Photosynthetic Diffusional Constraints Affect Yield in Drought Stressed Rice Cultivars during Flowering

Global production of rice (Oryza sativa) grain is limited by water availability and the low ‘leaf-level’ photosynthetic capacity of many cultivars. Oryza sativa is extremely susceptible to water-deficits; therefore, predicted increases in the frequency and duration of drought events, combined with future rises in global temperatures and food demand, necessitate the development of more productive and drought tolerant cultivars. We investigated the underlying physiological, isotopic and morphological responses to water-deficit in seven common varieties of O. sativa, subjected to prolonged drought of varying intensities, for phenotyping purposes in open field conditions. Significant variation was observed in leaf-level photosynthesis rates (A) under both water treatments. Yield and A were influenced by the conductance of the mesophyll layer to CO₂ (g _m) and not by stomatal conductance (g _s). Mesophyll conductance declined during drought to differing extents among the cultivars; those varieties that maintained g _m during water-deficit sustained A and yield to a greater extent. However, the variety with the highest g _m and yield under well-watered conditions (IR55419-04) was distinct from the most effective cultivar under drought (Vandana). Mesophyll conductance most effectively characterises the photosynthetic capacity and yield of O. sativa cultivars under both well-watered and water-deficit conditions; however, the desired attributes of high g _m during optimal growth conditions and the capacity for g _m to remain constant during water-deficit may be mutually exclusive. Nonetheless, future genetic and physiological studies aimed at enhancing O. sativa yield and drought stress tolerance should investigate the biochemistry and morphology of the interface between the sub-stomatal pore and mesophyll layer.     

Light-harvesting in photosystem I

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.     

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.     

Drought response of mesophyll conductance in forest understory species – impacts on water‐use efficiency and interactions with leaf water movement

Regulation of stomatal (g_s) and mesophyll conductance (g_m) is an efficient means for optimizing the relationship between water loss and carbon uptake in plants. We assessed water‐use efficiency (WUE)‐based drought adaptation strategies with respect to mesophyll conductance of different functional plant groups of the forest understory. Moreover we aimed at assessing the mechanisms of and interactions between water and CO₂ conductance in the mesophyll. The facts that an increase in WUE was observed only in the two species that increased g_m in response to moderate drought, and that over all five species examined, changes in mesophyll conductance were significantly correlated with the drought‐induced change in WUE, proves the importance of g_m in optimizing resource use under water restriction. There was no clear correlation of mesophyll CO₂ conductance and the tortuosity of water movement in the leaf across the five species in the control and drought treatments. This points either to different main pathways for CO₂ and water in the mesophyll either to different regulation of a common pathway.     

Mesophyll conductance decreases in the wild type but not in an ABA‐deficient mutant (aba1) of Nicotiana plumbaginifolia under drought conditions

Under drought conditions, leaf photosynthesis is limited by the supply of CO₂. Drought induces production of abscisic acid (ABA), and ABA decreases stomatal conductance (g_s). Previous papers reported that the drought stress also causes the decrease in mesophyll conductance (g_m). However, the relationships between ABA content and g_m are unclear. We investigated the responses of g_m to the leaf ABA content [(ABA)_L] using an ABA‐deficient mutant, aba1, and the wild type (WT) of Nicotiana plumbaginifolia. We also measured leaf water potential (Ψ_L) because leaf hydraulics may be related to g_m. Under drought conditions, g_m decreased with the increase in (ABA)_L in WT, whereas both (ABA)_L and g_m were unchanged by the drought treatment in aba1. Exogenously applied ABA decreased g_m in both WT and aba1 in a dose‐dependent manner. Ψ_L in WT was decreased by the drought treatment to ?0.7 MPa, whereas Ψ_L in aba1 was around ?0.8 MPa even under the well‐watered conditions and unchanged by the drought treatment. From these results, we conclude that the increase in (ABA)_L is crucial for the decrease in g_m under drought conditions. We discuss possible relationships between the decrease in g_m and changes in the leaf hydraulics.     

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.     

