植物光合作用叶肉导度及主要限制因素研究进展

叶肉导度(g_m)被用来衡量CO_2从植物叶片气孔下腔到叶绿体羧化位点的传输效率,其主要受解剖结构和生化因素的调控。近年来,g_m的研究在光合作用领域受到普遍关注;光合速率的限制因素已不再简单地划分为气孔限制和非气孔限制,而需要从气孔限制、叶肉限制和羧化限制3个限制因素开展研究工作。该文分析了植物细胞壁、细胞膜、细胞质、叶绿体膜和叶绿体基质对g_m的调控机制,指出细胞壁厚度以及面向细胞间隙的叶绿体面积(S_c)是影响g_m的重要结构因素。阐述了水孔蛋白和碳酸酐酶参与的生化过程对g_m的调控机制。同时,对外界环境因素,如温度、光强、干旱、氮等对g_m的调控机制进行了总结。在此基础上,探讨了g_m与水力导度的耦合关系。最后对g_m研究中的科学问题进行了展望。     

植物叶肉导度的测定方法及其研究进展

叶肉导度(g_m)表征CO₂从植物气孔下腔到叶绿体内光合作用羧化位点的传输导度,是光合作用的重要限制因子,也是改良作物光合作用和资源利用效率的关键参数。¹³C分馏在线联测法是研究植物光合生理的重要手段,也是目前测定C3植物g_m的主流方法之一。但该方法对实验设备要求较高、难度较大,因此尚未被广泛采用。本文综述了当前测定g_m的主流方法的适用性和特点;介绍了¹³C分馏在线联测法的理论基础——光合¹³C分馏模型,¹³C分馏在线联测法的实测方法、计算公式和测定系统的搭建,以及C3植物叶肉导度的调控因素。最后对¹³C分馏在线联测法的改进、操作要点和未来应用场景进行了展望。     

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。     

北方粳稻光合速率、气孔导度对光强和CO2浓度的响应

 以东北地区主栽的粳稻（Oryza sativa var. japonica）品种为对象,用美国LI-cor公司生产的Li 6400光合作用测定仪控制光强、CO2浓度和温度等环境条件,阐述了光合作用和气孔导度对光和CO2浓度的响应特征及其耦合关系。结果表明,光合速率随光强或CO2浓度的提高而增大,均遵循米氏响应;在不同CO2浓度下,表观量子效率随CO2浓度的提高而增大,但CO2浓度达到800 μmol•mol－1以上时,表观量子效率有所减小;在不同光强下,表观羧化效率也随光的增强而增大,但光强达到1 600 μmol•m-2•s-1以上时,表观羧化效率也有所减小;在光强和CO2浓度协同作用下,光合速率的响应遵循双底物的米氏方程,在光强和CO2浓度均趋于饱和时,北方粳稻（品种：辽粳294）剑叶的潜在最大光合速率为71.737 8 μmol•m-2•s-1,表观量子效率为0.056 0 μmolCO2•μmol-1 photons,表观羧化效率为0.103 1 μmol•m-2•s-1/μmol•mol－1。气孔导度也随光的增强而增大,对光强的响应规律也可以用Michaelis-Menten曲线模拟,而叶面CO2浓度的提高会使气孔导度减小,气孔导度（Gs）对叶面CO2浓度（Cs）的响应可以用Gs=Gmax,c/(1+Cs/Cs0)的双曲线方程模拟。在光强(PFD)和CO2浓度协同作用下,气孔导度可以用式Gs=Gmax(PFD/PFDc)/［(1+PFD/PFDc)(1+Cs/Cs0)］+Gct估算,当CO2浓度趋于0而光强趋于饱和时,北方粳稻的潜在最大气孔导度（Gmax）为0.670 9 mol•m-2•s-1。在光强和CO2浓度协同作用下,Ball-Berry模型及其修正形式依然能很好地表达气孔导度-光合速率的耦合关系,并且用叶面饱和水汽压差（Ds）修正耦合关系中的相对湿度可以提高模拟精度。     

