Increase of photosynthesis and starch in potato under elevated CO2 is dependent on leaf age

Potato plants (Solanum tuberosum cv. Bintje) were grown in open top chambers under ambient (400 microL L(-1)) and elevated CO2 (720 microL L(-1)). After 50 days one half of each group was transferred to the other CO2 concentration and the effects were studied in relation to leaf age (old, middle-aged and young leaves) in each of the four groups. Under long-term exposure to elevated CO2, photosynthesis increased between 10% and 40% compared to ambient CO2. A subsequent shift of the same plants to ambient CO2 caused a 20-40% decline in photosynthetic rate, which was most pronounced in young leaves. After shifting from long-term ambient to elevated CO2, photosynthesis also increased most strongly in young leaves (90%); these experiments show that photosynthesis was downregulated in the upper young fully expanded leaves of potato growing long-term under elevated CO2. Soluble sugar content in all leaf classes under long-term exposure was stable irrespective of the CO2 treatment, however under elevated CO2 young leaves showed a strongly increased starch accumulation (up to 400%). In all leaf classes starch levels dropped in response to the shift from 720 to 400 microL L(-1) approaching ambient CO2 levels. After the shift to 720 microL L(-1), sucrose and starch levels increased, principally in young Leaves. There is clear evidence that leaves of different age vary in their responses to changes in atmospheric CO2 concentration.     

Effects of Elevated CO2 and High Temperature on Single Leaf and Canopy Photosynthesis of Rice

The increase of atmospheric CO2 concentration is indisputable. In such condition, photosynthetic response of leaf is relatively well studied, while the comparison of that between single leaf and whole canopy is less emphasized. The stimulation of elevated CO2 on canopy photosynthesis may be different from that on single leaf level. In this study, leaf and canopy photosynthesis of rice ( Oryza sativa L. ) were studied throughout the growing season. High CO2 and temperature had a synergetic stimulation on single leaf photosynthetic rate until grain filling. Photosynthesis of leaf was stimulated by high CO2, although the stimulation was decreased by higher temperature at grain filling stage. On the other hand, the simulation of elevated CO2 on canopy photosynthesis leveled off with time. Stimulation at canopy level disappeared by grain filling stage in beth temperature treatments. Green leaf area index was not significantly affected by CO2 at maturity, but greater in plants grown at higher temperature. Leaf nitrogen content decreased with the increase of CO2 concentration although it was not statistically significant at maturity. Canopy respiration rate increased at flowering stage indicating higher carbon loss. Shading effect caused by leaf development reached maximum at flowering stage. The CO2 stimulation on photosynthesis was greater in single leaf than in canopy. Since enhanced CO2 significantly increased biomass of rice stems and panicles, increase in canopy respiration caused diminishment of CO2 stimulation in canopy net photosynthesis, keaf nitrogen in the canopy level decreased with CO2 concentration and may eventually hasten CO2 stimulation on canopy photosynthesis. Early senescence of canopy leaves in high CO2 is also a possible cause.     

高浓度二氧化碳对植物影响的研究进展

工业革命后全球大气CO2浓度持续上升,不仅对全球气候的变迁产生重大影响,而且对植物的形态、水分利用、蛋白质合成、光合、抗性、生长及生物量等都有不同程度的影响。高浓度CO2促进植物根、幼苗的生长,叶片增厚,降低气孔密度、气孔导度及蒸腾速率,增加水分利用效率、作物的产量及生物量,促进乙烯生物合成,增强植物的抗氧化能力。不同光合途径(C3、C4及CAM)及不同植被类型的植物对高浓度CO2的响应不同。长期和短期的高浓度CO2处理,植物响应方式有很大的差异,如短期高CO2处理使光合能力增强,而长期处理则使光合能力下调。     

大气CO2浓度和温度升高对水稻叶片及群体光合作用的影响

大气ＣＯ２浓度升高对植物光合作用的影响研究多集中在单叶水平,在高ＣＯ２及高温下对植物单叶及群体光合进行比较的研究少有报道,而群体水平的研究则是预测生态系统反应所不可缺少的。采用田间开顶式培养室研究了大气ＣＯ２浓度和温度升高对水稻（ＯｒｙｚａｓａｔｉｖａＬ．）叶片及群体光合作用的影响。发现ＣＯ２浓度和温度对水稻叶片光合作用有协同促进作用,而对群体光合作用的促进则随时间的推移而减弱;单叶光合受到的促进作用大于群体光合;叶面积指数只在营养生长期受到促进,冠层叶片含氮量受ＣＯ２影响降低。群体呼吸（包括茎杆）增加及冠层叶片早衰可能是后期ＣＯ２对群体光合促进作用下降的原因。     