Methods of mesophyll conductance estimation: its impact on key biochemical parameters and photosynthetic limitations in phosphorus‐stressed soybean across CO2

Despite the development of various methods, the rapid estimation of mesophyll conductance (g_m) for a large number of samples is still a daunting challenge. Although the accurate estimation of g_m is critical to partition photosynthetic limitations by stomatal (L_s) and mesophyll (L_m) conductance and by photo‐biochemical (L_b) processes, the impact of various g_m estimation methods on this is ambiguous. As phosphorus (P) starvation and elevated CO₂ (eCO₂) strongly affect photosynthetic processes, their combined effect on the proportional changes in these limitations are not well understood. To investigate this, while also evaluating distinct recent methods of g_m estimation sharing few common theories and assumptions, soybean was grown under a range of P nutrition at ambient and eCO₂. Methods significantly affected g_m and carboxylation efficiency (V_Cmax) but not other photosynthetic parameters. In all the methods, all photosynthetic parameters responded similarly to treatments. However, the percentage difference between V_Cmax assuming finite and infinite g_m was highly inconsistent among methods. The primary mechanism responsible for P limitation to soybean photosynthesis was not CO₂ diffusion limitations but L_b comprised of reduced chlorophyll, photochemistry and biochemical processes. The eCO₂ decreased L_b but increased L_m without affecting L_s across leaf P concentration. Although each method explored advances of our understanding about g_m variability, they all require assumptions of varying degrees, which lead to the discrepancy in the g_m values. Among the methods, the oxygen sensitivity‐based g_m estimation appeared to be suitable for the quick assessment of a large number of samples or genotypes. Digital tools are provided for the easy estimation of g_m for some methods.     

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.     

Nonstomatal Inhibition of Net CO(2) Uptake by (+/-) Abscisic Acid in Pharbitis nil

(±) Abscisic acid (ABA) injected into petioles of attached transpiring leaves of Pharbitis nil Chois. cv violet reduced the photosynthetic capacity of the mesophyll of these leaves as well as the stomatal conductance to CO₂ diffusion. Greater than 75% of the injected ABA was recovered as ABA, suggesting that ABA rather than some metabolite thereof was the active compound. The nonstomatal effect of ABA increased from 30% reduction in photosynthesis at 0.25 micromolar ABA in the leaf blade to 90% reduction at 18 micromolar. Despite the effect of ABA on the nonstomatal component of leaf net CO₂ uptake, it was calculated that a substantial part of the reduction in leaf net CO₂ uptake (50-80%) could be accounted for by the effect of ABA on stomatal conductance.     

On the Use of Leaf Spectral Indices to Assess Water Status and Photosynthetic Limitations in Olea europaea L. during Water-Stress and Recovery

Diffusional limitations to photosynthesis, relative water content (RWC), pigment concentrations and their association with reflectance indices were studied in olive (Olea europaea) saplings subjected to water-stress and re-watering. RWC decreased sharply as drought progressed. Following rewatering, RWC gradually increased to pre-stress values. Photosynthesis (A), stomatal conductance (g_s), mesophyll conductance (g_m), total conductance (g_t), photochemical reflectance index (PRI), water index (WI) and relative depth index (RDI) closely followed RWC. In contrast, carotenoid concentration, the carotenoid to chlorophyll ratio, water content reflectance index (WCRI) and structural independent pigment index (SIPI) showed an opposite trend to that of RWC. Photosynthesis scaled linearly with leaf conductance to CO₂; however, A measured under non-photorespiratory conditions (A_1%O2) was approximately two times greater than A measured at 21% [O₂], indicating that photorespiration likely increased in response to drought. A_1%O2 also significantly correlated with leaf conductance parameters. These relationships were apparent in saturation type curves, indicating that under non-photorespiratory conditions, CO₂ conductance was not the major limitations to A. PRI was significant correlated with RWC. PRI was also very sensitive to pigment concentrations and photosynthesis, and significantly tracked all CO₂ conductance parameters. WI, RDI and WCRI were all significantly correlated with RWC, and most notably to leaf transpiration. Overall, PRI correlated more closely with carotenoid concentration than SIPI; whereas WI tracked leaf transpiration more effectively than RDI and WCRI. This study clearly demonstrates that PRI and WI can be used for the fast detection of physiological traits of olive trees subjected to water-stress.     

