竞争⁃耐胁迫⁃杂草型植物对策理论及其应用研究进展

Research progress of plant competitor⁃stress tolerator⁃ruderal(CSR)theory and its application

竞争⁃耐胁迫⁃杂草型植物(CSR)对策理论主要研究植物对环境胁迫和干扰的适应性特化。从功能生态学角度讲,CSR理论是通过解释各种功能性状协同变异的主导维度来反映植物的生态对策。而物种所采取的生态对策将决定其在群落中的生态位,并最终会影响生态系统过程与功能。自1974年Grime首次提出CSR对策的概念以来,CSR对策思想就受到了生态学家很大的关注,经过近半个世纪的发展,目前已经形成了比较完整的理论和研究方法体系。试图在回顾CSR理论及其研究方法发展的基础上,阐述其基本内涵,并重点从功能性状变异空间、植物适应性、群落过程和生态系统特性等4个方面概述CSR理论及其分析模型的验证及应用,梳理其最新研究进展。CSR理论已经得到了很多研究的广泛验证并应用于多种尺度上的植物生态学研究,但仍有一些方面值得进一步发展完善,如第四区缺失问题、功能性状变异主轴以外的变异、CSR对策与功能性状种内变异和谱系关系的联系等方面需要深入研究。

Competitor,stress⁃tolerator,ruderal(CSR)theory,regarded as one of the best developed plant ecological strategy schemes,mainly studies the adaptive specialisation that plants evolve to achieve fitness in response to natural selection pressures,resulting in a triangular distribution of species optima along axes of habitat productivity and disturbance.From the perspective of functional ecology,the CSR theory reflects plant ecological strategies by explaining the principal functional spectra in the multidimensional trait space,that is,the world⁃wide economics spectrum and the size spectrum as fundamental gradients of plant evolution are major components of the CSR strategy variation.The ecological strategies adopted by plants then determine their ecological niche in the community and ultimately affect ecosystem processes and functions.Thus the CSR theory could not only be used as a framework for functionally classifying plants and predicting impacts of environmental change on vegetation,but also serves as a conceptual framework for investigating and interpreting community processes and ecosystem functioning.Since the concept of CSR strategies was first raised by Grime in 1974,the idea of CSR strategies has attracted much attention from ecologists,and the CSR theory has formed a relatively complete system of theories and research methods over the last half century.In this paper,we introduce the development and characteristics of the CSR theory and its methods to expound the core philosophy of the theory,review the verification and application of the CSR theory and its models by addressing the relevance of CSR strategies to functional trait variations,species adaptation,community processes and ecosystem properties,and summarize the advances in this theory.The CSR theory has been extensively validated by many studies and widely applied as a very useful analysis tool to plant ecology research on a variety of scales,but it needs further research,such as the missing fourth corner in the Grime′s C⁃S⁃R triangular model,the rest functional variations beyond the principally functional trait spectra that are not explained by the CSR scheme,and the relationships between CSR strategies and the intraspecific trait variation and phylogenetic signal.Especially for the rest functional variations,the CSR theory does not preclude the existence of additional axes of variability subordinate to the principal functional spectra,nor that these may be certainly key to fitness and survival in specific cases.The CSR theory could be seen as a general adaptive strategy,since it is consistent with economics and size tradeoffs evident throughout the tree of life,hinting at overarching natural selection filters operating universally within the biosphere.