大气一氧化碳浓度升高对植物生长的影响

大气ＣＯ２浓度同对植物生长有促进作用,对Ｃ３植物生长的促进作用最大。短期ＣＯ２浓度升高时,植物光和速率增加;在长期ＣＯ２浓度升高条件下,植物光鸽上降并发生光合适应现象。这可能是植物在长期ＣＯ２浓度升高条件下植物源库关系不平衡引起的反馈抑制作用以及营养吸收不能满足光合速率增加的需要所引起Ｒｕｂｉｓｅｏ活必和含量下降。在ＣＯ２浓度升高条件下植物的呼吸也会发生变化,根的分枝和数量增多,根系的分泌量和吸收     

Elevated CO(2) induces biochemical and ultrastructural changes in leaves of the C(4) cereal sorghum

We analyzed the impact of growth at either 350 (ambient) or 700 (elevated) microL L(-1) CO(2) on key elements of the C(4) pathway (photosynthesis, carbon isotope discrimination, and leaf anatomy) using the C(4) cereal sorghum (Sorghum bicolor L. Moench.). Gas-exchange analysis of the CO(2) response of photosynthesis indicated that both carboxylation efficiency and the CO(2) saturated rate of photosynthesis were lower in plants grown at elevated relative to ambient CO(2). This was accompanied by a 49% reduction in the phosphoenolpyruvate carboxylase content of leaves (area basis) in the elevated CO(2)-grown plants, but no change in Rubisco content. Despite the lower phosphoenolpyruvate carboxylase content, there was a 3-fold increase in C isotope discrimination in leaves of plants grown at elevated CO(2) and bundle sheath leakiness was estimated to be 24% and 33%, respectively, for the ambient and elevated CO(2)-grown plants. However, we could detect no difference in quantum yield. The ratio of quantum yield of CO(2) fixation to PSII efficiency was lower in plants grown at elevated CO(2), but only when leaf internal was below 50 microL L(-1). This suggests a reduction in the efficiency of the C(4) cycle when [CO(2)] is low, and also implies increased electron transport to acceptors other than CO(2). Analysis of leaf sections using a transmission electron microscope indicated a 2-fold decrease in the thickness of the bundle sheath cell walls in plants grown at elevated relative to ambient CO(2). These results suggest that significant acclimation to increased CO(2) concentrations occurs in sorghum.     

Experimental Study and Mathematical Simulation of Water Use Efficiency in Wheat Leaves Affected by Some Environmental Factors

By incorporating Ball-Berry model of stomatal conductance into the models of photosynthesis and transpiration, a model of leaf water use efficiency (WUE) as affected by several environmental variables [irradiance (Ⅰ), vapor pressure deficit (VPD) and atmospheric CO2 concentration (Ca) ] was constructed. Because the environmental variables influenced the photosynthetic rate and transpiration rate in different ways, the changes of leaf WUE with these factors were quite complicated. The rates of photosynthesis and transpiration of wheat leaves were also measured in the phytotron where the environmental variables were kept within certain ranges, and leaf WUE was calculated therefrom. The results of simulation fit quite well with the measurements except at high Ca.     

Major diffusion leaks of clamp‐on leaf cuvettes still unaccounted: how erroneous are the estimates of Farquhar et al. model parameters?