Decrease of mesophyll conductance to CO<Subscript>2</Subscript> is a possible mechanism of abscisic acid influence on photosynthesis in seedlings of pea and wheat

Treatment of plants by phytohormones is a perspective method of regulation of the plant stress resistance and productivity. However, the mechanisms of phytohormone effects on physiological processes require investigations. The aim of this work was the analysis of the influence of exogenous abscisic acid (ABA) on photosynthesis in seedlings of pea and wheat, and in particular, an estimation of the involvement of the mesophyll conductance to CO₂ in the realization of the ABA effects. A standard system for recording of photosynthetic parameters and a system for intracellular measurements of electrical activity were used in the experiments. It was shown that the effect of exogenous ABA on photosynthesis was most prominent 1 day after spraying of a plant. A detailed analysis of photosynthetic processes showed that ABA decreased photosynthetic assimilation of CO₂ and increased cyclic electron flow in plants under study; these processes were interconnected. A decrease of the mesophyll conductance to CO₂ was probably a mechanism of the decrease in the photosynthetic assimilation of CO₂, because ABA did not significantly influence water conductance of a leaf and parameters of the CO₂ fixation in Calvin–Benson cycle. It is likely that the decrease of the mesophyll conductance to CO₂ was related with a decrease of the plasma membrane conductance to carbon dioxide. Inactivation of H⁺-ATP-ase and changes in extracellular pH can be a mechanism of this decrease. A decrease of the metabolic component of the electrical resting potential after the ABA treatment testifies in favor of this possibility.     

Significance of mesophyll conductance for photosynthetic capacity and water-use efficiency in response to alkaline stress in Populus cathayana seedlings

Cuttings of Populus cathayana were exposed to three different alkaline regimes (0, 75, and 150 mM Na₂CO₃) in a semicontrolled environment. The net photosynthesis rate (P _N), mesophyll conductance (g _m), the relative limitations posed by stomatal conductance (L _s) and by mesophyll conductance (L _m), photosynthetic nitrogen-use efficiency (PNUE), carbon isotope composition (δ¹³C), as well as specific leaf area (SLA) were measured. P _N decreased due to alkaline stress by an average of 25% and g _m decreased by an average of 57%. Alkaline stress caused an increase of L _m but not L _s, with average L _s of 26%, and L _m average of 38% under stress conditions. Our results suggested reduced assimilation rate under alkaline stress through decreased mesophyll conductance in P. cathayana. Moreover, alkaline stress increased significantly δ¹³C and it drew down CO₂ concentration from the substomatal cavities to the sites of carboxylation (C _i-C _c), but decreased PNUE. Furthermore, a relationship was found between PNUE and C _i-C _c. Meanwhile, no correlation was found between δ¹³C and C _i/C _a, but a strong correlation was proved between δ¹³C and C _c/C _a, indicating that mesophyll conductance was also influencing the ¹³C/¹²C ratio of leaf under alkaline stress.     

Stomatal,mesophyll conductance,and biochemical limitations to photosynthesis during induction

The dynamics of leaf photosynthesis in fluctuating light affects carbon gain by plants. Mesophyll conductance (g_m) limits CO₂ assimilation rate (A) under the steady state, but the extent of this limitation under non-steady-state conditions is unknown. In the present study, we aimed to characterize the dynamics of g_m and the limitations to A imposed by gas diffusional and biochemical processes under fluctuating light. The induction responses of A, stomatal conductance (g_s), g_m, and the maximum rate of RuBP carboxylation (V_cmax) or electron transport (J) were investigated in Arabidopsis (Arabidopsis thaliana (L.)) and tobacco (Nicotiana tabacum L.). We first characterized g_m induction after a change from darkness to light. Each limitation to A imposed by g_m, g_s and V_cmax or J was significant during induction, indicating that gas diffusional and biochemical processes limit photosynthesis. Initially, g_s imposed the greatest limitation to A, showing the slowest response under high light after long and short periods of darkness, assuming RuBP-carboxylation limitation. However, if RuBP-regeneration limitation was assumed, then J imposed the greatest limitation. g_m did not vary much following short interruptions to light. The limitation to A imposed by g_m was the smallest of all the limitations for most of the induction phase. This suggests that altering induction kinetics of mesophyll conductance would have little impact on A following a change in light. To enhance the carbon gain by plants under naturally dynamic light environments, attention should therefore be focused on faster stomatal opening or activation of electron transport.

Gas diffusional and biochemical processes impose significant limitations to CO2 assimilation during photosynthetic induction.     

