FvCB生物化学光合模型及A-C_i曲线测定

由Farquhar、von Caemmerer和Berry提出的生物化学光合模型(以下简称FvCB模型)是一个基于光合碳反应过程的CO_2响应模型。此模型认为C3植物叶片光合速率(A)由3个生物化学过程速率中的最低者——核酮糖-1,5-双磷酸羧化酶/加氧酶(Rubisco)所能支持的羧化速率、电子传递所能支持的核酮糖-1,5-双磷酸(Ru BP)再生速率和磷酸丙糖(TP)利用速率决定。利用改进的FvCB模型对光合速率-胞间CO_2浓度(A-C_i)曲线进行拟合,能有效地估计最大羧化速率、最大电子传递速率、TP利用速率、明呼吸速率、叶肉细胞导度等生化参数,促进我们对植物光合生理及其响应环境变化的理解和预测。该文首先详细地描述了FvCB模型,并分析了此模型分段性和过参数化的特点。然后介绍利用FvCB模型对A-C_i曲线进行拟合,从而估计叶片光合生化参数的研究进展。光合生化参数估计经历了主观分段、分段拟合到客观分段、整体拟合几个阶段,目标函数的最小化方法也从传统的最小二乘法为主转向基于现代计算机技术的迭代算法(如遗传算法、模拟退火算法)。然而,如要进一步提高参数估计的可靠性和精确性,还需加强Rubisco动力学属性和温度依赖性方面的研究。最后,为了获取能更有效地进行参数估计的光合数据,根据目前对FvCB模型拟合的认知,整合并改进了A-C_i曲线的测定方法。     

水分胁迫对小麦叶片光合作用的影响及其与抗旱性的关系

在水分胁迫初期,两个小麦品种叶片光合速率,气孔导度和细胞间隙CO_2浓度降低,气孔限制值增加,光合速率的降低主要是气孔因素的限制。中度到严重水分胁迫使叶片光合速率、气孔导度和气孔限制值降低,细胞间隙CO_2浓度明显增加,且叶圆片放氧能力,叶绿体Hill反应、叶绿素荧光强度和表观量子产额降低,此时光合速率的降低主要是叶肉细胞光合活性的下降引起的。抗旱性弱的郑引一号叶肉细胞光合活性比抗旱性强的丰抗13更容易受到水分胁迫的影响。     

采煤塌陷影响下土壤含水量变化对柠条气孔导度、蒸腾与光合作用速率的影响

采煤塌陷引起的土壤环境因子的变化对矿区植物生长的影响越来越受到人们的关注,气孔导度、蒸腾与光合作用作为环境变化响应的敏感因子,研究植物气孔导度、蒸腾与光合作用的变化是揭示荒漠矿区自然环境变化及其规律的重要手段之一。研究采煤塌陷条件下植物光合生理的变化是探究煤炭开采对植物叶片水分蒸腾散失和CO_2同化速率影响的关键环节,是探讨采煤塌陷影响下植物能量与水分交换动态的基础,而采煤矿区植物叶片气孔导度、蒸腾与光合作用速率对采煤塌陷影响下土壤含水量变化的响应如何尚不清楚。选取神东煤田大柳塔矿区52302工作面为实验场地,以生态修复物种柠条为研究对象,对采煤塌陷区和对照区柠条叶片气孔导度、蒸腾和光合作用速率以及土壤体积含水量进行监测,分析了采煤塌陷条件下土壤含水量的变化以及其对柠条叶片气孔导度、蒸腾与光合作用速率的影响。结果显示:(1)煤炭井工开采在地表形成大量裂缝,破坏了土体结构,潜水位埋深降低,土壤含水量均低于沉陷初期,相对于对照区,硬梁和风沙塌陷区土壤含水量分别降低了18.61%、21.12%;(2)柠条叶片气孔导度、蒸腾和光合作用速率均与土壤含水量呈正相关关系;煤炭开采沉陷增加了地表水分散失,加剧了土壤水分胁迫程度,为了减少蒸腾导致的水分散失,柠条叶片气孔阻力增加,从而气孔导度降低,阻碍了光合作用CO_2的供应,从而导致柠条叶片光合作用速率的降低,蒸腾速率也显著降低。     

