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1.
Mesophyll conductance (gm) is known to affect plant photosynthesis. However, gm is rarely explicitly considered in land surface models (LSMs), with the consequence that its role in ecosystem and large‐scale carbon and water fluxes is poorly understood. In particular, the different magnitudes of gm across plant functional types (PFTs) are expected to cause spatially divergent vegetation responses to elevated CO2 concentrations. Here, an extensive literature compilation of gm across major vegetation types is used to parameterize an empirical model of gm in the LSM JSBACH and to adjust photosynthetic parameters based on simulated An ? Ci curves. We demonstrate that an explicit representation of gm changes the response of photosynthesis to environmental factors, which cannot be entirely compensated by adjusting photosynthetic parameters. These altered responses lead to changes in the photosynthetic sensitivity to atmospheric CO2 concentrations which depend both on the magnitude of gm and the climatic conditions, particularly temperature. We then conducted simulations under ambient and elevated (ambient + 200 μmol/mol) CO2 concentrations for contrasting ecosystems and for historical and anticipated future climate conditions (representative concentration pathways; RCPs) globally. The gm‐explicit simulations using the RCP8.5 scenario resulted in significantly higher increases in gross primary productivity (GPP) in high latitudes (+10% to + 25%), intermediate increases in temperate regions (+5% to + 15%), and slightly lower to moderately higher responses in tropical regions (?2% to +5%), which summed up to moderate GPP increases globally. Similar patterns were found for transpiration, but with a lower magnitude. Our results suggest that the effect of an explicit representation of gm is most important for simulated carbon and water fluxes in the boreal zone, where a cold climate coincides with evergreen vegetation.  相似文献   

2.
Mesophyll conductance (gm) generally correlates with photosynthetic capacity, although the causal relationship between the two is unclear. The response of gm to various CO2 regimes was measured to determine its relationship to environmental changes that affect photosynthesis. The overall effect of CO2 growth environment on gm was species and experiment dependent. The data did not statistically differ from the previously shown Agm relationship and was unaffected by CO2 treatment. The consequences of the CO2 effect on gm for interpreting photosynthesis in individual cases were investigated. Substantial effects of assumed versus calculated gm on leaf properties estimated from gas‐exchange measurements were found. This differential error resulted in an underestimation in ratio of maximum carboxylation to electron transport, especially in plants with high photosynthetic capacity. Including gm in the calculations also improved the agreement between maximum carboxylation rates and in vitro Rubisco measurements. It is concluded that gm is finite and varies with photosynthetic capacity. Including gm when calculating photosynthesis parameters from gas‐exchange data will avoid systematic errors.  相似文献   

3.
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 (gm) and detected a drastic decline in gm following the electrical signal, whereas the photosynthetic electron transport rate (ETR) was only marginally affected. In accordance with the drop in net photosynthesis (AN), 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 AN, ETR, the chloroplast CO2 concentration (Cc) and gm to be independent of the external CO2 regime, whereas the effect of the electrical signal on stomatal conductance for CO2 (gs) 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 gm.  相似文献   

4.
基于FvCB模型的叶片光合生理对环境因子的响应研究进展   总被引:7,自引:0,他引:7  
唐星林  曹永慧  顾连宏  周本智 《生态学报》2017,37(19):6633-6645
为提高叶片光合速率并更好地理解叶片光合生理对环境因子变化的响应机制,FvCB模型(C_3植物光合生化模型)常用于分析不同环境条件下CO_2响应曲线并预测叶片活体内光合系统的内在变化状况。系统介绍了FvCB模型的建立、发展过程和拟合方法等基本理论,综述了该模型在叶片光合生理对光、CO_2、水、温度和N营养等环境因子变化的响应机制中的应用研究。为进一步完善FvCB模型并更好地理解叶片活体内光合系统对环境因子变化的响应机制,未来拟加强以下研究:1)羧化速率与光合电子传递速率之间的联系;2)叶肉导度的具体组分及其对FvCB模型参数估计的影响;3)叶片气孔导度和叶肉导度对环境因子变化的调控机制。  相似文献   