Diffusional conductances to CO2 as a target for increasing photosynthesis and photosynthetic water-use efficiency

A key objective for sustainable agriculture and forestry is to breed plants with both high carbon gain and water-use efficiency (WUE). At the level of leaf physiology, this implies increasing net photosynthesis (A _N) relative to stomatal conductance (g _s). Here, we review evidence for CO₂ diffusional constraints on photosynthesis and WUE. Analyzing past observations for an extensive pool of crop and wild plant species that vary widely in mesophyll conductance to CO₂ (g _m), g _s, and foliage A _N, it was shown that both g _s and g _m limit A _N, although the relative importance of each of the two conductances depends on species and conditions. Based on Fick’s law of diffusion, intrinsic WUE (the ratio A _N/g _s) should correlate on the ratio g _m/g _s, and not g _m itself. Such a correlation is indeed often observed in the data. However, since besides diffusion A _N also depends on photosynthetic capacity (i.e., V _c,max), this relationship is not always sustained. It was shown that only in a very few cases, genotype selection has resulted in simultaneous increases of both A _N and WUE. In fact, such a response has never been observed in genetically modified plants specifically engineered for either reduced g _s or enhanced g _m. Although increasing g _m alone would result in increasing photosynthesis, and potentially increasing WUE, in practice, higher WUE seems to be only achieved when there are no parallel changes in g _s. We conclude that for simultaneous improvement of A _N and WUE, genetic manipulation of g _m should avoid parallel changes in g _s, and we suggest that the appropriate trait for selection for enhanced WUE is increased g _m/g _s.     

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.     

Evolution of protein transport to the chloroplast envelope membranes

Comparing with other angiosperms, most members within the family Orchidaceae have lower photosynthetic capacities. However, the underlying mechanisms remain unclear. Cypripedium and Paphiopedilum are closely related phylogenetically in Orchidaceae, but their photosynthetic performances are different. We explored the roles of internal anatomy and diffusional conductance in determining photosynthesis in three Cypripedium and three Paphiopedilum species, and quantitatively analyzed their diffusional and biochemical limitations to photosynthesis. Paphiopedilum species showed lower light-saturated photosynthetic rate (A _N), stomatal conductance (g _s), and mesophyll conductance (g _m) than Cypripedium species. A _N was positively correlated with g _s and g _m. And yet, in both species A _N was more strongly limited by g _m than by biochemical factors or g _s. The greater g _s of Cypripedium was mainly affected by larger stomatal apparatus area and smaller pore depth, while the less g _m of Paphiopedilum was determined by the reduced surface area of mesophyll cells and chloroplasts exposed to intercellular airspace per unit of leaf area, and much thicker cell wall thickness. These results suggest that leaf anatomical structure is the key factor affecting g _m, which is largely responsible for the difference in photosynthetic capacity between those two genera. Our findings provide new insight into the photosynthetic physiology and functional diversification of orchids.     

Diffusion limitations and metabolic factors associated with inhibition and recovery of photosynthesis from drought stress in a C3 perennial grass species

Stomatal closure and metabolic impairment under drought stress limits photosynthesis. The objective of this study was to determine major stomatal and metabolic factors involved in photosynthetic responses to drought and recovery upon re‐watering in a C₃ perennial grass species, Kentucky bluegrass (Poa pratensis L.). Two genotypes differing in drought resistance, ‘Midnight’ (tolerant) and ‘Brilliant’ (sensitive), were subjected to drought stress for 15 days and then re‐watered for 10 days in growth chambers. Single‐leaf net photosynthetic rate (A), stomatal conductance (g_s) and transpiration rate (Tr) decreased during drought, with a less rapid decline in ‘Midnight’ than in ‘Brilliant’. Photochemical efficiency, Rubisco activity and activation state declined during drought, but were significantly higher in ‘Midnight’ than in ‘Brilliant’. The relationship between A and internal leaf CO₂ concentration (A/Ci curve) during drought and re‐watering was analyzed to estimate the relative influence of stomatal and non‐stomatal components on photosynthesis. Stomatal limitation (Ls %), non‐stomatal limitation (Lns %), CO₂ compensation point (CP) and dark respiration (Rd) increased with stress duration in both genotypes, but to a lesser extent in ‘Midnight’. Maximum CO₂ assimilation rate (A_max), carboxylation efficiency (CE) and mesophyll conductance (g_m) declined, but ‘Midnight’ had significantly higher levels of A_max, CE and g_m than ‘Brilliant’. Maximum carboxylation rate of Rubisco (V_cmax) and ribulose‐1,5‐bisphospate (RuBP) regeneration capacity mediated by maximum electron transport rate (J_max) decreased from moderate to severe drought stress in both genotypes, but to a greater extent in ‘Brilliant’ than in ‘Midnight’. After re‐watering, RWC restored to about 90% of the control levels in both genotypes, whereas A, g_s, Tr and Fv/Fm was only partially recovered, with a higher recovery level in ‘Midnight’ than in ‘Brilliant’. Rubisco activity and activation state restored to the control level after re‐watering, with more rapid increase in ‘Midnight’ than in ‘Brilliant’. The values of Ls, Lns, CP and Rd declined, and A_max, CE, V_cmax, J_max and g_m increased after re‐watering, with more rapid change in all parameters in ‘Midnight’ than in ‘Brilliant’. These results indicated that the maintenance of higher A and A_max under drought stress in drought‐tolerant Kentucky bluegrass could be attributed to higher Rubico activation state, higher CE and less stomatal limitation. The ability to resume metabolic activity (A_max, CE, Fv/Fm and Rubisco) was observed in the drought‐tolerant genotype and is the most likely cause for the increased recuperative ability of photosynthesis. Incomplete recovery of photosynthesis upon re‐watering could be attributable to lasting stomatal limitations caused by severe drought damage in both genotypes. Promoting rapid stomatal recovery from drought stress may be critical for plants to resume full photosynthetic capacity in C₃ perennial grass species.     