干旱生境中接种丛枝菌根真菌对三叶鬼针草

为阐明丛枝菌根真菌对石灰岩地区适生植物三叶鬼针草(Bidens pilosa L.)光合作用的影响,设置正常浇水(A)、中度干旱胁迫(B)和重度干旱胁迫(C)3个水分处理梯度,比较了不同水分处理条件下接种丛枝菌根真菌Glomus mosseae和未接种三叶鬼针草之间净光合速率、气孔导度、蒸腾速率、胞间CO2浓度、羧化效率、水分利用效率等特征的差异.结果表明,水分胁迫显著降低三叶鬼针草的净光合速率、气孔导度、蒸腾速率和羧化效率.胞间CO2浓度在处理的前期(7d)因干旱胁迫而降低,在后期随土壤含水量的降低而升高;水分利用效率则是中度胁迫的植株、正常浇水处理植株、重度胁迫植株依次降低.在正常浇水条件下接种G. mosseae 对三叶鬼针草光合参数没有产生显著性影响;在中度胁迫条件下,接种植株较未接种植株在水分处理的前28d有更高的净光合速率、气孔导度、蒸腾速率和羧化效率;在重度胁迫条件下,虽然净光合速率、气孔导度、蒸腾速率和羧化效率接种植株高于未接种植株,但是二者并不显著.研究认为,干旱胁迫对三叶鬼针草光合作用的影响在水分处理的前期表现为气孔因素制约,在后期则主要是非气孔因素的影响;在正常浇水条件下接种G. mosseae 对三叶鬼针草的光合作用没有显著性影响,在干旱胁迫条件下,丛枝菌根真菌通过改善三叶鬼针草气孔导度和羧化效率等减弱干旱胁迫对植株的伤害,但是这种保护作用因为土壤水分的严重匮乏以及土壤干旱的时间延长而受到限制.     

杨品种‘84K’光合生理的不同过程对干旱和复水的响应

以杨品种‘84K’（Populus alba×P.glandulosa）为材料,对其干旱胁迫及复水后光合生理特性的变化进行了研究。结果显示,在干旱胁迫及复水过程中,杨品种‘84K’光合作用相关的主要反应对此过程响应不同步。在干旱胁迫过程中,‘84K’的羧化反应速度、气孔导度（G_s）、叶肉导度（G_m）均显著下降,但前者下降幅度小于后两者,此时的光合作用主要受G_s和G_m制约。复水之后,G_m很快得到恢复,而光化学淬灭过程、羧化反应速度均没有恢复到对照水平,此时是光化学淬灭和（或）羧化反应制约了‘84K’的碳固定及光合作用。     

叶肉细胞导度研究进展

叶肉细胞导度指叶片叶肉细胞内部的CO2扩散能力,在植物生理生态及全球气候变化和陆地生态系统相互关系的研究中具有重要作用。系统介绍了叶肉细胞导度的发现、发展过程及其研究进展、几种目前国际上常用的叶肉细胞导度测度方法的原理、计算过程;强调了叶肉细胞导度作为光合作用扩散过程一部分的重要意义,明确了叶肉细胞导度的定义及分布范围。并探讨了不同方法的优缺点及注意事项。总结分析了叶肉细胞导度对不同环境因子(温度、水分及环境中CO2和O3浓度等)的响应,从不同角度对叶肉细胞导度的生态学意义进行了简单的概括。对叶肉细胞导度的未来研究进行了展望。     

杨品种‘84K’光合生理的不同过程对干旱和复水的响应

以杨品种‘84K’(Populus alba×P. glandulosa)为材料,对其干旱胁迫及复水后光合生理特性的变化进行了研究。结果显示,在干旱胁迫及复水过程中,杨品种‘84K’光合作用相关的主要反应对此过程响应不同步。在干旱胁迫过程中,‘84K’的羧化反应速度、气孔导度(Gs)、叶肉导度(Gm)均显著下降,但前者下降幅度小于后两者,此时的光合作用主要受Gs和Gm制约。复水之后,Gm很快得到恢复,而光化学淬灭过程、羧化反应速度均没有恢复到对照水平,此时是光化学淬灭和(或)羧化反应制约了‘84K’的碳固定及光合作用。     