Estimates of leaf gas-exchange characteristics using standard clamp-on leaf chambers are prone to errors because of diffusion leaks. While some consideration has been given to CO(2) diffusion leaks, potential water vapour diffusion leaks through chamber gaskets have been neglected. We estimated diffusion leaks of two clamp-on Li-Cor LI-6400 (Li-Cor, Inc., Lincoln, NE, USA) leaf chambers with polymer foam gaskets and enclosing either 2 or 6 cm(2) leaf area, and conducted a sensitivity analysis of the diffusion leak effects on Farquhar et al. photosynthesis model parameters - the maximum carboxylase activity of ribulose 1 x 5-bisphosphate carboxylase/oxygenase (Rubisco) (V(cmax)), capacity for photosynthetic electron transport (J(max)) and non-photorespiratory respiration rate in light (R(d)). In addition, net assimilation rate (A(n)) versus intercellular CO(2) (C(i)) responses were measured in leaves of Mediterranean evergreen species Quercus ilex L. enclosing the whole leaf chamber in a polyvinyl fluoride bag flushed with the exhaust air of leaf chamber, thereby effectively reducing the CO(2) and water vapour gradients between ambient air and leaf chamber. For the empty chambers, average diffusion leak for CO(2), K(CO2), (molar flow rate corresponding to unit CO(2) mole fraction difference) was ca. 0.40 micromol s(-1). K(CO2) increased ca. 50% if a dead leaf was clamped between the leaf chamber. Average diffusion leak for H(2)O was ca. 5- to 10-fold larger than the diffusion leak for CO(2). Sensitivity analyses demonstrated that the consequence of a CO(2) diffusion leak was apparent enhancement of A(n) at high CO(2) mole fraction and reduction at lower CO(2) mole fraction, and overall compression of C(i) range. As the result of these modifications, Farquhar et al. model parameters were overestimated. The degree of overestimation increased in the order of V(cmax) < J(max) < R(d), and was larger for smaller chambers and for leaves with lower photosynthetic capacity, leading to overestimation of all three parameters by 70-290% for 2 cm(2), and by 10-60% for 6 cm(2) chamber. Significant diffusion corrections (5-36%) were even required for leaves with high photosynthetic capacity measured in largest chamber. Water vapour diffusion leaks further enhanced the overestimation of model parameters. For small chambers and low photosynthetic capacities, apparent C(i) was simulated to decrease with increasing A(n) because of simultaneous CO(2) and H(2)O diffusion leaks. Measurements in low photosynthetic capacity Quercus ilex leaves enclosed in 2 cm(2) leaf chamber exhibited negative apparent C(i) values at highest A(n). For the same leaves measured with the entire leaf chamber enclosed in the polyvinyl fluoride bag, C(i) and A(n) increased monotonically. While the measurements without the bag could be corrected for diffusion leaks, the required correction in A(n) and transpiration rates was 100-500%, and there was large uncertainty in Farquhar et al. model parameters derived from 'corrected'A(n)/C(i) response curves because of uncertainties in true diffusion leaks. These data demonstrate that both CO(2) and water vapour diffusion leaks need consideration in measurements with clamp-on leaf cuvettes. As plants in natural environments are often characterized by low photosynthetic capacities, cuvette designs need to be improved for reliable measurements in such species.     

热带季节雨林冠层树种绒毛番龙眼的光合生理生态特性

采用Li-6400便携式光合作用测定仪,对西双版纳热带季节雨林冠层树种绒毛番龙眼成树树冠上、中、下3层叶片进行了测定,分析西双版纳热带季节雨林冠层树木的光合作用.结果表明,绒毛番龙眼成树具有喜光的光合特性,光饱和点较高(1 000～1 500 μmol·m^-2·s^-1),而光补偿点较低(7.7～15.3 μmol·m^-2·s^-1),对光环境有较强的适应和调节能力,光合有效辐射是影响绒毛番龙眼光合日进程的关键因子;12月,叶片处于成熟期,生长良好,光合能力较强,树冠上层净光合速率（P_n）日变化为单峰型,最大净光合速率（A_max）约为8.9 μmol CO₂·m^-2·s^-1;4月处于新老树叶更替期,光合能力下降,树冠上层P_n日变化为双峰型,中午出现“午休”现象,树冠上层A_max约为4.3 μmol CO₂·m^-2·s^-1;7月上、中层叶片P_n为单峰型,下层出现“午休”.如人为使CO₂浓度在短期内迅速升高,则绒毛番龙眼的P_n会增加,而气孔导度和蒸腾速率降低;CO₂浓度从400 μmol·mol^-1升高到800 μmol·mol^-1时,干季水分利用效率（WUE）提高约50%～100％,雨季WUE较低.     

Interactive effects of elevated C02 and temperature on the anatomical characteristics of leaves in eleven species

The anatomical features of leaves in 11 species of plants grown in a temperature gradient and a temperature + CO2 gradient were studied.The palisade parenchyma thickness,the spongy parenchyma thickness and the total leaf thickness were measured and analyzed to investigate the effects of elevated temperature and CO2 on the anatomical characteristics of the leaves.Our results show that with the increase of temperature,the leaf thickness of C4 species increased while the leaf thickness of C3 species showed no constant changes.With increased CO2,seven out of nine C3 species exhibited increased total leaf thickness.In C4 species,leaf thickness decreased.As for the trend on the multi-grades,the plants exhibited linear or non-linear changes.With the increase of temperature or both temperature and CO2 for the 11 species investigated,leaf thickness varied greatly in different plants (species) and even in different branches on the same plant.These results demonstrated that the effect of increasing CO2 and temperature on the anatomical features of the leaves were species-specific.Since plant structures are correlated with plant functions,the changes in leaf anatomical characteristics in elevated temperature and CO2 may lead to functional differences.     

