2 0 %～ 72 %。随着 CO2 浓度的增加 ,针叶净光合速率和水分利用效率呈线性上升 ,约到 4 5μmol/ L以后 ,增加速率减慢 ,但 CO2 饱和点可达 1 1 6μmol/ L以上。杉木针叶对CO2 增长的这种反应 ,对大气 CO2 不断增长的条件下杉木生长是有利的 ,但对其长期反应还有待进一步研究。     

CO2增长对杉木中龄林针叶光合生理生态的影响

通过对１７ａ生杉木人工林小枝的活体测定,研究了大气ＣＯ２增长对杉木中龄林净光合、呼吸、气孔导度和水分利用效率等生理生态特性的影响。结果表明,在ＣＯ２浓度为４５μｍｏｌ／Ｌ左右时,杉木针叶净光合速率比正常大气ＣＯ２下提高１倍以卢,气孔导度和蒸速率有不同程度的降低,水分利用效率提高约１￣２倍;同时使光补偿点降低,饱和点和光抑制点提高,光量子效率提高４０％￣２９５％且阳酝 大于阴枝,针叶暗呼吸降低２０％     

Stomatal behaviour, photosynthesis and transpiration under rising CO2

Definitions of the variables used and the units are given in Table 1

The literature reports enormous variation between species in the extent of stomatal responses to rising CO₂. This paper attempts to provide a framework within which some of this diversity can be explained. We describe the role of stomata in the short-term response of leaf gas exchange to increases in ambient CO₂ concentration by developing the recently proposed stomatal model of Jarvis & Davies (1998 ). In this model stomatal conductance is correlated with the functioning of the photosynthetic system so that the effects of increases in CO₂ on stomata are experienced through changes in the rate of photosynthesis in a simple and mechanistically transparent way. This model also allows us to consider the effects of evaporative demand and soil moisture availability on stomatal responses to photosynthesis and therefore provides a means of considering these additional sources of variation. We emphasize that the relationship between the rate of photosynthesis and the internal CO₂ concentration and also drought will have important effects on the relative gains to be achieved under rising CO₂.  

  Table 1 . Abbreviations     

8.

两个品种转基因抗虫棉光合生理的CO2响应   总被引：1，自引：0，他引：1       下载免费PDF全文

孙彩霞  郝健均  王杰  缪璐  陈振华  齐华 《生态学报》2010,30(2):504-510

栽培环境条件的改变会对转基因作物产生深远影响。以2种不同转基因棉花及其亲本对照为材料,研究了盆栽种植条件下不同棉花品种在蕾期和吐絮期光合生理特性CO2响应特征。结果表明,与各自的常规棉对照比较,两种转基因抗虫棉单叶净光合速率CO2响应的特征参数表观初始羧化效率(CE)、表观暗呼吸速率(Rd)和最大净光合速率(Pmax,c)虽有一定程度的变化,但其差异均未达到显著水平。在高CO2浓度范围内(700μmol.mol-1),转基因抗虫棉单叶净光合速率和水分利用率(WUE)的CO2响应曲线特征发生变化,且与品种及生育时期有关。两种转基因抗虫棉在不同生育时期的气孔导度(Gs)对CO2浓度的响应特征与其常规棉对照相似,短期CO2浓度增高对转基因抗虫棉的气孔导度没有显著性影响。     

9.

Stomatal and non-stomatal limitation to photosynthesis in two trembling aspen (Populus tremuloides Michx.) clones exposed to elevated CO₂ and/or O₃   总被引：3，自引：2，他引：3  

A. Noormets  A. Sôber  E. J. Pell  R. E. Dickson  G. K. Podila  J. Sôber  J. G. Isebrands  & D. F. Karnosky 《Plant, cell & environment》2001,24(3):327-336