海拔梯度对巴郎山奇花柳叶片δ13C的影响

2010年在四川卧龙自然保护区选择海拔为2350、2700、3150和3530 m的4个分布地点,研究了巴郎山海拔梯度对奇花柳叶片13C、光合、CO2扩散导度、氮含量、光合氮利用效率( PNUE)和比叶面积(SLA)的影响.结果表明:随着海拔的升高,目标树种叶片氮含量(尤其是单位面积氮含量)及PNUE增加,叶片δ13C值也随之显著增加,且海拔每升高1000 m,δ13C增加1.4‰;CO2扩散导度(气孔导度和叶肉细胞导度)的增加,在一定程度上阻碍了叶片δ13C值随海拔升高,但不足以改变δ13C值随海拔升高的趋势;羧化能力是羧化位点与外界CO2分压比( Pc/Pa),甚至δ13C的限制因子.在海拔2350～2700m,奇花柳光合系统内部氮素分配主要受温度的影响,而2700～3530 m的光照作用可能更大.奇花柳的SLA随海拔无显著变化.     

水分胁迫条件下气孔与非气孔因素对小麦光合的限制

较长时间、中度以上土壤干旱条件下,盆栽小麦叶片光合下降,气孔导度虽然降低,但叶内部CO_2浓度升高,叶肉CO_2导度下降,电子传递和羧化反应受到抑制,表明叶片光合的下降并非气孔部分关闭所致,叶肉细胞光合能力降低是干旱下小麦光合下降的原因。     

Rapid variations of mesophyll conductance in response to changes in CO2 concentration around leaves

The effects of short-term (minutes) variations of CO2 concentration on mesophyll conductance to CO2 (gm) were evaluated in six different C3 species by simultaneous measurements of gas exchange, chlorophyll fluorescence, online carbon isotope discrimination and a novel curve-fitting method. Depending on the species, gm varied from five- to ninefold, along the range of sub-stomatal CO2 concentrations typically used in photosynthesis CO2-response curves (AN)-Ci curves; where AN is the net photosynthetic flux and Ci is the CO2 concentrations in the sub-stomatal cavity), that is, 50 to 1500 micromol CO2 mol(-1) air. Although the pattern was species-dependent, gm strongly declined at high Ci, where photosynthesis was not limited by CO2, but by regeneration of ribulose-1,5-bisphosphate or triose phosphate utilization. Moreover, these changes on gm were found to be totally independent of the velocity and direction of the Ci changes. The response of gm to Ci resembled that of stomatal conductance (gs), but kinetic experiments suggested that the response of gm was actually faster than that of gs. Transgenic tobacco plants differing in the amounts of aquaporin NtAQP1 showed different slopes of the gm-Ci response, suggesting a possible role for aquaporins in mediating CO2 responsiveness of gm. The importance of these findings is discussed in terms of their effects on parameterization of AN-Ci curves.     

Potassium mediates coordination of leaf photosynthesis and hydraulic conductance by modifications of leaf anatomy

Typical symptoms of potassium deficiency, characterized as chlorosis or withered necrosis, occur concomitantly with downregulated photosynthesis and impaired leaf water transport. However, the prominent limitations and mechanisms underlying the concerted decreases of leaf photosynthesis and hydraulic conductance are poorly understood. Monocots and dicots were investigated based on responses of photosynthesis and hydraulic conductance and their components and the correlated anatomical determinants to potassium deficiency. We found a conserved pattern in which leaf photosynthesis and hydraulic conductance concurrently decreased under potassium starvation. However, monocots and dicots showed two different hydraulic‐redesign strategies: Dicots tended to show a decreased minor vein density, whereas monocots reduced the size of the bundle sheath and its extensions, rather than the minor vein density; both of these strategies may restrain xylem and outside‐xylem hydraulic conductance. Additionally, potassium‐deprived leaves developed with fewer mesophyll cell‐to‐cell connections, leading to a reduced area being available for liquid‐phase flow. Further quantitative analysis revealed that mesophyll conductance to CO₂ and outside‐xylem hydraulic resistance were the major contributors to photosynthetic limitation and increased hydraulic resistance, at more than 50% and 60%, respectively. These results emphasize the importance of potassium in the coordinated regulation of leaf photosynthesis and hydraulic conductance through modifications of leaf anatomy.     