The response of photosynthesis and stomatal conductance to rising [CO2]: mechanisms and environmental interactions

This review summarizes current understanding of the mechanisms that underlie the response of photosynthesis and stomatal conductance to elevated carbon dioxide concentration ([CO2]), and examines how downstream processes and environmental constraints modulate these two fundamental responses. The results from free-air CO2 enrichment (FACE) experiments were summarized via meta-analysis to quantify the mean responses of stomatal and photosynthetic parameters to elevated [CO2]. Elevation of [CO2] in FACE experiments reduced stomatal conductance by 22%, yet, this reduction was not associated with a similar change in stomatal density. Elevated [CO2] stimulated light-saturated photosynthesis (Asat) in C3 plants grown in FACE by an average of 31%. However, the magnitude of the increase in Asat varied with functional group and environment. Functional groups with ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco)-limited photosynthesis at elevated [CO2] had greater potential for increases in Asat than those where photosynthesis became ribulose-1,5-bisphosphate (RubP)-limited at elevated [CO2]. Both nitrogen supply and sink capacity modulated the response of photosynthesis to elevated [CO2] through their impact on the acclimation of carboxylation capacity. Increased understanding of the molecular and biochemical mechanisms by which plants respond to elevated [CO2], and the feedback of environmental factors upon them, will improve our ability to predict ecosystem responses to rising [CO2] and increase our potential to adapt crops and managed ecosystems to future atmospheric [CO2].     

Carbonic anhydrase and its influence on carbon isotope discrimination during C4 photosynthesis. Insights from antisense RNA in Flaveria bidentis

In C4 plants, carbonic anhydrase (CA) facilitates both the chemical and isotopic equilibration of atmospheric CO2 and bicarbonate (HCO3-) in the mesophyll cytoplasm. The CA-catalyzed reaction is essential for C4 photosynthesis, and the model of carbon isotope discrimination (Delta13C) in C4 plants predicts that changes in CA activity will influence Delta13C. However, experimentally, the influence of CA on Delta13C has not been demonstrated in C4 plants. Here, we compared measurements of Delta13C during C4 photosynthesis in Flaveria bidentis wild-type plants with F. bidentis plants with reduced levels of CA due to the expression of antisense constructs targeted to a putative mesophyll cytosolic CA. Plants with reduced CA activity had greater Delta13C, which was also evident in the leaf dry matter carbon isotope composition (delta13C). Contrary to the isotope measurements, photosynthetic rates were not affected until CA activity was less than 20% of wild type. Measurements of Delta13C, delta13C of leaf dry matter, and rates of net CO2 assimilation were all dramatically altered when CA activity was less than 5% of wild type. CA activity in wild-type F. bidentis is sufficient to maintain net CO2 assimilation; however, reducing leaf CA activity has a relatively large influence on Delta13C, often without changes in net CO2 assimilation. Our data indicate that the extent of CA activity in C4 leaves needs to be taken into account when using Delta13C and/or delta13C to model the response of C4 photosynthesis to changing environmental conditions.     

Influence of Elevated CO2 Concentration and Nitrogen Source on Photosynthetic Traits in the Invasive Species Eupatorium adenophorum (Asteraceae)