基于FvCB模型的叶片光合生理对环境因子的响应研究进展

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。     

Differential tissue-specific expression of NtAQP1 in Arabidopsis thaliana reveals a role for this protein in stomatal and mesophyll conductance of CO2 under standard and salt-stress conditions

The regulation of plant hydraulic conductance and gas conductance involves a number of different morphological, physiological and molecular mechanisms working in harmony. At the molecular level, aquaporins play a key role in the transport of water, as well as CO₂, through cell membranes. Yet, their tissue-related function, which controls whole-plant gas exchange and water relations, is less understood. In this study, we examined the tissue-specific effects of the stress-induced tobacco Aquaporin1 (NtAQP1), which functions as both a water and CO₂ channel, on whole-plant behavior. In tobacco and tomato plants, constitutive overexpression of NtAQP1 increased net photosynthesis (A _N), mesophyll CO₂ conductance (g _m) and stomatal conductance (g _s) and, under stress, increased root hydraulic conductivity (L _pr) as well. Our results revealed that NtAQP1 that is specifically expressed in the mesophyll tissue plays an important role in increasing both A _N and g _m. Moreover, targeting NtAQP1 expression to the cells of the vascular envelope significantly improved the plants’ stress response. Surprisingly, NtAQP1 expression in the guard cells did not have a significant effect under any of the tested conditions. The tissue-specific involvement of NtAQP1 in hydraulic and gas conductance via the interaction between the vasculature and the stomata is discussed.     

Carbon dioxide diffusion across stomata and mesophyll and photo-biochemical processes as affected by growth CO2 and phosphorus nutrition in cotton

Nutrients such as phosphorus may exert a major control over plant response to rising atmospheric carbon dioxide concentration (CO₂), which is projected to double by the end of the 21st century. Elevated CO₂ may overcome the diffusional limitations to photosynthesis posed by stomata and mesophyll and alter the photo-biochemical limitations resulting from phosphorus deficiency. To evaluate these ideas, cotton (Gossypium hirsutum) was grown in controlled environment growth chambers with three levels of phosphate (Pi) supply (0.2, 0.05 and 0.01 mM) and two levels of CO₂ concentration (ambient 400 and elevated 800 μmol mol⁻¹) under optimum temperature and irrigation. Phosphate deficiency drastically inhibited photosynthetic characteristics and decreased cotton growth for both CO₂ treatments. Under Pi stress, an apparent limitation to the photosynthetic potential was evident by CO₂ diffusion through stomata and mesophyll, impairment of photosystem functioning and inhibition of biochemical process including the carboxylation efficiency of ribulose-1,5-bisphosphate carboxylase/oxyganase and the rate of ribulose-1,5-bisphosphate regeneration. The diffusional limitation posed by mesophyll was up to 58% greater than the limitation due to stomatal conductance (g_s) under Pi stress. As expected, elevated CO₂ reduced these diffusional limitations to photosynthesis across Pi levels; however, it failed to reduce the photo-biochemical limitations to photosynthesis in phosphorus deficient plants. Acclimation/down regulation of photosynthetic capacity was evident under elevated CO₂ across Pi treatments. Despite a decrease in phosphorus, nitrogen and chlorophyll concentrations in leaf tissue and reduced stomatal conductance at elevated CO₂, the rate of photosynthesis per unit leaf area when measured at the growth CO₂ concentration tended to be higher for all except the lowest Pi treatment. Nevertheless, plant biomass increased at elevated CO₂ across Pi nutrition with taller plants, increased leaf number and larger leaf area.