Estimation of Mesophyll Conductance to CO(2) Flux by Three Different Methods

The resistance to diffusion of CO₂ from the intercellular airspaces within the leaf through the mesophyll to the sites of carboxylation during photosynthesis was measured using three different techniques. The three techniques include a method based on discrimination against the heavy stable isotope of carbon, ¹³C, and two modeling methods. The methods rely upon different assumptions, but the estimates of mesophyll conductance were similar with all three methods. The mesophyll conductance of leaves from a number of species was about 1.4 times the stomatal conductance for CO₂ diffusion determined in unstressed plants at high light. The relatively low CO₂ partial pressure inside chloroplasts of plants with a low mesophyll conductance did not lead to enhanced O₂ sensitivity of photosynthesis because the low conductance caused a significant drop in the chloroplast CO₂ partial pressure upon switching to low O₂. We found no correlation between mesophyll conductance and the ratio of internal leaf area to leaf surface area and only a weak correlation between mesophyll conductance and the proportion of leaf volume occupied by air. Mesophyll conductance was independent of CO₂ and O₂ partial pressure during the measurement, indicating that a true physical parameter, independent of biochemical effects, was being measured. No evidence for CO₂-accumulating mechanisms was found. Some plants, notably Citrus aurantium and Simmondsia chinensis, had very low conductances that limit the rate of photosynthesis these plants can attain at atmospheric CO₂ level.     

The role of mesophyll conductance in the economics of nitrogen and water use in photosynthesis

A recent resurgence of interest in formal optimisation theory has begun to improve our understanding of how variations in stomatal conductance and photosynthetic capacity control the response of whole plant photosynthesis and growth to the environment. However, mesophyll conductance exhibits similar variation and has similar impact on photosynthesis as stomatal conductance; yet, the role of mesophyll conductance in the economics of photosynthetic resource use has not been thoroughly explored. In this article, we first briefly summarise the knowledge of how mesophyll conductance varies in relation to environmental factors that also affect stomatal conductance and photosynthetic capacity, and then we use a simple analytical approach to begin to explore how these important controls on photosynthesis should mutually co-vary in a plant canopy in the optimum. Our analysis predicts that when either stomatal or mesophyll conductance is limited by fundamental biophysical constraints in some areas of a canopy, e.g. reduced stomatal conductance in upper canopy leaves due to reduced water potential, the other of the two conductances should increase in those leaves, while photosynthetic capacity should decrease. Our analysis also predicts that if mesophyll conductance depends on nitrogen investment in one or more proteins, then nitrogen investment should shift away from Rubisco and towards mesophyll conductance if hydraulic or other constraints cause chloroplastic CO₂ concentration to decline. Thorough exploration of these issues awaits better knowledge of whether and how mesophyll conductance is itself limited by nitrogen investment, and about how these determinants of photosynthetic CO₂ supply and demand co-vary among leaves in real plant canopies.     

Theoretical Considerations when Estimating the Mesophyll Conductance to CO(2) Flux by Analysis of the Response of Photosynthesis to CO(2)

The conductance for CO₂ diffusion in the mesophyll of leaves can limit photosynthesis. We have studied two methods for determining the mesophyll conductance to CO₂ diffusion in leaves. We generated an ideal set of photosynthesis rates over a range of partial pressures of CO₂ in the stroma and studied the effect of altering the mesophyll diffusion conductance on the measured response of photosynthesis to intercellular CO₂ partial pressure. We used the ideal data set to test the sensitivity of the two methods to small errors in the parameters used to determine mesophyll conductance. The two methods were also used to determine mesophyll conductance of several leaves using measured rather than ideal data sets. It is concluded that both methods can be used to determine mesophyll conductance and each method has particular strengths. We believe both methods will prove useful in the future.     

Effects of Powdery Mildew Infection on the Efficiency of CO(2) Fixation and Light Utilization by Sugar Beet Leaves

Sugar beet leaves (Beta vulgaris L.) infected with powdery mildew (Erysiphe polygoni D.C.) show declining rates of net photosynthesis as the disease develops; relative to healthy controls, reductions of 35, 70, and 75% were observed at 9, 16, and 22 days after inoculation, respectively. A leaf gas exchange procedure in which an air stream flowed through the leaf showed that mesophyll conductance declined in parallel with photosynthesis in mildew-infected leaves. Viscous flow conductance of diseased leaves also declined over the same period suggesting that stomatal aperture was reduced. From the magnitude and time course of disease effects on stomatal aperture and mesophyll conductance, it appears that the effects at the mesophyll level were primarily responsible for mediating the decline in net photosynthesis. Changes in mesophyll conductance were closely correlated with reduced activity of ribulose-1,5-bisphosphate carboxylase on a leaf area basis. This decrease could be attributed to a reduction in the concentration of the enzyme, a reduction which was greater than the reduction in total soluble protein. The quantum efficiency of light use was also decreased by the disease. Mildew-infected leaves had quantum yields that were reduced, relative to healthy leaves, by 17 and 22% at 14 and 18 days after inoculation, respectively.     