Increases in the concentration of atmospheric CO2 and plant invasion are two important problems that face humans worldwide. In some plants, exposure to a short term elevated concentration of CO2 (SE［CO2］) promotes photosynthesis, but the promotion of elevated ［CO2］ (E ［CO2］) to photosynthesis might disappear after long term treatment (so called “CO2 acclimation”); this might result from the associated inhibition of nitrate assimilation. The present study investigated the physiological effects of short term (8 days) and long term (40 days) exposure to E［CO2］ when these were combined with different forms of inorganic N (full N; nitrate (NO3-) N) in the invasive species Eupatorium adenophorum. Exposure to E［CO2］ increased the biomass of Eadenophorum, regardless of the duration of exposure to E［CO2］ and the type of inorganic N that was supplied. E［CO2］ could promote the photosynthesis of Eadenophorum seedlings fertilised with non depleted Hoagland solutions (full N). For plants fertilised with NH4+ depleted Hoagland solution (NO3- N), LE［CO2］ treatment promoted the photosynthesis of Eadenop horum, but the promotion of photosynthesis by E［CO2］ disappeared under SE［CO2］ conditions. Photosynthetic pigments contents were determined to estimate potential changes in the photosynthetic capacity of Eadenophorum. For plants fertilised with non depleted Hoagland solution, there were no significant differences in chlorophyll among the three ［CO2］ treatments, but the treatment of SE［CO2］ increased the levels of chlorophyll in leaves. The apparent promotion of biomass accumulation and photosynthesis at LE［CO2］ without a decrease in chlorophyll indicates that Eadenophorum might not acclimate to long term exposure to E［CO2］. NH4+ depletion did not affect the capacity of LE［CO2］ to promote the photosynthesis of Eadenophorum. Thus, considering some plants fertilised with NO3- acclimating to LE［CO2］, Eadenophorum might be more competitive in areas where the soils are relatively poor in NH4+ as levels of atmospheric CO2 continue to rise.     

Crassulacean acid metabolism enhances underwater photosynthesis and diminishes photorespiration in the aquatic plant Isoetes australis

? Underwater photosynthesis by aquatic plants is often limited by low availability of CO(2), and photorespiration can be high. Some aquatic plants utilize crassulacean acid metabolism (CAM) photosynthesis. The benefits of CAM for increased underwater photosynthesis and suppression of photorespiration were evaluated for Isoetes australis, a submerged plant that inhabits shallow temporary rock pools. ? Leaves high or low in malate were evaluated for underwater net photosynthesis and apparent photorespiration at a range of CO(2) and O(2) concentrations. ? CAM activity was indicated by 9.7-fold higher leaf malate at dawn, compared with at dusk, and also by changes in the titratable acidity (μmol H(+) equivalents) of leaves. Leaves high in malate showed not only higher underwater net photosynthesis at low external CO(2) concentrations but also lower apparent photorespiration. Suppression by CAM of apparent photorespiration was evident at a range of O(2) concentrations, including values below air equilibrium. At a high O(2) concentration of 2.2-fold the atmospheric equilibrium concentration, net photosynthesis was reduced substantially and, although it remained positive in leaves containing high malate concentrations, it became negative in those low in malate. ? CAM in aquatic plants enables higher rates of underwater net photosynthesis over large O(2) and CO(2) concentration ranges in floodwaters, via increased CO(2) fixation and suppression of photorespiration.     

Senescence and hyperspectral reflectance of cotton leaves exposed to ultraviolet-B radiation and carbon dioxide

The objectives of this study were to determine the effects of UV-B radiation and atmospheric carbon dioxide concentrations ([CO(2)]) on leaf senescence of cotton by measuring leaf photosynthesis and chlorophyll content and to identify changes in leaf hyperspectral reflectance occurring due to senescence and UV-B radiation. Plants were grown in controlled-environment growth chambers at two [CO(2)] (360 and 720 micro mol mol(-1)) and three levels of UV-B radiation (0, 7.7 and 15.1 kJ m(-2) day(-1)). Photosynthesis, chlorophyll, carotenoids and phenolic compounds along with leaf hyperspectral reflectance were measured on three leaves aged 12, 21 and 30 days in each of the treatments. No interaction was detected between [CO(2)] and UV-B for any of the measured parameters. Significant interactions were observed between UV-B and leaf age for photosynthesis and stomatal conductance. Elevated [CO(2)] enhanced leaf photosynthesis by 32%. On exposure to 0, 7.7 and 15.1 kJ of UV-B, the photosynthetic rates of 30-day-old leaves compared with 12-day-old leaves were reduced by 52, 76 and 86%, respectively. Chlorophyll pigments were not affected by leaf age at UV-B radiation of 0 and 7.7 kJ, but UV-B of 15.1 kJ reduced the chlorophylls by 20, 60 and 80% in 12, 21 and 30-day-old leaves, respectively. The hyperspectral reflectance between 726 and 1142 nm showed interaction for UV-B radiation and leaf age. In cotton, leaf photosynthesis can be used as an indicator of leaf senescence, as it is more sensitive than photosynthetic pigments on exposure to UV-B radiation. This study revealed that, cotton leaves senesced early on exposure to UV-B radiation as indicated by leaf photosynthesis, and leaf hyperspectral reflectance can be used to detect changes caused by UV-B and leaf ageing.     

