植物光合作用叶肉导度及主要限制因素研究进展

doi:10.17521/cjpe.2016.0337

摘要/Abstract

摘要：

叶肉导度(g_m)被用来衡量CO₂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。

关键词: 光合作用, CO2传输, 叶肉导度, 解剖结构, 生化因素, 环境变化, 水力导度

Abstract:

Mesophyll conductance (g_m) represents the CO₂ diffusion facility from sub-stomatal internal cavities to carboxylation sites in chloroplasts, and the variation of g_m across genotypes as well as environmental conditions is expected to be related to the anatomical structures and biochemical properties of leaves. In recent years, the variation of g_m has attracted wide attention. The limiting factors in photosynthetic rate are no longer divided simply into stomatal limitation and non-stomatal limitation, but splitted in stomatal limitation, mesophyll limitation and carboxylation limitation. In this review, we summarize the potential influences of cell wall, cell membrane, cytoplasm, chloroplast envelope and stroma on g_m, and indicate that cell wall thickness and the surface area of chloroplast exposed to intercellular air space (S_c) are the most important factors influencing the g_m. We also analyze the probable effects of biochemical process related with aquaporins and carbonic anhydrase on g_m. Meanwhile, the regulation mechanisms of long- and short-term environment changes (including temperature, light intensity, drought, and nutrients) on g_m are also summarized. The relationship between g_m and hydraulic conductance (K_leaf) is debated. Finally, we discuss the scientific problems related with g_m.

Key words: photosynthesis, CO2 diffusion, mesophyll conductance, anatomical structure, biochemical factor, environmental change, hydraulic conductance

韩吉梅, 张旺锋, 熊栋梁, 张亚黎. 植物光合作用叶肉导度及主要限制因素研究进展. 植物生态学报, 2017, 41(8): 914-924. DOI: 10.17521/cjpe.2016.0337

Ji-Mei HAN, Wang-Feng ZHANG, Dong-Liang XIONG, Jaume FLEXAS, Ya-Li ZHANG. Mesophyll conductance and its limiting factors in plant leaves. Chinese Journal of Plant Ecology, 2017, 41(8): 914-924. DOI: 10.17521/cjpe.2016.0337

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2016.0337

https://www.plant-ecology.com/CN/Y2017/V41/I8/914

图/表 3

图1 CO2运输模式图。AQPs, 水孔蛋白; Ca, 大气CO2浓度; Ci, 胞间CO2浓度; CA, 碳酸酐酶; gias, 气相导度; glip, 液相导度。

Fig. 1 CO2 transport model. AQPs, aquaporins; Ca, the atmospheric CO2 concentration; Ci, intercellular CO2 concentration; CA, carbonic anhydrase; gias, the gas phase conductance; glip, the liquid phase conductance.

图2 gm反映的CO2扩散路径。A, 光学显微镜拍摄的棉花叶片解剖结构图,代表的是CO2从外界大气进入叶片细胞间隙,完成气相传输。B, 电子显微镜拍摄的棉花叶片超微结构图。代表的是CO2从细胞间隙进入叶绿体羧化位点所经过的部位,完成液相传输。图中简单介绍了影响传输路径的因素及需要进一步深入研究的问题。AQPs, 水孔蛋白; Ci, 胞间CO2浓度; Cc, 叶绿体羧化位点CO2浓度; CA, 碳酸酐酶, gias, 液相导度。

Fig. 2 The diffusion path of CO2 reflected by gm. A, The leaf anatomical structure in cotton by optical microscope, which represents the CO2 gas phase diffusion from the atmosphere into the leaf intercellular air layer; B, The leaf ultra-micro structure in cotton by electron microscope, which represents the CO2 liquid phase diffusion from intercellular into the chloroplast carboxylation site. AQPs, aquaporins; Ci, intercellular CO2 concentration; Cc, CO2 concentration at chloroplast carboxylation site; CA, carbonic anhydrase; gias, the gas phase conductance.

表1 CO2通过叶肉细胞中各超微组分的扩散方式、运输形态、阻力来源、动力来源和对外界环境的响应时间等的差异

Table 1 Diffusion way, transportation form, resistance source, power source when CO2 passes through the ultrastructure components of mesophyll cells and the different response time to the external environment

	CO₂扩散方式 CO₂ diffusion way	CO₂运输形态 CO₂transportation form	阻力来源 Resistance source	动力来源 Power source	对外界环境的响应时间 Response time to the external environment
细胞壁 Cell wall	物理和生化方式 Physics and biochemical mode	CO₂	厚度、孔隙度、果胶等组分 Thickness, porosity, pectin etc.	CO₂浓度差 Difference of CO₂ concentration	最长 Longest
细胞膜 Cell membrane	物理和生化方式 Physics and biochemical mode	CO₂	水孔蛋白、膜两侧pH差值 AQPs, the difference of pH on both sides of the membrane	CO₂浓度差、跨膜蛋白主动运输 Difference of CO₂ concentration, active transport of transmembrane protein	较短 Shorter
细胞液 Cytoplasm	生化和物理方 Biochemical and physical mode	CO₂, HCO₃^-	CA、pH、细胞液组分 CA, pH, cytosol component	pH、CA的催化 pH, catalysis of CA	较短 Shorter
叶绿体膜 Chloroplast membranes	生化和物理方 Biochemical and physical mode	CO₂	水孔蛋白、膜两侧CO₂浓度差 AQPs, the difference of CO₂ concentration on both sides of the membrane	跨膜蛋白主动运输 Active transport of transmembrane protein	较短 Shorter
叶绿体基质 Stroma	生化和物理方式 Biochemical and physical mode)	CO₂, HCO₃^-	CA, pH	pH、CA的催化 pH, catalysis of CA	最短 Shortest

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