红树林植物叶片形态和解剖特征对叶肉导度、叶片导水率的影响

叶肉导度和叶片导水率是影响光合作用的两个重要过程,叶肉导度通过影响从气孔下腔到Rubisco酶位点的二氧化碳浓度梯度直接影响光合作用,而叶片导水率则通过影响水分供应或气孔行为来影响光合作用,然而对这两个生理过程之间的协同性研究较少。本研究选择9种红树林植物为研究对象,探讨盐生环境下植物叶肉导度和叶片导水率的协同性及其与叶片解剖结构特征之间的相关性。结果表明,9种红树林植物叶片导水率(0.78~5.83 mmol·m~(-2)·s~(-1)·MPa-1)、叶肉导度(0.06~0.36 mol·m~(-2)·s~(-1))、最大光合速率(7.23~23.71μmol·m~(-2)·s~(-1))等特征的差别较大;叶肉导度与最大光合速率呈显著正相关,而与比叶重无显著相关性,其原因是由于比叶重与叶片厚度、叶片密度不存在相关性;叶脉密度与气孔密度呈较强的相关性,说明红树林植物叶片水分运输与散失相关的叶片结构之间存在协同关系;叶片导水率不受叶脉密度影响,并且与叶肉导度、最大光合速率也不存在相关性,这很可能与红树林植物叶片的肉质化、有发达的储水组织有关,体现了红树林植物叶片结构和功能的特殊性。     

羌活光合作用日变化及其与生理生态因子的关系

以大田引种栽培的羌活植株为材料,用Li-6400便携式光合作用测定系统原位测定自然条件下生长的羌活孕蕾期叶片的光合速率光响应、净光合速率及生理生态因子日变化,并通过相关分析、通径分析和逐步回归分析探讨净光合速率与生理生态因子的关系.结果表明:(1)羌活净光合速率(P_n)、 蒸腾速率(T_r)、气孔导度(G_s)日变化均呈双峰曲线,其净光合速率具有典型的"午休"现象,并主要由非气孔限制因素造成;(2)影响羌活叶片Pn日变化的主要决定生理因子是G_s,主要限制因子是胞间CO_2浓度(C_i);主要决定生态因子是空气相对湿度(RH),限制因子是气温(T_a); G_s是影响净光合速率最重要的生理生态因子.研究发现,羌活有较强的适应弱光环境的能力,属于阳性耐荫植物,宜选择海拔和荫蔽度均较高的环境进行引种驯化栽培,以利于其生长和存活.     

半干旱黄土丘陵区沙棘的光合特性及其影响因子

结合1998年半干旱黄土丘陵区安塞的观测资料,对沙棘(Hippophae rhamnoides L.)的光合特性及影响因子进行了分析,结果表明(1)沙棘光合速率具有明显的日变化和季节变化,月均值为11.64 CO2μmol/m2·s.(2)沙棘光合速率与环境因子(气温、相对湿度、光合有效辐射及CO2浓度)间有显著的相关关系,复相关系数为0.716 8～0.874 5;沙棘光合速率与林地土壤水分间有极显著的相关关系,复相关系数达0.992 5,且沙棘光合速率的季节动态滞后于土壤水分的月变化.(3)沙棘光合速率与植物因子(气孔导度及细胞间CO2浓度)间有十分显著的相关关系,气孔导度或细胞间CO2浓度增大,沙棘光合速率增大,反之则减小,复相关系数达0.971 5和0.970 8.这为分析沙棘光合作用对环境因子的响应程度,分析沙棘最适生理生态条件,提高沙棘生产力提供科学依据.     