开放式空气CO2浓度升高与作物/杂草的竞争关系

CO2浓度升高对植物的光合作用、呼吸作用和水分利用等生理过程产生直接影响,进而影响植物的生长繁殖,CO2浓度升高对于具有C3光合途径的植物较具C4光合途径的植物更为有益,由于许多重要的杂草是C4植物,而许多重要的作用是C3植物,CO2浓度升高对杂草/作物的相互关系将有重要影响,本文就全球CO2浓度升高和气候变化对杂草/作物之间竞争关系影响进行综述,同时针对目前研究现状和可持续农业的需要,提出CO2学浓度升高条件下杂草/作物之间竞争关系及未来农田杂草治理方面理论与实践中有待解决的问题。     

System-level adjustments to elevated CO2 in model spruce ecosystems

Atmospheric carbon dioxide enrichment and increasing nitrogen deposition are often predicted to increase forest productivity based on currently available data for isolated forest tree seedlings or their leaves. However, it is highly uncertain whether such seedling responses will scale to the stand level. Therefore, we studied the effects of increasing CO₂ (280, 420 and 560 μL L^-1) and increasing rates of wet N deposition (0, 30 and 90 kg ha^-1 y^-1) on whole stands of 4-year-old spruce trees (Picea abies). One tree from each of six clones, together with two herbaceous understory species, were established in each of nine 0.7 m² model ecosystems in nutrient poor forest soil and grown in a simulated montane climate for two years. Shoot level light-saturated net photosynthesis measured at growth CO₂ concentrations increased with increasing CO₂, as well as with increasing N deposition. However, predawn shoot respiration was unaffected by treatments. When measured at a common CO₂ concentration of 420 μL L^-1 37% down-regulation of photosynthesis was observed in plants grown at 560 μL CO₂ L^-1. Length growth of shoots and stem diameter were not affected by CO₂ or N deposition. Bud burst was delayed, leaf area index (LAI) was lower, needle litter fall increased and soil CO₂ efflux increased with increasing CO₂. N deposition had no effect on these traits. At the ecosystem level the rate of net CO₂ exchange was not significantly different between CO₂ and N treatments. Most of the responses to CO₂ studied here were nonlinear with the most significant differences between 280 and 420 μL CO₂ L^-1 and relatively small changes between 420 and 560 μL CO₂ L^-1. Our results suggest that the lack of above-ground growth responses to elevated CO₂ is due to the combined effects of physiological down-regulation of photosynthesis at the leaf level, allometric adjustment at the canopy level (reduced LAI), and increasing strength of below-ground carbon sinks. The non-linearity of treatment effects further suggests that major responses of coniferous forests to atmospheric CO₂ enrichment might already be under way and that future responses may be comparatively smaller.     

Specification of adaxial and abaxial stomata, epidermal structure and photosynthesis to CO2 enrichment in maize leaves

Acclimation to CO2 enrichment was studied in maize plants grown to maturity in either 350 or 700 microl l-1 CO2. Plants grown with CO2 enrichment were significantly taller than those grown at 350 microl l-1 CO2 but they had the same number of leaves. High CO2 concentration led to a marked decrease in whole leaf chlorophyll and protein. The ratio of stomata on the adaxial and abaxial leaf surfaces was similar in all growth conditions, but the stomatal index was considerably increased in plants grown at 700 microl l-1 CO2. Doubling the atmospheric CO2 content altered epidermal cell size leading to fewer, much larger cells on both leaf surfaces. The photosynthesis and transpiration rates were always higher on the abaxial surface than the adaxial surface. CO2 uptake rates increased as atmospheric CO2 was increased up to the growth concentrations on both leaf surfaces. Above these values, CO2 uptake on the abaxial surface was either stable or increased as CO2 concentration increased. In marked contrast, CO2 uptake rates on the adaxial surface were progressively inhibited at concentrations above the growth CO2 value, whether light was supplied directly to this or the abaxial surface. These results show that maize leaves adjust their stomatal densities through changes in epidermal cell numbers rather than stomatal numbers. Moreover, the CO2-response curve of photosynthesis on the adaxial surface is specifically determined by growth CO2 abundance and tracks transpiration. Conversely, photosynthesis on the abaxial surface is largely independent of CO2 concentration and rather independent of stomatal function.