C3和C4植物光合途径的适应性变化和进化

doi:10.3773/j.issn.1005-264x.2009.01.023

摘要/Abstract

摘要：

高等植物大多为C₃植物, C₄植物和景天酸代谢(Crassulacean acid metabolism, CAM)植物是由C₃植物进化而来的。C₄途径的多源进化表明, 光合途径由C₃途径向C₄途径的转变相对简单。该文分析研究了植物光合途径的进化前景, 指出C₄植物是从C₃植物进化而来的高光效种类, 且地质时期以来降低的大气CO₂浓度和升高的大气温度以及干旱和盐渍化是C₄途径进化的外部动力。C₃植物的C₄途径的发现说明植物的光合途径并非是一成不变的, C₃和C₄植物的光合特征具有极大的可塑性, 某些环境的变化会引起植物光合途径在C₃和C₄途径之间转变。C₃植物具有的C₄途径是环境调控的产物, 是对逆境的适应性进化结果, 因而光合途径的转变也适用于干旱地区植被的适应性生存机理研究。该文还利用国外最新的C₄光合进化模型介绍了植物在进化C₄途径中所经历的7个重要时期(从分子基础到形态基础、结构基础, 再到物质代谢水平、光合酶活水平, 直到C₃和C₄途径协调运转时期, 最后达到形态与功能最优化阶段), 并结合全球气候变化的特点对国内外相关领域的研究进行了分析, 总结了植物光合途径的适应性转变和进化的研究成果, 为今后的相关工作提出建议。

关键词: C₃植物, C₄植物, C₄途径, 干旱, 适应性进化

Abstract:

Three photosynthetic pathways are used by plants: C₃, which most plants use, C₄ and crassulacean acid metabolism (CAM). C₃ is the ancestral pathway, with C₄ and CAM representing recently diverged forms. We conclude that the variation and evolution of photosynthetic pathway of C₃ and C₄ are adaptations to environmental stresses. First, we discuss the evolutional future of photosynthetic pathways. It has been suggested that low atmosphere CO₂, enhanced temperatures, drought and salinity are external drivers of C₄ photosynthetic evolution. Second, we analyze the possibility of evolution from C₃to C₄. The polyphyletic evolution of the C₄ pathway suggests that the transition from C₃ to C₄ was relatively simple. This suggestion is supported by the observation that both C₃ and C₄ plants possess inherent plasticity in their photosynthetic characteristics. The stress causing the shift from C₃ to C₄ was involved in the environmental regulation of plants, and the C₄ pathway in C₃ plants evolved as an adaptation. The environmental stresses may have involved plant capacities for survivorship and competition in arid areas. Third, we present a simplistic model of the main phases of C₄ photosynthetic evolution and discuss the variation and evolution of C₃and C₄photosynthetic carbon assimilating pathways. Evolution was not directly to C₄ photosynthesis, so each step had to be stable, beginning with numerous preconditions needed for an evolutionary lineage to begin evolving C₄ characteristics. A current complication of C₄ photosynthetic evolution is global climate change and human manipulation of the biosphere. It is possible that C₄ photosynthesis can be used to resist adversity and increase the yield of C₃ crops. In conclusion, study of the evolution of photosynthetic pathway in plants provided insight into the photosynthetic physiology of plants under stress and provided new theory to reconstruct vegetation, enhance crop yield, and explain adaptation of C₃ species in arid areas.

Key words: C₃ plants, C₄ plants, C₄ pathway, drought, adaptable evolution

龚春梅, 宁蓬勃, 王根轩, 梁宗锁. C₃和C₄植物光合途径的适应性变化和进化. 植物生态学报, 2009, 33(1): 206-221. DOI: 10.3773/j.issn.1005-264x.2009.01.023

GONG Chun-Mei, NING Peng-Bo, WANG Gen-Xuan, LIANG Zong-Suo. A REVIEW OF ADAPTABLE VARIATIONS AND EVOLUTION OF PHOTOSYNTHETIC CARBON ASSIMILATING PATHWAY IN C₃ AND C₄ PLANTS. Chinese Journal of Plant Ecology, 2009, 33(1): 206-221. DOI: 10.3773/j.issn.1005-264x.2009.01.023

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链接本文: https://www.plant-ecology.com/CN/10.3773/j.issn.1005-264x.2009.01.023

https://www.plant-ecology.com/CN/Y2009/V33/I1/206

图/表 4

图1 第三纪以来大气CO2浓度的变化(引自Sage(2004), 略有改动) ○: 硼同位素测定的大气CO2浓度 Atmospheric CO2 concentration measured by using boron isotopes method : 烯酮片断中的碳同位素比率 Carbon isotope ratio in alkenone fragments : 4千万年前大气CO2浓度的变化 The best-fit regression to the CO2 data older than 40 million years : C4植物发生发展的关键时期 Key developmental period in the history of C4 plants

Fig. 1 Changes in atmospheric CO2 concentration since the beginning of the Tertiary Period (ca. 65 million years ago)(Adapted from Sage, 2004)

图2 C4途径在被子植物分类学上的进化路线 (引自Sage (2004), 略有改动) 被子植物进化C4途径的目的顺序用粗体字表示, 直线右侧是每一目主要的C4科、属, 括号中的数字是估计的C4属/种数或物种数 Angiosperm orders with C4 photosynthetic carbon assimilating pathway are shown in bold. Lines to the right of these orders indicate families and principal C4 genera within a lineage. Numbers in parentheses refer to estimates of genera/species numbers or, where relevant, just species numbers

Fig. 2 Evolutional course of C4 pathway in the taxonomic orders of angiosperms (Adapted from Sage, 2004)

图3 两种单一光合细胞的C4植物叶解剖结构(引自Sage (2004), 略有改动) A: 异子蓬 Borszczowia aralocaspica B: Bienertia cycloptera P: PEPCase的分布区 Distribution region of PEPCase R: Rubisco的分布区 Distribution region of Rubisco

Fig. 3 Leaf anatomy of the two single-celled plants with the C4 pathway of organic acid flux in each cell highlighted (Adapted from Sage, 2004)

图4 包含C4途径主要进化时期的最简金字塔模型(引自Sage (2004), 略有改动)

Fig. 4 An ascending pyramid of C4 pathway: a simplistic model of the main phases of C4 evolution (Adapted from Sage, 2004)

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C₃和C₄植物光合途径的适应性变化和进化

A REVIEW OF ADAPTABLE VARIATIONS AND EVOLUTION OF PHOTOSYNTHETIC CARBON ASSIMILATING PATHWAY IN C₃ AND C₄ PLANTS

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