香樟冠层光环境对叶片功能性状和光合特性的影响

为了解香樟冠层不同光环境叶片光合能力的适应机制，提升冠层碳汇功能，本文选取香樟单株冠层南向外侧(100%全光)、南向内侧(34%全光)和北向(21%全光)3个方位的叶片，研究光环境对叶片形态结构、营养和生理性状以及光合特性的影响，分析弱光环境下导致光合能力下调的主要限制因子。结果表明：随着冠层光强减弱，叶片的比叶重、表皮角质层厚度、下表皮厚度、栅栏组织厚度、栅栏组织细胞数和细胞宽度、栅栏组织与海绵组织的厚度比、细胞结构紧密度均显著降低，而海绵组织厚度、栅栏组织细胞长宽比、细胞结构疏松度均显著升高。与全光环境相比，2个弱光环境下叶片的碳含量、可溶性糖、淀粉和可溶性蛋白含量均显著降低，而氮含量在北向显著升高。弱光导致净光合速率(P_n)、暗呼吸速率、CO₂气孔导度(g_sc)、叶肉导度(g_m)、CO₂总导度(g_tot)、胞间CO₂浓度(C_i)、叶绿体CO₂浓度(C_c)等气体交换...     

虎杖光合生理生态特性日变化研究

用Li-6400便携式光合测定系统对5个虎杖材料的光合生理特性日变化及其与气象因子关系进行了研究。结果表明：（1）虎杖的净光合速率日变化呈‘单峰’型曲线,日最大净光合速率（15．0μmol·m^-2·s^-1）值出现在9：00左右;（2）叶片水压亏缺、气孔导度和蒸腾速率的峰值在同一时间出现（13：00）,胞间CO2浓度不随气孔导度的降低而减小,控制虎杖光合速率因子为非气孔限制;（3）供试5个材料的净光合速率日变化趋势基本一致,并以地栽组培苗的净光合速率最高,而‘贵州凯里’的最低。光合有效辐射对光合特征参数的变化影响最大,且对净光合速率起决定性作用（r-0．534^＊＊）,环境因子主要通过对蒸腾速率、叶片水压亏缺和叶面温度的作用来影响虎杖叶片净光合速率。     

基于Farquhar改进模型的北亚热带森林常见树种光合限速因子研究

为探讨北亚热带地区植物的光合限速因子,利用改进的Farquhar模型研究了9种常见树种的光合特性。结果表明,与常绿树种相比,落叶树种枫香(Liquidambar formosana)和乌桕(Sapium sebiferum)的最大净光合速率(Pmax)和表观羧化速率(CE)较大;核酮糖-1,5-二磷酸羧化酶/加氧酶(Rubisco)活性是他们的光合限速因子。地带性树种青冈栎(Cyclobalanopsis glauca)的Pmax和CE在常绿木本植物中最大,青冈较强的光合能力可能是来源于Vcmax和TPU。耐阴灌木八角金盘(Fatsia japonica)和美人茶(Camellia uraku)的Pmax较小,其光合限速因子是叶肉细胞导度和呼吸速率。低光照下植物较低的光合能力是由于较小的叶肉导度(gm)和TPU导致的;有效光合辐射短时间的降低使得物种的gm平均减少了60.14%。因此,不同树种在不同环境条件下的光合限速因子不尽相同,应根据树种不同的光合生理特性来合理布局,科学育林。     