5.
Light gradients within tree canopies play a major role in the distribution of plant resources that define the photosynthetic capacity of sun and shade leaves. However, the biochemical and diffusional constraints on gas exchange in sun and shade leaves in response to light remain poorly quantified, but critical for predicting canopy carbon and water exchange. To investigate the CO2 diffusion pathway of sun and shade leaves, leaf gas exchange was coupled with concurrent measurements of carbon isotope discrimination to measure net leaf photosynthesis (An), stomatal conductance (gs) and mesophyll conductance (gm) in Eucalyptus tereticornis trees grown in climate controlled whole‐tree chambers. Compared to sun leaves, shade leaves had lower An, gm, leaf nitrogen and photosynthetic capacity (Amax) but gs was similar. When light intensity was temporarily increased for shade leaves to match that of sun leaves, both gs and gm increased, and An increased to values greater than sun leaves. We show that dynamic physiological responses of shade leaves to altered light environments have implications for up‐scaling leaf level measurements and predicting whole canopy carbon gain. Despite exhibiting reduced photosynthetic capacity, the rapid up‐regulation of gm with increased light enables shade leaves to respond quickly to sunflecks.  相似文献   

6.
In C3 leaves, the mesophyll conductance to CO2 diffusion, gm, determines the drawdown in CO2 concentration from intercellular airspace to the chloroplast stroma. Both gm and stomatal conductance limit photosynthetic rate and vary in response to the environment. We investigated the response of gm 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 gm by 26–40% in all three species, whereas following a step decrease from 1000 to 200 ppm, the 26–38% increase in gm was not statistically significant. The response of gm 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 gm.  相似文献   

7.
The rate of photosynthesis (A) of plants exposed to water deficit is a function of stomatal (gs) and mesophyll (gm) 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 gm 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, gs and gm in all four species. Reduced gm occurred within one hour of ABA treatment in three of the four analysed species; indicating that the effect of ABA on gm occurs on a shorter timescale than previously considered. These declines in gm 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 (Ci). 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 gm is highly responsive to fluctuations in the external environment, and stress signals such as ABA induce co-ordinated modifications of both gs and gm in the regulation of photosynthesis.  相似文献   

8.
The relationship between single leaf photosynthesis and conductance was examined in cotton (Gossypium hirsutum L.) across a range of environmental conditions. The purpose of this research was to separate and define the degree of stomatal and nonstomatal limitations in the photosynthetic process of field-grown cotton.

Photosynthetic rates were related to leaf conductance of upper canopy leaves in a curvilinear manner. Increases in leaf conductance of CO2 in excess of 0.3 to 0.4 mole per square meter per second did not result in significant increases in gross or net photosynthetic rates. No tight coupling between environmental influences on photosynthetic rates and those affecting conductance levels was evident, since photosynthesis per unit leaf conductance did not remain constant. Slowly developing water stress caused greater reductions in photosynthesis than in leaf conductance, indicating nonstomatal limitations of photosynthesis.

Increases in external CO2 concentration to levels above ambient did not produce proportional increases in photosynthesis even though substomatal or intercellular CO2 concentration increased. The lack of a linear increase in photosynthetic rate in response to increases in leaf conductance and in response to increases in external CO2 concentration demonstrated that nonstomatal factors are major photosynthetic rate determinants of cotton under field conditions.

  相似文献   

9.
Mesophyll conductance (gm) 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 gm 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 (AN) under drought conditions was related to a decline in gm and in stomatal conductance (gs). gm and gs 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 gm, its decline in cotton leaves under drought was mostly regulated by the chloroplast surface area exposed to leaf intercellular air spaces per leaf area (Sc/S) and might also be regulated by the expression of leaf CARBONIC ANHYDRASE (CA1). Meanwhile, cotton leaves can minimize the decrease in gm under drought by maintaining cell wall thickness (Tcw). Our results indicated that modification of chloroplasts might be a target trait in future attempts to improve cotton drought tolerance.  相似文献   

10.
Photosynthesis in C3 plants is significantly limited by mesophyll conductance (gm), which can vary with leaf anatomical traits and nitrogen (N) supplements. Several studies have investigated the response of gm to N supplements; however, none examined the implications of N supplements on the response of gm to rapid environmental changes. Here we investigated the effect of N supplement on gm and the response of gm 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 gm. The gm 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 gm 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 gm response to rapid changes in environmental factors.  相似文献   

11.
Rising atmospheric carbon dioxide concentration ([CO2]) significantly influences plant growth, development, and biomass. Increased photosynthesis rate, together with lower stomatal conductance, has been identified as the key factors that stimulate plant growth at elevated [CO2] (e[CO2]). However, variations in photosynthesis and stomatal conductance alone cannot fully explain the dynamic changes in plant growth. Stimulation of photosynthesis at e[CO2] is always associated with post‐photosynthetic secondary metabolic processes that include carbon and nitrogen metabolism, cell cycle functions, and hormonal regulation. Most studies have focused on photosynthesis and stomatal conductance in response to e[CO2], despite the emerging evidence of e[CO2]'s role in moderating secondary metabolism in plants. In this review, we briefly discuss the effects of e[CO2] on photosynthesis and stomatal conductance and then focus on the changes in other cellular mechanisms and growth processes at e[CO2] in relation to plant growth and development. Finally, knowledge gaps in understanding plant growth responses to e[CO2] have been identified with the aim of improving crop productivity under a CO2 rich atmosphere.  相似文献   