Leaf gas exchange parameters and the content of ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Rubisco) in the leaves of two 2‐year‐old aspen (Populus tremuloides Michx.) clones (no. 216, ozone tolerant and no. 259, ozone sensitive) were determined to estimate the relative stomatal and mesophyll limitations to photosynthesis and to determine how these limitations were altered by exposure to elevated CO₂ and/or O₃. The plants were exposed either to ambient air (control), elevated CO₂ (560 p.p.m.) elevated O₃ (55 p.p.b.) or a mixture of elevated CO₂ and O₃ in a free air CO₂ enrichment (FACE) facility located near Rhinelander, Wisconsin, USA. Light‐saturated photosynthesis and stomatal conductance were measured in all leaves of the current terminal and of two lateral branches (one from the upper and one from the lower canopy) to detect possible age‐related variation in relative stomatal limitation (leaf age is described as a function of leaf plastochron index). Photosynthesis was increased by elevated CO₂ and decreased by O₃ at both control and elevated CO₂. The relative stomatal limitation to photosynthesis (l_s) was in both clones about 10% under control and elevated O₃. Exposure to elevated CO₂ + O₃ in both clones and to elevated CO₂ in clone 259, decreased l_s even further – to about 5%. The corresponding changes in Rubisco content and the stability of C_i/C_a ratio suggest that the changes in photosynthesis in response to elevated CO₂ and O₃ were primarily triggered by altered mesophyll processes in the two aspen clones of contrasting O₃ tolerance. The changes in stomatal conductance seem to be a secondary response, maintaining stable C_i under the given treatment, that indicates close coupling between stomatal and mesophyll processes.     

10.

Ontogeny affects response of northern red oak seedlings to elevated CO₂ and water stress: I. Carbon assimilation and biomass production     

PAUL D. ANDERSON  & PATRICIA T. TOMLINSON 《The New phytologist》1998,140(3):477-491

     

11.

Stomatal acclimation over a subambient to elevated CO₂ gradient in a C₃/C₄ grassland   总被引：1，自引：1，他引：1  

H. Maherali  C. D. Reid  H. W. Polley  H. B. Johnson  & R. B. Jackson 《Plant, cell & environment》2002,25(4):557-566

An investigation to determine whether stomatal acclimation to [CO₂] occurred in C₃/C₄ grassland plants grown across a range of [CO₂] (200–550 µmol mol^?1) in the field was carried out. Acclimation was assessed by measuring the response of stomatal conductance (g_s) to a range of intercellular CO₂ (a g_s–C_i curve) at each growth [CO₂] in the third and fourth growing seasons of the treatment. The g_s–C_i response curves for Solanum dimidiatum (C₃ perennial forb) differed significantly across [CO₂] treatments, suggesting that stomatal acclimation had occurred. Evidence of non–linear stomatal acclimation to [CO₂] in this species was also found as maximum g_s (g_smax; g_s measured at the lowest C_i) increased with decreasing growth [CO₂] only below 400 µmol mol^?1. The substantial increase in g_s at subambient [CO₂] for S. dimidiatum was weakly correlated with the maximum velocity of carboxylation (V_cmax; r² = 0·27) and was not associated with CO₂ saturated photosynthesis (A_max). The response of g_s to C_i did not vary with growth [CO₂] in Bromus japonicus (C₃ annual grass) or Bothriochloa ischaemum (C₄ perennial grass), suggesting that stomatal acclimation had not occurred in these species. Stomatal density, which increased with rising [CO₂] in both C₃ species, was not correlated with g_s. Larger stomatal size at subambient [CO₂], however, may be associated with stomatal acclimation in S. dimidiatum. Incorporating stomatal acclimation into modelling studies could improve the ability to predict changes in ecosystem water fluxes and water availability with rising CO₂ and to understand their magnitudes relative to the past.     

12.

  总被引：1，自引：0，他引：1  

H. MAHERALI  H. B. JOHNSON  & R. B. JACKSON 《Plant, cell & environment》2003,26(8):1297-1306

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.     

13.

Predicting leaf gas exchange and δ¹³C responses to the past 30000 years of global environmental change     

D. J. BEERLING 《The New phytologist》1994,128(3):425-433

     

14.