温室茄子（Solanum melongena L.）光合数学模型与光合生化模型模拟分析

针对植物光合与内外环境因子间的关系以及光合“午睡”现象中的气孔限制与非气孔限制问题,以温室茄子‘茄杂一号’为试材,对叶室温光组合方式下测定的净光合速率Pn对胞间CO2浓度Ci响应曲线,和人工增施CO2处理下测定的Pn日变化进程,进行了光合数学模型和Farquhar、von Caemmerer和Berry的光合生化动力学模型（简称为FvCB模型）模拟分析。采用美国思爱迪生态仪器有限公司的CI-301PS光合作用测定仪进行净光合速率（Pn）、光合有效辐射（PAR）、气温（Ta）、叶温（Tl）、环境二氧化碳浓度（Ca）、胞间二氧化碳浓度（Ci）和空气相对湿度（Hr）参数测定。其结果表明,无论是Pn对Ci的响应曲线还是光合日进程中,数学模型对Pn的拟合度明显优于为FvCB模型。因此,通过数学模型可以解析出光合日进程受单一环境因子（PAR、Ta、Ca、Hr）及其复合环境因子的综合影响。然而,FvCB模型模拟结果显示出,温光组合下受Rubisco（即RuBP羧化/加氧酶）数量与活性及动力学特性限制的羧化速率Ac、受RuBP（1,5-二磷酸核酮糖）再生限制的羧化速率Aj以及受TPU（磷酸丙糖）可利用量限制的羧化速率Ap对Ci响应的主控作用呈现交替变化趋势。其交替变化转折点胞间二氧化碳浓度Cicj在强光高温组合中较高,而在弱光低温组合中较低;同时还发现,Cicj和Cijp受叶温的影响强于光照。光合日进程中的FvCB模型模拟分析揭示出,早晨和傍晚弱光下为Aj限制时段;晴天上午和中午前后的充足日照下为Ac限制时段。多云和阴天下Aj的限制时段延长。增施CO2会延长Aj的限制时段,同时相应缩短Ac的限制时段;冬季2次增施CO2的出现了Ap限制时段。     

Drought‐introduced variability of mesophyll conductance in Gossypium and its relationship with leaf anatomy

Mesophyll conductance (g_m) is one of the major determinants of photosynthetic rate, for which it has an impact on crop yield. However, the regulatory mechanisms behind the decline in g_m of cotton (Gossypium. spp) by drought are unclear. An upland cotton (Gossypium hirsutum) genotype and a pima cotton (Gossypium barbadense) genotype were used to determine the gas exchange parameters, leaf anatomical structure as well as aquaporin and carbonic anhydrase gene expression under well‐watered and drought treatment conditions. In this study, the decrease of net photosynthetic rate (A_N) under drought conditions was related to a decline in g_m and in stomatal conductance (g_s). g_m and g_s coordinate with each other to ensure optimum state of CO₂ diffusion and achieve the balance of water and CO₂ demand in the process of photosynthesis. Meanwhile, mesophyll limitations to photosynthesis are equally important to the stomatal limitations. Considering g_m, its decline in cotton leaves under drought was mostly regulated by the chloroplast surface area exposed to leaf intercellular air spaces per leaf area (S_c/S) and might also be regulated by the expression of leaf CARBONIC ANHYDRASE (CA1). Meanwhile, cotton leaves can minimize the decrease in g_m under drought by maintaining cell wall thickness (T_cw). Our results indicated that modification of chloroplasts might be a target trait in future attempts to improve cotton drought tolerance.     

Do all leaf photosynthesis parameters of rice acclimate to elevated CO2, elevated temperature,and their combination,in FACE environments?

Leaf photosynthesis of crops acclimates to elevated CO₂ and temperature, but studies quantifying responses of leaf photosynthetic parameters to combined CO₂ and temperature increases under field conditions are scarce. We measured leaf photosynthesis of rice cultivars Changyou 5 and Nanjing 9108 grown in two free‐air CO₂ enrichment (FACE) systems, respectively, installed in paddy fields. Each FACE system had four combinations of two levels of CO₂ (ambient and enriched) and two levels of canopy temperature (no warming and warmed by 1.0–2.0°C). Parameters of the C₃ photosynthesis model of Farquhar, von Caemmerer and Berry (the FvCB model), and of a stomatal conductance (g_s) model were estimated for the four conditions. Most photosynthetic parameters acclimated to elevated CO₂, elevated temperature, and their combination. The combination of elevated CO₂ and temperature changed the functional relationships between biochemical parameters and leaf nitrogen content for Changyou 5. The g_s model significantly underestimated g_s under the combination of elevated CO₂ and temperature by 19% for Changyou 5 and by 10% for Nanjing 9108 if no acclimation was assumed. However, our further analysis applying the coupled g_s–FvCB model to an independent, previously published FACE experiment showed that including such an acclimation response of g_s hardly improved prediction of leaf photosynthesis under the four combinations of CO₂ and temperature. Therefore, the typical procedure that crop models using the FvCB and g_s models are parameterized from plants grown under current ambient conditions may not result in critical errors in projecting productivity of paddy rice under future global change.     

