外来入侵植物对环境梯度和天敌逃逸的适应进化

doi:10.3724/SP.J.1003.2009.09115

摘要/Abstract

摘要：

进化假说认为, 在入侵地外来种能发生遗传变化, 以适应新的环境, 最终成功定殖和扩散。种内或种间杂交、遗传漂变、新环境带来的新的选择压力等是促使外来种发生进化的重要原因。在入侵地, 响应来自非生物和生物因素的选择压力是外来种发生适应进化的主要原因。本文主要介绍外来植物如何通过进化适应入侵地的纬度、海拔等非生物环境和天敌逃逸等生物环境。关于外来入侵植物对纬度和海拔等环境梯度的适应进化, 本文在强调表型进化研究应与分子标记研究相结合的基础上, 介绍了一些同质种植园实验和交互移植实验。关于外来入侵植物对天敌逃逸的适应进化, 本文主要介绍增强竞争能力的进化假说和修正的增强竞争能力的进化假说, 及其在理论上和验证方法上存在的问题。最后, 本文介绍了氮分配的进化假说, 该假说认为天敌逃逸可使外来入侵植物降低叶氮向防御的分配, 同时增加氮向光合的分配。

关键词: 生物入侵, 进化, 局域适应, 遗传多样性, 修正的EICA假说, 氮分配的进化假说

Abstract:

Evolution hypothesis predicts that alien species may experience genetic changes in introduced ranges, facilitating adaptation to new habitats and range expansion. Responses to the selection pressures incurred by the novel abiotic and biotic factors in introduced ranges are primary causes for alien species evolution, although other factors such as intra- and inter-specific hybridizations, genetic shifts can also cause evolution. In this paper, we mainly analyze how alien plant species evolutionarily adapt to environmental changes related to different latitudes and altitudes, and to enemy release in introduced ranges. With respect to evolutionary adaptation to abiotic environments, we introduce some important common garden experiments and reciprocal transplant experiments, emphasizing the necessity to integrate the studies on evolution of phenotypic traits with those on neutral molecular makers. With respect to evolutionary adaptation to enemy release, we mainly discuss evolution of increased competitive ability (EICA) hypothesis, the refinement of EICA hypothesis, and their weaknesses in theory and practices. Finally, we introduce the hypothesis of the evolution of nitrogen allocation, which predicts that invasive plants may decrease leaf nitrogen allocations to defenses but increase allocations to photosynthesis in response to enemy release in introduced ranges.

Key words: biological invasions, refinement of EICA hypothesis, genetic diversity, local adaptation, the hypothesis of the evolution of nitrogen allocation

冯玉龙, 廖志勇, 张茹, 郑玉龙, 李扬苹, 类延宝. 外来入侵植物对环境梯度和天敌逃逸的适应进化[J]. 生物多样性, 2009, 17(4): 340-352.

Yulong Feng, Zhiyong Liao, Ru Zhang, Yulong Zheng, Yangping Li, Yanbao Lei. Adaptive evolution in response to environmental gradients and enemy release in invasive alien plant species[J]. Biodiv Sci, 2009, 17(4): 340-352.

图/表 1

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特性 Characteristics	EICA假说 EICA Hypothesis	修正的EICA假说 The Refinement of EICA Hypothesis	氮分配的进化假说 The Hypothesis of the Evolution of Nitrogen Allocation
是否区分专性天敌和广谱天敌的作用 Whether differentiate the roles of specialist and generalist enemies	不区分 Not differentiate	区分 Differentiate	区分 Differentiate
逃逸的天敌类型 Types of enemies escaped by invasive plants	专性天敌和广谱天敌Specialists and generalists	专性天敌 Specialists	专性天敌和广谱天敌(不总是) Specialists and generalists (not always)
专性天敌抗性(量的防御物质含量)的变化 Changes in resistance to specialists (contents of chemicals related to quantitative defense)	降低 Decrease	降低 Decrease	降低 Decrease
广谱天敌抗性(质的防御物质含量)的变化 Changes in resistance to generalists (contents of chemicals related to qualitative defense)	降低 Decrease	升高 Increase	降低(当天敌不危害入侵种时)或升高(当天敌危害入侵种时) Decrease (when enemies do not attack invaders) or increase (when enemies attack invaders)
竞争能力的变化 Changes in competitive ability	升高 Increase	升高 Increase	升高 Increase
再分配的资源 Resources reallocated	生物量(碳和能量) Biomass (carbon and energy)	生物量(碳和能量) Biomass (carbon and energy)	氮 Nitrogen
分配的部位 Where allocated	“生长”和“防御” “Growth” and “Defenses”	“生长”和“防御” “Growth” and “Defenses”	叶绿体、细胞壁中的蛋白质、含氮化学防御物质 Chloroplast, proteins in cell walls, and nitrogen-based defensive chemicals
是否阐明竞争能力提高的机制 Whether elucidate the mechanisms underlying increased competitive ability	未阐明 Not elucidate	未阐明 Not elucidate	阐明了,即通过提高资源捕获能力和利用效率 Elucidate, i.e. via increasing resource capture ability and use efficiency

特性 Characteristics	EICA假说 EICA Hypothesis	修正的EICA假说 The Refinement of EICA Hypothesis	氮分配的进化假说 The Hypothesis of the Evolution of Nitrogen Allocation
是否区分专性天敌和广谱天敌的作用 Whether differentiate the roles of specialist and generalist enemies	不区分 Not differentiate	区分 Differentiate	区分 Differentiate
逃逸的天敌类型 Types of enemies escaped by invasive plants	专性天敌和广谱天敌Specialists and generalists	专性天敌 Specialists	专性天敌和广谱天敌(不总是) Specialists and generalists (not always)
专性天敌抗性(量的防御物质含量)的变化 Changes in resistance to specialists (contents of chemicals related to quantitative defense)	降低 Decrease	降低 Decrease	降低 Decrease
广谱天敌抗性(质的防御物质含量)的变化 Changes in resistance to generalists (contents of chemicals related to qualitative defense)	降低 Decrease	升高 Increase	降低(当天敌不危害入侵种时)或升高(当天敌危害入侵种时) Decrease (when enemies do not attack invaders) or increase (when enemies attack invaders)
竞争能力的变化 Changes in competitive ability	升高 Increase	升高 Increase	升高 Increase
再分配的资源 Resources reallocated	生物量(碳和能量) Biomass (carbon and energy)	生物量(碳和能量) Biomass (carbon and energy)	氮 Nitrogen
分配的部位 Where allocated	“生长”和“防御” “Growth” and “Defenses”	“生长”和“防御” “Growth” and “Defenses”	叶绿体、细胞壁中的蛋白质、含氮化学防御物质 Chloroplast, proteins in cell walls, and nitrogen-based defensive chemicals
是否阐明竞争能力提高的机制 Whether elucidate the mechanisms underlying increased competitive ability	未阐明 Not elucidate	未阐明 Not elucidate	阐明了,即通过提高资源捕获能力和利用效率 Elucidate, i.e. via increasing resource capture ability and use efficiency