Effects of elevated carbon dioxide concentration on photosynthesis and growth of small birch plants (Betula pendula Roth.) at optimal nutrition   总被引：3，自引：3，他引：3  

R. PETTERSSON  A. J. S. McDONALD 《Plant, cell & environment》1992,15(8):911-919

Small birch plants (Betula pendula Roth.) were grown from seed for periods of up to 70d in a climate chamber at optimal nutrition and at present (350 μmol mol^?1) or elevated (700 μmol mol^?1) concentrations of atmospheric CO₂. Nutrients were sprayed over the roots in Ingestad-type units. Relative growth rate and net assimilation rate were slightly higher at elevated CO₂, whereas leaf area ratio was slightly lower. Smaller leaf area ratio was associated with lower values of specific leaf area. Leaves grown at elevated CO₂ had higher starch concentrations (dry weight basis) than leaves grown at present levels of CO₂. Biomass allocation showed no change with CO₂, and no large effects on stem height, number of side shoots and number of leaves were found. However, the specific root length of fine roots was higher at elevated CO₂. No large difference in the response of carbon assimilation to intercellular CO₂ concentration (A/C_i curves) were found between CO₂ treatments. When measured at the growth environments, the rates of photosynthesis were higher in plants grown at elevated CO₂ than in plants grown at present CO₂. Water use efficiency of single leaves was higher in the elevated treatment. This was mainly attributable to higher carbon assimilation rate at elevated CO₂. The difference in water use efficiency diminished with leaf age. The small treatment difference in relative growth rate was maintained throughout the experiment, which meant that the difference in plant size became progressively greater. Thus, where plant nutrition is sufficient to maintain maximum growth, small birch plants may potentially increase in size more rapidly at elevated CO₂.     

15.

    

Steven L. Voelker  J. Renée Brooks  Frederick C. Meinzer  Rebecca Anderson  Martin K.‐F. Bader  Giovanna Battipaglia  Katie M. Becklin  David Beerling  Didier Bert  Julio L. Betancourt  Todd E. Dawson  Jean‐Christophe Domec  Richard P. Guyette  Christian Körner  Steven W. Leavitt  Sune Linder  John D. Marshall  Manuel Mildner  Jérôme Ogée  Irina Panyushkina  Heather J. Plumpton  Kurt S. Pregitzer  Matthias Saurer  Andrew R. Smith  Rolf T. W. Siegwolf  Michael C. Stambaugh  Alan F. Talhelm  Jacques C. Tardif  Peter K. Van de Water  Joy K. Ward  Lisa Wingate 《Global Change Biology》2016,22(2):889-902

Rising atmospheric [CO₂], c_a, is expected to affect stomatal regulation of leaf gas‐exchange of woody plants, thus influencing energy fluxes as well as carbon (C), water, and nutrient cycling of forests. Researchers have proposed various strategies for stomatal regulation of leaf gas‐exchange that include maintaining a constant leaf internal [CO₂], c_i, a constant drawdown in CO₂ (c_a ? c_i), and a constant c_i/c_a. These strategies can result in drastically different consequences for leaf gas‐exchange. The accuracy of Earth systems models depends in part on assumptions about generalizable patterns in leaf gas‐exchange responses to varying c_a. The concept of optimal stomatal behavior, exemplified by woody plants shifting along a continuum of these strategies, provides a unifying framework for understanding leaf gas‐exchange responses to c_a. To assess leaf gas‐exchange regulation strategies, we analyzed patterns in c_i inferred from studies reporting C stable isotope ratios (δ¹³C) or photosynthetic discrimination (?) in woody angiosperms and gymnosperms that grew across a range of c_a spanning at least 100 ppm. Our results suggest that much of the c_a‐induced changes in c_i/c_a occurred across c_a spanning 200 to 400 ppm. These patterns imply that c_a ? c_i will eventually approach a constant level at high c_a because assimilation rates will reach a maximum and stomatal conductance of each species should be constrained to some minimum level. These analyses are not consistent with canalization toward any single strategy, particularly maintaining a constant c_i. Rather, the results are consistent with the existence of a broadly conserved pattern of stomatal optimization in woody angiosperms and gymnosperms. This results in trees being profligate water users at low c_a, when additional water loss is small for each unit of C gain, and increasingly water‐conservative at high c_a, when photosystems are saturated and water loss is large for each unit C gain.     

16.