Rapid responses of mesophyll conductance to changes of CO2 concentration,temperature and irradiance are affected by N supplements in rice

Photosynthesis in C₃ plants is significantly limited by mesophyll conductance (g_m), which can vary with leaf anatomical traits and nitrogen (N) supplements. Several studies have investigated the response of g_m to N supplements; however, none examined the implications of N supplements on the response of g_m to rapid environmental changes. Here we investigated the effect of N supplement on g_m and the response of g_m to change of CO₂, temperature and irradiance in rice. High N supplement (HN) increased mesophyll cell wall surface area and chloroplast surface area exposed to intercellular airspace per leaf area, and reduced cell wall thickness. These changes resulted in increased g_m. The g_m of leaves with HN was more sensitive to changes in CO₂ concentration, temperature and irradiance. The difference in leaf structural features between low N supplement and HN indicates that a rapid change in g_m is related to the regulation of diffusion through biological membranes rather than leaf structural features. These results will contribute to an understanding of the determinants of g_m response to rapid changes in environmental factors.     

Photosynthesis Inhibition During Gas Exchange Oscillations in ABA-Treated Helianthus annuus: Relative Role of Stomatal Patchiness and Leaf Carboxylation Capacity

Environmental factors that induce spatial heterogeneity of stomatal conductance, g _s, called stomatal patchiness, also reduce the photochemical capacity of CO₂ fixation, yet current methods cannot distinguish between the relative effect of stomatal patchiness and biochemical limitations on photosynthetic capacity. We evaluate effects of stomatal patchiness and the biochemical capacity of CO₂ fixation on the sensitivity of net photosynthetic rate (P _N) to stomatal conductance (g _s), θ (θ = δP _N/g _s). A qualitative model shows that stomatal patchiness increases the sensitivity θ while reduced biochemical capacity of CO₂ fixation lowers θ. We used this feature to distinguish between stomatal patchiness and mesophyll impairments in the photochemistry of CO₂ fixation. We compared gas exchange of sunflower (Helianthus annuus L.) plants grown in a growth chamber and fed abscisic acid, ABA (10⁻⁵ M), for 10 d with control plants (-ABA). P _N and g _s oscillated more frequently in ABA-treated than in control plants when the leaves were placed into the leaf chamber and exposed to a dry atmosphere. When compared with the initial CO₂ response measured at the beginning of the treatment (day zero), both ABA and control leaves showed reduced P _N at particular sub-stomatal CO₂ concentration (c _i) during the oscillations. A lower reduction of P _N at particular g _s indicated overestimation of c _i due to stomatal patchiness and/or omitted cuticular conductance, g _c. The initial period of damp oscillation was characterised by inhibition of chloroplast processes while stomatal patchiness prevailed at the steady state of gas exchange. The sensitivity θ remained at the original pre-treatment values at high g _s in both ABA and control plants. At low g _s, θ decreased in ABA-treated plants indicating an ABA-induced impairment of chloroplast processes. In control plants, g _c neglected in the calculation of g _s was the likely reason for apparent depression of photosynthesis at low g _s. This revised version was published online in August 2006 with corrections to the Cover Date.     