12.
Decline in mesophyll conductance (gm) plays a key role in limiting photosynthesis in plants exposed to elevated ozone (O3). Leaf anatomical traits are known to influence gm, but the potential effects of O3-induced changes in leaf anatomy on gm have not yet been clarified. Here, two poplar clones were exposed to elevated O3. The effects of O3 on the photosynthetic capacity and anatomical characteristics were assessed to investigate the leaf anatomical properties that potentially affect gm. We also conducted global meta-analysis to explore the general response patterns of gm and leaf anatomy to O3 exposure. We found that the O3-induced reduction in gm was critical in limiting leaf photosynthesis. Changes in liquid-phase conductance rather than gas-phase conductance drive the decline in gm under elevated O3, and this effect was associated with thicker cell walls and smaller chloroplast sizes. The effects of O3 on palisade and spongy mesophyll cell traits and their contributions to gm were highly genotype-dependent. Our results suggest that, while anatomical adjustments under elevated O3 may contribute to defense against O3 stress, they also cause declines in gm and photosynthesis. These results provide the first evidence of anatomical constraints on gm under elevated O3.  相似文献   

13.
Improving Rubisco catalysis is considered a promising way to enhance C3-photosynthesis and photosynthetic water use efficiency (WUE) provided the introduced changes have little or no impact on other processes affecting photosynthesis such as leaf photochemistry or leaf CO2 diffusion conductances. However, the extent to which the factors affecting photosynthetic capacity are co-regulated is unclear. The aim of the present study was to characterize the photochemistry and CO2 transport processes in the leaves of three transplantomic tobacco genotypes expressing hybrid Rubisco isoforms comprising different Flaveria L-subunits that show variations in catalysis and differing trade-offs between the amount of Rubisco and its activation state. Stomatal conductance (g s) in each transplantomic tobacco line matched wild-type, while their photochemistry showed co-regulation with the variations in Rubisco catalysis. A tight co-regulation was observed between Rubisco activity and mesophyll conductance (g m) that was independent of g s thus producing plants with varying g m/g s ratios. Since the g m/g s ratio has been shown to positively correlate with intrinsic WUE, the present results suggest that altering photosynthesis by modifying Rubisco catalysis may also be useful for targeting WUE.  相似文献   

14.
Natural selection on photosynthetic performance is a primary factor determining leaf phenotypes. The complex CO2 diffusion path from substomatal cavities to the chloroplasts – the mesophyll conductance (gm) – limits photosynthetic rate in many species and hence shapes variation in leaf morphology and anatomy. Among sclerophyllous and succulent taxa, structural investment in leaves, measured as the leaf dry mass per area (LMA), has been implicated in decreased gm. However, in herbaceous taxa with high gm, it is less certain how LMA impacts CO2 diffusion and whether it significantly affects photosynthetic performance. We addressed these questions in the context of understanding the ecophysiological significance of leaf trait variation in wild tomatoes, a closely related group of herbaceous perennials. Although gm was high in wild tomatoes, variation in gm significantly affected photosynthesis. Even in these tender‐leaved herbaceous species, greater LMA led to reduced gm. This relationship between gm and LMA is partially mediated by cell packing and leaf thickness, although amphistomy (equal distribution of stomata on both sides of the leaf) mitigates the effect of leaf thickness. Understanding the costs of increased LMA will inform future work on the adaptive significance of leaf trait variation across ecological gradients in wild tomatoes and other systems.  相似文献   

15.
Biochemical models are used to predict and understand the response of photosynthesis to rising temperatures and CO2 partial pressures. These models require the temperature dependency of ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Rubisco) kinetics and mesophyll conductance to CO2 (gm). However, it is not known how the temperature response of Rubisco kinetics differs between species, and comprehensive in vivo Rubisco kinetics that include gm have only been determined in the warm‐adapted Nicotiana tabacum. Here, we measured the temperature response of Rubisco kinetics and gm in N. tabacum and the cold‐adapted Arabidopsis thaliana using gas exchange and 13CO2 isotopic discrimination on plants with genetically reduced levels of Rubisco. While the individual Rubisco kinetic parameters in N. tabacum and A. thaliana were similar across temperatures, they collectively resulted in significantly different modelled rates of photosynthesis. Additionally, gm increased with temperature in N. tabacum but not in A. thaliana. These findings highlight the importance of considering species‐dependent differences in Rubisco kinetics and gm when modelling the temperature response of photosynthesis.  相似文献   