Development of Water Stress under Increased Atmospheric CO2 Concentration     

J. Pospíšilová  J. Čatský 《Biologia Plantarum》1999,42(1):1-24

The increase in water use efficiency (the ratio of photosynthetic to transpiration rates) is likely to be the commonest positive effect of long-term elevation in CO₂ concentration (CE). This may not necessarily lead to decrease in long-term water use owing to increased leaf area. However, some plant species seem to cope better with drought stress under CE, because increased production of photosynthates might enhance osmotic adjustment and decreased stomatal conductance and transpiration rate under CE enable plants to maintain a higher leaf water potential during drought. In addition, at the same stomatal conductance, internal CO₂ concentration might be higher under CE which results in higher photosynthetic rate. Therefore plants under CE of the future atmosphere will probably survive eventual higher drought stress and some species may even be able to extend their biotope into less favourable sites. This revised version was published online in July 2006 with corrections to the Cover Date.     

17.

    

Lasse Tarvainen  Mats Räntfors  Göran Wallin 《Plant, cell & environment》2015,38(11):2487-2496

Previous leaf‐scale studies of carbon assimilation describe short‐term resource‐use efficiency (RUE) trade‐offs where high use efficiency of one resource requires low RUE of another. However, varying resource availabilities may cause long‐term RUE trade‐offs to differ from the short‐term patterns. This may have important implications for understanding canopy‐scale resource use and allocation. We used continuous gas exchange measurements collected at five levels within a Norway spruce, Picea abies (L.) karst., canopy over 3 years to assess seasonal differences in the interactions between shoot‐scale resource availability (light, water and nitrogen), net photosynthesis (A_n) and the use efficiencies of light (LUE), water (WUE) and nitrogen (NUE) for carbon assimilation. The continuous data set was used to develop and evaluate multiple regression models for predicting monthly shoot‐scale A_n. These models showed that shoot‐scale A_n was strongly dependent on light availability and was generally well described with simple one‐ or two‐parameter models. WUE peaked in spring, NUE in summer and LUE in autumn. However, the relative importance of LUE for carbon assimilation increased with canopy depth at all times. Our results suggest that accounting for seasonal and within‐canopy trade‐offs may be important for RUE‐based modelling of canopy carbon uptake.     

18.

    

Dafeng Hui  Yiqi Luo  Weixin Cheng  J. S. Coleman  Dale W. Johnson  Daniel A. Sims 《Global Change Biology》2001,7(1):75-91

This study used an environmentally controlled plant growth facility, EcoCELLs, to measure canopy gas exchanges directly and to examine the effects of elevated [CO₂] on canopy radiation‐ and water‐use efficiencies. Sunflowers (Helianthus annus var. Mammoth) were grown at ambient (399 μmol mol^?1) and elevated [CO₂] (746 μmol mol^?1) for 53 days in EcoCELLs. Whole canopy carbon‐ and water‐fluxes were measured continuously during the period of the experiment. The results indicated that elevated [CO₂] enhanced daily total canopy carbon‐ and water‐fluxes by 53% and 11%, respectively, on a ground‐area basis, resulting in a 54% increase in radiation‐use efficiency (RUE) based on intercepted photosynthetic active radiation and a 26% increase in water‐use efficiency (WUE) by the end of the experiment. Canopy carbon‐ and water‐fluxes at both CO₂ treatments varied with canopy development. They were small at 22 days after planting (DAP) and gradually increased to the maxima at 46 DAP. When canopy carbon‐ and water‐fluxes were expressed on a leaf‐area basis, no effect of CO₂ was found for canopy water‐flux while elevated [CO₂] still enhanced canopy carbon‐flux by 29%, on average. Night‐time canopy carbon‐flux was 32% higher at elevated than at ambient [CO₂]. In addition, RUE and WUE displayed strong diurnal variations, high at noon and low in the morning or afternoon for WUE but opposite for RUE. This study provided direct evidence that plant canopy may consume more, instead of less, water but utilize both water and radiation more efficiently at elevated than at ambient [CO₂], at least during the exponential growth period as illustrated in this experiment.     

19.