Variable mesophyll conductance revisited: theoretical background and experimental implications

The CO₂ concentration at the site of carboxylation inside the chloroplast stroma depends not only on the stomatal conductance, but also on the conductance of CO₂ between substomatal cavities and the site of CO₂ fixation. This conductance, commonly termed mesophyll conductance (g_m), significantly constrains the rate of photosynthesis. Here we show that estimates of g_m are influenced by the amount of respiratory and photorespiratory CO₂ from the mitochondria diffusing towards the chloroplasts. This results in an apparent CO₂ and oxygen sensitivity of g_m that does not imply a change in intrinsic diffusion properties of the mesophyll, but depends on the ratio of mitochondrial CO₂ release to chloroplast CO₂ uptake. We show that this effect (1) can bias the estimation of the CO₂ photocompensation point and non‐photorespiratory respiration in the light; (2) can affect the estimates of ribulose 1·5‐bisphosphate carboxylase/oxygenase (Rubisco) kinetic constants in vivo; and (3) results in an apparent obligatory correlation between stomatal conductance and g_m. We further show that the amount of photo(respiratory) CO₂ that is refixed by Rubisco can be directly estimated through measurements of g_m.     

Effects of instantaneous and growth CO2 levels and abscisic acid on stomatal and mesophyll conductances

C₃ photosynthesis is often limited by CO₂ diffusivity or stomatal (g_s) and mesophyll (g_m) conductances. To characterize effects of stomatal closure induced by either high CO₂ or abscisic acid (ABA) application on g_m, we examined g_s and g_m in the wild type (Col‐0) and ost1 and slac1‐2 mutants of Arabidopsis thaliana grown at 390 or 780 μmol mol^?1 CO₂. Stomata of these mutants were reported to be insensitive to both high CO₂ and ABA. When the ambient CO₂ increased instantaneously, g_m decreased in all these plants, whereas g_s in ost1 and slac1‐2 was unchanged. Therefore, the decrease in g_m in response to high CO₂ occurred irrespective of the responses of g_s. g_m was mainly determined by the instantaneous CO₂ concentration during the measurement and not markedly by the CO₂ concentration during the growth. Exogenous application of ABA to Col‐0 caused the decrease in the intercellular CO₂ concentration (C_i). With the decrease in C_i, g_m did not increase but decreased, indicating that the response of g_m to CO₂ and that to ABA are differently regulated and that ABA content in the leaves plays an important role in the regulation of g_m.     

Using tunable diode laser spectroscopy to measure carbon isotope discrimination and mesophyll conductance to CO₂ diffusion dynamically at different CO₂ concentrations

In C₃ leaves, the mesophyll conductance to CO₂ diffusion, g_m, determines the drawdown in CO₂ concentration from intercellular airspace to the chloroplast stroma. Both g_m and stomatal conductance limit photosynthetic rate and vary in response to the environment. We investigated the response of g_m to changes in CO₂ in two Arabidopsis genotypes (including a mutant with open stomata, ost1), tobacco and wheat. We combined measurements of gas exchange with carbon isotope discrimination using tunable diode laser absorption spectroscopy with a CO₂ calibration system specially designed for a range of CO₂ and O₂ concentrations. CO₂ was initially increased from 200 to 1000 ppm and then decreased stepwise to 200 ppm and increased stepwise back to 1000 ppm, or the sequence was reversed. In 2% O₂ a step increase from 200 to 1000 ppm significantly decreased g_m by 26–40% in all three species, whereas following a step decrease from 1000 to 200 ppm, the 26–38% increase in g_m was not statistically significant. The response of g_m to CO₂ was less in 21% O₂. Comparing wild type against the ost1 revealed that mesophyll and stomatal conductance varied independently in response to CO₂. We discuss the effects of isotope fractionation factors on estimating g_m.     