Salt impact on photosynthesis and leaf ultrastructure of <Emphasis Type="Italic">Aeluropus littoralis</Emphasis>

The effects of salinity (400 mM NaCl) on growth, biomass partitioning, photosynthesis, and leaf ultrastructure were studied in hydroponically grown plants of Aeluropus littoralis (Willd) Parl. NaCl produced a significant inhibition of the main growth parameters and a reduction in leaf gas exchange (e.g. decreased rates of photosynthesis and stomatal conductance). However, NaCl salinity affected neither the composition of photosynthesis pigments nor leaf water content. The reduction in leaf gas exchange seemed to correlate with a decrease in mesophyll thickness as well as a severe disorganisation of chloroplast structure, with misshapen chloroplasts and dilated thylakoid membranes. Conspicuously, mesophyll chloroplasts were more sensitive to salt treatment than those of bundle sheath cells. The effects of NaCl toxicity on leaf structure and ultrastructure and the associated physiological implications are discussed in relation to the degree of salt resistance of A. littoralis.     

Stomatal responses to long-term high vapor pressure deficits mediated most limitation of photosynthesis in tomatoes

Plants grown at high vapor pressure deficit (VPD) usually present decreased photosynthesis, but stomatal and mesophyll limitation to photosynthesis remain poorly quantified. To better understand the regulation of high VPD on photosynthesis and plant growth in tomatoes, we investigated the limitation of stomatal conductance and mesophyll conductance to photosynthesis and relative importance of stomatal morphology and function in stomatal conductance. Both the net photosynthesis rate and total biomass were significantly limited by high VPD. Meanwhile, stomatal conductance and mesophyll conductance were decreased under high VPD. The stomatal conductance limitation was responsible for 60% of the total photosynthetic limitation. Moreover, a reduction in stomatal density and stomatal size occurred under high VPD, which was significantly correlated with the down-regulation of stomatal conductance. The stomatal morphology contributed to more than half the change in stomatal conductance. Nevertheless, stomatal movement was also an important factor in regulating stomatal conductance. The decrease of hydraulic conductance and transpiration rate with no significant difference in relative water content, leaf water potential, and/or osmotic potential suggested passive hydraulic regulation in the feedforward responses of stomata to high VPD.     

The effects of soil and atmospheric drought on photosynthesis and stomatal control of gas exchange in three coniferous species

The responses of steady-state CO₂ assimilation rate (A), transpiration rate (E), and stomatal conductance (g_s) to changes in leaf-to-air vapour pressure difference (δW) on one hand and to increasing soil drought on the other hand were examined in 2-year-old seedlings of Pseudotsuga menziesii, Pseudotsuga macrocarpa and Cedrus atlantica. Analysing the data through A vs intercellular CO₂ molar fraction (c_i) graphs, we could determine stomatal and mesophyll contributions to changes in A as δW or soil drought were increased. Increasing soil drought affected g_s and mesophyll photosynthesis independently, since clearly distinct predawn leaf water potential (ψ_p) regions appeared in which either stomatal or mesophyll effects prevailed for explaining the changes in A. The two Pseudotsuga species exhibited a large ψ_P range (between ca -0.8 and -1.5 to -1.9 MPa) in which only stomata were responsible for the decrease in A. A dramatic decline in mesophyll photosynthesis was noticed starting from values as high as -1.2 MPa ( C. atlantica ), -1.5 MPa ( P. macrocarpa ) and -1.9 MPa ( P. menziesii ). Increasing ΔW at high soil water content led to a sharp decline in A primarily due to an alteration of mesophyll photosynthesis. Stomatal conductance for CO₂ diffusion was affected in a lesser extent and in close correlation with the changes in mesophyll photosynthesis, which could suggest the existence of a functional linkage between mesophyll photosynthesis and stomata. Surprisingly, the drought resistant P. macrocarpa exhibited the least conservative water use efficiency in response to the two types of drought. In this species drought adaptation seems to be mainly due to its high root growth and soil prospection ability.