    

Melissa A. Pastore  Tali D. Lee  Sarah E. Hobbie  Peter B. Reich 《Plant, cell & environment》2020,43(8):1862-1878

Global changes can interact to affect photosynthesis and thus ecosystem carbon capture, yet few multi-factor field studies exist to examine such interactions. Here, we evaluate leaf gas exchange responses of five perennial grassland species from four functional groups to individual and interactive global changes in an open-air experiment in Minnesota, USA, including elevated CO₂ (eCO₂), warming, reduced rainfall and increased soil nitrogen supply. All four factors influenced leaf net photosynthesis and/or stomatal conductance, but almost all effects were context-dependent, i.e. they differed among species, varied with levels of other treatments and/or depended on environmental conditions. Firstly, the response of photosynthesis to eCO₂ depended on species and nitrogen, became more positive as vapour pressure deficit increased and, for a C₄ grass and a legume, was more positive under reduced rainfall. Secondly, reduced rainfall increased photosynthesis in three functionally distinct species, potentially via acclimation to low soil moisture. Thirdly, warming had positive, neutral or negative effects on photosynthesis depending on species and rainfall. Overall, our results show that interactions among global changes and environmental conditions may complicate predictions based on simple theoretical expectations of main effects, and that the factors and interactions influencing photosynthesis vary among herbaceous species.     

20.

Stomatal conductance of forest species after long-term exposure to elevated CO₂ concentration: a synthesis   总被引：9，自引：4，他引：9  

B. E. Medlyn  C. V. M. Barton  M. S. J. Broadmeadow  R. Ceulemans  P. De Angelis  M. Forstreuter  M. Freeman  S. B. Jackson  S. Kellomäki  E. Laitat  A. Rey  P. Roberntz  B. D. Sigurdsson  J. Strassemeyer  K. Wang  P. S. Curtis  P. G. Jarvis 《The New phytologist》2001,149(2):247-264

     

两个品种转基因抗虫棉光合生理的CO2响应

栽培环境条件的改变会对转基因作物产生深远影响。以2种不同转基因棉花及其亲本对照为材料,研究了盆栽种植条件下不同棉花品种在蕾期和吐絮期光合生理特性CO2响应特征。结果表明,与各自的常规棉对照比较,两种转基因抗虫棉单叶净光合速率CO2响应的特征参数表观初始羧化效率(CE)、表观暗呼吸速率(Rd)和最大净光合速率(Pmax,c)虽有一定程度的变化,但其差异均未达到显著水平。在高CO2浓度范围内(700μmol.mol-1),转基因抗虫棉单叶净光合速率和水分利用率(WUE)的CO2响应曲线特征发生变化,且与品种及生育时期有关。两种转基因抗虫棉在不同生育时期的气孔导度(Gs)对CO2浓度的响应特征与其常规棉对照相似,短期CO2浓度增高对转基因抗虫棉的气孔导度没有显著性影响。     

Stomatal and non-stomatal limitation to photosynthesis in two trembling aspen (Populus tremuloides Michx.) clones exposed to elevated CO2 and/or O3

Leaf gas exchange parameters and the content of ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Rubisco) in the leaves of two 2‐year‐old aspen (Populus tremuloides Michx.) clones (no. 216, ozone tolerant and no. 259, ozone sensitive) were determined to estimate the relative stomatal and mesophyll limitations to photosynthesis and to determine how these limitations were altered by exposure to elevated CO₂ and/or O₃. The plants were exposed either to ambient air (control), elevated CO₂ (560 p.p.m.) elevated O₃ (55 p.p.b.) or a mixture of elevated CO₂ and O₃ in a free air CO₂ enrichment (FACE) facility located near Rhinelander, Wisconsin, USA. Light‐saturated photosynthesis and stomatal conductance were measured in all leaves of the current terminal and of two lateral branches (one from the upper and one from the lower canopy) to detect possible age‐related variation in relative stomatal limitation (leaf age is described as a function of leaf plastochron index). Photosynthesis was increased by elevated CO₂ and decreased by O₃ at both control and elevated CO₂. The relative stomatal limitation to photosynthesis (l_s) was in both clones about 10% under control and elevated O₃. Exposure to elevated CO₂ + O₃ in both clones and to elevated CO₂ in clone 259, decreased l_s even further – to about 5%. The corresponding changes in Rubisco content and the stability of C_i/C_a ratio suggest that the changes in photosynthesis in response to elevated CO₂ and O₃ were primarily triggered by altered mesophyll processes in the two aspen clones of contrasting O₃ tolerance. The changes in stomatal conductance seem to be a secondary response, maintaining stable C_i under the given treatment, that indicates close coupling between stomatal and mesophyll processes.     

