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在叶水平中,通常至少由具有同化结构的叶片和悬臂结构的叶柄组成,叶片是植物光合作用的主要构件,是植物自身能量的主要来源[1-3]。叶柄是植物叶支撑系统的主要构件,对植物的水分和营养的运输起关键性作用[4]。一般来说,若叶的总生物量多,分配给同化结构的生物量多,叶片的光合作用强度大,增强了叶的碳获取能力,同时叶内支撑投资随着叶内生物量的增加递增[5-6],两者之间存在权衡关系,管道模型(pipe model)理论证实了以上观点,即为满足叶片对水分的需求,输导结构的横截面积应与其上的叶面积成等比例。但是叶柄除了为叶片输送水分之外,还需要为叶面提供机械支撑,承受自身及叶片引起的重力,具有支撑整个叶的静态重力和动态的机械拉扯引起的外力,如风力等外力为支撑结构增加了负荷。由此可以看出叶内生物量分配策略错综复杂,叶片和叶柄之间存在异速生长关系[7-9]。通过近几年学者们的研究发现,存在这种关系的原因主要是叶内生物量分配的差异,这些差异受一些因素的响应。研究主要通过生活型、叶型、叶大小、叶形状及叶含水量这几方面论述对叶内生物量权衡过程的影响,对未来叶生物量分配策略的研究奠定基础。
Research progress on the response of intra-leaf biomass allocation strategies to life forms and leaf traits
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摘要: 叶片是光合作用的主要场所,通过光合作用固定的能量用于呼吸消耗和同化作用,进行物质生产的同时进行叶内生物量的分配,这种分配并不是柄叶之间生物量发生等比例变化,而是由于自身及外界因素的影响使柄叶之间出现异速生长关系,即柄叶权衡。目前未发现叶内生物量分配策略对生活型及叶性状响应的评述,研究主要对叶内生物量分配对不同生活型、叶型、叶大小、叶形状和叶含水量的响应机制进行了论述。并提出今后需要加强研究的几方面:(1)探索全部或者部分支持构件与叶内生物量分配之间的关系,揭示不同占比的支持结构是如何影响叶内生物量分配策略的,各个支持结构在叶内生物量分配中扮演什么样的“角色”;(2)探索叶序与生物量分配的关系,揭示光合构件在不同排列方式下进行碳收益过程中叶内生物量如何分配,不同排列方式下的叶片水分养分输导过程对叶内生物量分配的影响又是如何;(3)探索叶柄及其他支撑结构与叶性状的作用关系;(4)探索叶经济谱(LES)作用范围,揭示LES是否包括叶片和叶柄的作用关系。Abstract: Leaves are the main places for photosynthesis. Through photosynthesis, the energy fixed by photosynthesis is used for respiration consumption and assimilation, and the biomass in leaves is allocated at the same time of material production. This allocation is not a proportional change in biomass between stalks and leaves, but an allometric growth relationship between stalks and leaves, that is, the balance between stalks and leaves, due to the influence of their own and external factors. At present, there is no review on the response of intra-leaf biomass allocation strategies to life forms and leaf traits. This paper mainly discusses the response mechanisms of intra-leaf biomass allocation to different life forms, leaf types, leaf sizes, leaf shapes and leaf water content. Several aspects that need to be further studied in the future are proposed: (1) Explore the relationship between all or part of the supporting components and intra-leaf biomass allocation, and reveal how the supporting structure of different proportions affected the leaf biomass allocation strategies, and what kind of 'role' each supporting structure played in leaf biomass allocation; (2) Explore the relationship between phyllotaxis and biomass allocation, and reveal how the intra-leaf biomass was allocated during the carbon gain process of photosynthetic components under different arrangement modes, and how the leaf water and nutrient transport process under different arrangement modes affected the intra-leaf biomass allocation. (3) Explore the relationship between petiole and other supporting structures and leaf traits; (4) Explore the range of leaf economic spectrum (LES) and reveal whether LES includes the relationship between leaves and petioles.
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