Photosynthetic responses of soybean (Glycine max L.) to heat‐induced electrical signalling are predominantly governed by modifications of mesophyll conductance for CO2

In recent years, the effect of heat‐induced electrical signalling on plant photosynthetic activity has been demonstrated for many plant species. However, the underlying triggers of the resulting transient inhibition of photosynthesis still remain unknown. To further investigate on this phenomenon, we focused in our present study on soybean (Glycine max L.) on the direct effect of signal transmission in the leaf mesophyll on conductance for CO₂ diffusion in the mesophyll (g_m) and detected a drastic decline in g_m following the electrical signal, whereas the photosynthetic electron transport rate (ETR) was only marginally affected. In accordance with the drop in net photosynthesis (A_N), energy dispersive X‐ray analysis (EDXA) revealed a shift of K, Mg, O and P on leaf chloroplasts. Control experiments under elevated CO₂ conditions proved the transient reduction of A_N, ETR, the chloroplast CO₂ concentration (C_c) and g_m to be independent of the external CO₂ regime, whereas the effect of the electrical signal on stomatal conductance for CO₂ (g_s) turned out much less distinctive. We therefore conclude that the effect of electrical signalling on photosynthesis in soybean is triggered by its immediate effects on g_m.     

Abscisic Acid Induces Rapid Reductions in Mesophyll Conductance to Carbon Dioxide

The rate of photosynthesis (A) of plants exposed to water deficit is a function of stomatal (g_s) and mesophyll (g_m) conductance determining the availability of CO₂ at the site of carboxylation within the chloroplast. Mesophyll conductance often represents the greatest impediment to photosynthetic uptake of CO₂, and a crucial determinant of the photosynthetic effects of drought. Abscisic acid (ABA) plays a fundamental role in signalling and co-ordination of plant responses to drought; however, the effect of ABA on g_m is not well-defined. Rose, cherry, olive and poplar were exposed to exogenous ABA and their leaf gas exchange parameters recorded over a four hour period. Application with ABA induced reductions in values of A, g_s and g_m in all four species. Reduced g_m occurred within one hour of ABA treatment in three of the four analysed species; indicating that the effect of ABA on g_m occurs on a shorter timescale than previously considered. These declines in g_m values associated with ABA were not the result of physical changes in leaf properties due to altered turgor affecting movement of CO₂, or caused by a reduction in the sub-stomatal concentration of CO₂ (C_i). Increased [ABA] likely induces biochemical changes in the properties of the interface between the sub-stomatal air-space and mesophyll layer through the actions of cooporins to regulate the transport of CO₂. The results of this study provide further evidence that g_m is highly responsive to fluctuations in the external environment, and stress signals such as ABA induce co-ordinated modifications of both g_s and g_m in the regulation of photosynthesis.     

Leaf anatomy does not explain apparent short‐term responses of mesophyll conductance to light and CO2 in tobacco

Mesophyll conductance to CO₂ (g_m), a key photosynthetic trait, is strongly constrained by leaf anatomy. Leaf anatomical parameters such as cell wall thickness and chloroplast area exposed to the mesophyll intercellular airspace have been demonstrated to determine g_m in species with diverging phylogeny, leaf structure and ontogeny. However, the potential implication of leaf anatomy, especially chloroplast movement, on the short‐term response of g_m to rapid changes (i.e. seconds to minutes) under different environmental conditions (CO₂, light or temperature) has not been examined. The aim of this study was to determine whether the observed rapid variations of g_m in response to variations of light and CO₂ could be explained by changes in any leaf anatomical arrangements. When compared to high light and ambient CO₂, the values of g_m estimated by chlorophyll fluorescence decreased under high CO₂ and increased at low CO₂, while it decreased with decreasing light. Nevertheless, no changes in anatomical parameters, including chloroplast distribution, were found. Hence, the g_m estimated by analytical models based on anatomical parameters was constant under varying light and CO₂. Considering this discrepancy between anatomy and chlorophyll fluorescence estimates, it is concluded that apparent fast g_m variations should be due to artefacts in its estimation and/or to changes in the biochemical components acting on diffusional properties of the leaf (e.g. aquaporins and carbonic anhydrase).