Stomatal acclimation over a subambient to elevated CO2 gradient in a C3/C4 grassland

An investigation to determine whether stomatal acclimation to [CO₂] occurred in C₃/C₄ grassland plants grown across a range of [CO₂] (200–550 µmol mol^?1) in the field was carried out. Acclimation was assessed by measuring the response of stomatal conductance (g_s) to a range of intercellular CO₂ (a g_s–C_i curve) at each growth [CO₂] in the third and fourth growing seasons of the treatment. The g_s–C_i response curves for Solanum dimidiatum (C₃ perennial forb) differed significantly across [CO₂] treatments, suggesting that stomatal acclimation had occurred. Evidence of non–linear stomatal acclimation to [CO₂] in this species was also found as maximum g_s (g_smax; g_s measured at the lowest C_i) increased with decreasing growth [CO₂] only below 400 µmol mol^?1. The substantial increase in g_s at subambient [CO₂] for S. dimidiatum was weakly correlated with the maximum velocity of carboxylation (V_cmax; r² = 0·27) and was not associated with CO₂ saturated photosynthesis (A_max). The response of g_s to C_i did not vary with growth [CO₂] in Bromus japonicus (C₃ annual grass) or Bothriochloa ischaemum (C₄ perennial grass), suggesting that stomatal acclimation had not occurred in these species. Stomatal density, which increased with rising [CO₂] in both C₃ species, was not correlated with g_s. Larger stomatal size at subambient [CO₂], however, may be associated with stomatal acclimation in S. dimidiatum. Incorporating stomatal acclimation into modelling studies could improve the ability to predict changes in ecosystem water fluxes and water availability with rising CO₂ and to understand their magnitudes relative to the past.     

Effects of elevated carbon dioxide concentration on photosynthesis and growth of small birch plants (Betula pendula Roth.) at optimal nutrition

Small birch plants (Betula pendula Roth.) were grown from seed for periods of up to 70d in a climate chamber at optimal nutrition and at present (350 μmol mol^?1) or elevated (700 μmol mol^?1) concentrations of atmospheric CO₂. Nutrients were sprayed over the roots in Ingestad-type units. Relative growth rate and net assimilation rate were slightly higher at elevated CO₂, whereas leaf area ratio was slightly lower. Smaller leaf area ratio was associated with lower values of specific leaf area. Leaves grown at elevated CO₂ had higher starch concentrations (dry weight basis) than leaves grown at present levels of CO₂. Biomass allocation showed no change with CO₂, and no large effects on stem height, number of side shoots and number of leaves were found. However, the specific root length of fine roots was higher at elevated CO₂. No large difference in the response of carbon assimilation to intercellular CO₂ concentration (A/C_i curves) were found between CO₂ treatments. When measured at the growth environments, the rates of photosynthesis were higher in plants grown at elevated CO₂ than in plants grown at present CO₂. Water use efficiency of single leaves was higher in the elevated treatment. This was mainly attributable to higher carbon assimilation rate at elevated CO₂. The difference in water use efficiency diminished with leaf age. The small treatment difference in relative growth rate was maintained throughout the experiment, which meant that the difference in plant size became progressively greater. Thus, where plant nutrition is sufficient to maintain maximum growth, small birch plants may potentially increase in size more rapidly at elevated CO₂.     

Development of Water Stress under Increased Atmospheric CO2 Concentration

The increase in water use efficiency (the ratio of photosynthetic to transpiration rates) is likely to be the commonest positive effect of long-term elevation in CO₂ concentration (CE). This may not necessarily lead to decrease in long-term water use owing to increased leaf area. However, some plant species seem to cope better with drought stress under CE, because increased production of photosynthates might enhance osmotic adjustment and decreased stomatal conductance and transpiration rate under CE enable plants to maintain a higher leaf water potential during drought. In addition, at the same stomatal conductance, internal CO₂ concentration might be higher under CE which results in higher photosynthetic rate. Therefore plants under CE of the future atmosphere will probably survive eventual higher drought stress and some species may even be able to extend their biotope into less favourable sites. This revised version was published online in July 2006 with corrections to the Cover Date.