16.
Yu  Gui-Rui  Kobayashi  Tatsuaki  Zhuang  Jie  Wang  Qiu-Feng  Qu  Le-Qing 《Plant and Soil》2003,249(2):401-415
The study presents a theoretical basis of a stomatal behavior-based coupled model for estimating photosynthesis, A, and transpiration, E. Outputs of the model were tested against data observed in a maize (Zea mays L.) field. The model was developed by introducing the internal conductance, g ic, to CO2 assimilation, and the general equation of stomatal conductance, g sw, to H2O diffusion, into models of CO2 and H2O diffusion through the stomata of plant leaves. The coupled model is easier for practical use since the model only includes environmental variables, such as ambient CO2 concentration, leaf temperature, humidity and photosynthetic photon flux received at the leaves within the canopy. Moreover, concept of g ic, and factors controlling A and E were discussed, and applicability of the model was examined with the data collected in the maize field.  相似文献   

17.
Mesophyll conductance to CO2 (gm), 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 gm 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 gm 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 gm 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 gm 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 gm 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 gm 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).  相似文献   

18.
Very few studies have attempted to disentangle the respective role of ontogeny and water stress on leaf photosynthetic attributes. The relative significance of both effects on photosynthetic attributes has been investigated in leaves of field‐grown almond trees [Prunus dulcis (Mill.) D. A. Webb] during four growth cycles. Leaf ontogeny resulted in enhanced leaf dry weight per unit area (Wa), greater leaf dry‐to‐fresh weight ratio and lower N content per unit of leaf dry weight (Nw). Concomitantly, area‐based maximum carboxylation rate (Vcmax), maximum electron transport rate (Jmax), mesophyll conductance to CO2 diffusion (gm)′ and light‐saturated net photosynthesis (Amax) declined in both well‐watered and water‐stressed almond leaves. Although gm and stomatal conductance (gs) seemed to be co‐ordinated, a much stronger coordination in response to ontogeny and prolonged water stress was observed between gm and the leaf photosynthetic capacity. Under unrestricted water supply, the leaf age‐related decline of Amax was equally driven by diffusional and biochemical limitations. Under restricted soil water availability, Amax was mainly limited by gs and, to a lesser extent, by photosynthetic capacity and gm. When both ontogeny and water stress effects were combined, diffusional limitations was the main determinant of photosynthesis limitation, while stomatal and biochemical limitations contributed similarly.  相似文献   

19.
Mesophyll conductance (gm), the diffusion of CO2 from substomatal cavities to the carboxylation sites in the chloroplasts, is a highly complex trait driving photosynthesis (net CO2 assimilation, AN). However, little is known concerning the mechanisms by which it is dynamically regulated. The apoplast is considered as a ‘key information bridge’ between the environment and cells. Interestingly, most of the environmental constraints affecting gm also cause apoplastic responses, cell wall (CW) alterations and metabolic rearrangements. Since CW thickness is a key determinant of gm, we hypothesize that other changes in this cellular compartiment should also influence gm. We study the relationship between the antioxidant apoplastic system and CW metabolism and the gm responses in tobacco plants (Nicotiana sylvestris L.) under two abiotic stresses (drought and salinity), combining in vivo gas‐exchange measurements with analyses of antioxidant activities, CW composition and primary metabolism. Stress treatments imposed substantial reductions in AN (58–54%) and gm (59%), accompanied by a strong antioxidant enzymatic response at the apoplastic and symplastic levels. Interestingly, apoplastic but not symplastic peroxidases were positively related to gm. Leaf anatomy remained mostly stable; however, the stress treatments significantly affected the CW composition, specifically pectins, which showed significant relationships with AN and gm. The treatments additionally promoted a differential primary metabolic response, and specific CW‐related metabolites including galactose, glucosamine and hydroxycinnamate showed exclusive relationships with gm independent of the stress. These results suggest that gm responses can be attributed to specific changes in the apoplastic antioxidant system and CW metabolism, opening up more possibilities for improving photosynthesis using breeding/biotechnological strategies.  相似文献   

20.
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 (gm), significantly constrains the rate of photosynthesis. Here we show that estimates of gm 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 gm 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 gm. We further show that the amount of photo(respiratory) CO2 that is refixed by Rubisco can be directly estimated through measurements of gm.  相似文献   

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