生态学的再思考: 历史、理论和实践

doi:10.17521/cjpe.2025.0220

植物生态学报 ›› 2026, Vol. 50 ›› Issue (1): 1-23.DOI: 10.17521/cjpe.2025.0220

• 侯学煜评述 • 下一篇

生态学的再思考: 历史、理论和实践

方精云¹^,^*(), 石岳², 朱江玲¹, 沈海花², 王志恒¹

¹ 北京大学城市与环境学院, 北京大学生态研究中心, 植被结构功能与建造全国重点实验室, 北京 100871
² 中国科学院植物研究所植被与环境变化重点实验室, 北京 100093

收稿日期:2025-06-12 接受日期:2025-10-17 出版日期:2026-01-20 发布日期:2026-02-13
通讯作者: *方精云, E-mail: jyfang@urban.pku.edu.cn
作者简介:方精云, 北京大学教授、植被结构功能与建造全国重点实验室主任。中国科学院院士、欧洲科学院外籍院士、第三世界科学院院士。主要从事全球变化生态学、植被生态、生态遥感、草牧业等方面的研究。发表论文500余篇, 论文总引用8万余次。曾获国家自然科学奖二等奖、长江学者成就奖以及美国生态学会惠特克杰出生态学家奖等奖项
石岳, 中国科学院植物研究所助理研究员。主要从事生态系统碳收支、生态草牧业理论等领域的研究。围绕中国陆地生态系统碳汇、中国天然草地和主要人工牧草品质分布格局及影响因素等发表中英文论文20余篇
朱江玲, 北京大学高级工程师, 植被结构功能与建造全国重点实验室副主任, 主要从事生物多样性、全球变化等方面的研究。发表论文50余篇, 曾获北京市自然科学一等奖、中国出版政府奖提名奖等奖项
沈海花, 中国科学院植物研究所研究员、博士生导师。主要从事植物生态学、全球变化生态学以及草地—牲畜耦合系统等方面的研究。在国内外主流学术期刊发表多篇研究论文
王志恒, 北京大学城市与环境学院教授, 国家杰出青年科学基金获得者。主要从事生物多样性与保护生态学研究。出版专著2部, 发表论文160余篇, 论著总被引约1.6万次
基金资助:
国家自然科学基金(32588202)

Rethinking about ecology: development, theories, and applications

FANG Jing-Yun¹^,^*(), SHI Yue², ZHU Jiang-Ling¹, SHEN Hai-Hua², WANG Zhi-Heng¹

¹ Institute of Ecology, College of Urban and Environmental Sciences, and State Key Laboratory for Vegetation Structure, Function and Construction (VegLab), Peking University, Beijing 100871, China
² Key Laboratory of Vegetation and Environment Change, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China

Received:2025-06-12 Accepted:2025-10-17 Online:2026-01-20 Published:2026-02-13
Supported by:
National Natural Science Foundation of China(32588202)

摘要/Abstract

摘要：

生态学是研究宏观生命系统内在规律及变化的科学, 是生态文明建设的科学基础。为适应学科发展的需要, 我国将生态学从生物学中独立出来, 作为独立的一级学科。为此, 亟须对生态学的学科体系、科学内涵、研究范畴及应用拓展进行梳理和重塑。该文对生态学的定义是: 研究宏观生命系统的结构、功能及其动态的科学, 它为人类认识、保护和利用自然提供理论基础和解决方案。现代生态学的研究涉及从分子、器官、个体到生物圈等不同组织层次, 但核心是个体、种群、群落、生态系统和景观五个层次。生态学的主干分支包括植物生态学、动物生态学、微生物生态学、生态系统生态学、景观生态学、修复生态学和可持续生态学。经过近160年的发展, 生态学产生了一系列重要概念和重要理论, 对推动自然科学的发展和人类文明的进步产生了重要影响。现代生态学具有鲜明的特点: 一是研究内容和研究范畴不断向宏观和微观方向拓展; 二是分子生物学、信息科学等技术和方法得到广泛应用; 三是野外控制实验和观测网络已成为发展趋势, 区域性和全球尺度的观测平台相继建立; 四是应用生态学受到更多重视, 为人类社会解决区域性和全球性生态与环境问题提供更多支持。生态学是人类认识和理解生命世界的自然观, 也是保护和改造自然世界的实践论。生态学认识和改造生命世界的方法论可以概括为五点: 层次观(生命系统具有多样的层次结构)、整体论(从整体论视角认识不同层次的生命现象)、系统论(以系统的观点看待生命世界)、演变论(从动态和演化的视角理解生命系统)和实践观(为保护、利用和改造自然世界提供方案和技术)。生态学研究的具体方法包括: 野外调查与观测、室内和室外实验、模型模拟, 以及整合分析等。虽然生态学及其各主干方向具有完整的理论体系, 但缺乏技术体系, 因此创建和发展核心技术对学科的持续健康发展至关重要。

关键词: 生态学定义, 生态学原理, 生态学方法, 生态学理论, 生态学层次, 生态学分支, 生态学趋势, 生态文明

Abstract:

Ecology explores the fundamental principles and dynamics of macro living-systems and provides the scientific foundation of ecological civilization. China has separated the ecology as an independent subject from the biology subject to better promote its development. This transition calls for a broader conceptual framework for the subject of ecology. In this article, we define “ecology” as “the science that studies the structures, functions and dynamics of macro living-systems”, which provides a theoretical guidance and practical solutions for maintaining sustainable biosphere. Current ecology encompasses multiple living-system levels from molecules to the biosphere, with its core focus on five key levels: individual, population, community, ecosystem, and landscape. The sub-subject system of ecology comprises seven core disciplines: Plant Ecology, Animal Ecology, Microbial Ecology, Ecosystem Ecology, Landscape Ecology, Restoration Ecology, and Sustainable Ecology. Over nearly 160 years, ecology has generated seminal concepts and landmark theories that have profoundly influenced natural science advancement and human civilization. Current ecology is characterized with four distinctive features: (1) expansion of research scope to both macro- and micro-scales; (2) broad adoption of methodologies from other fields such as molecular biology or information science; (3) increased attention to field-based experiments and observational networks, with platforms now established at regional and global scales; and (4) enhanced emphasis on applied ecology to address ecological challenges of human society. Serving as both a natural philosophy for comprehending the living world and a praxeology for conserving and utilizing nature, ecology can be framed through five core perspectives: (1) hierarchical perspective (recognizing the multiple structural levels of living systems), (2) holistic perspective (approaching ecological phenomena from an integrative viewpoint), (3) systematic perspective (viewing the living world as interconnected networks), (4) evolutionary perspective (understanding life systems as dynamic and evolving), and (5) practical perspective (developing solutions for sustainable stewardship of nature). Methodologically, ecological research relies on four principal approaches: field investigations, laboratory and in-situ controlled experiments, model simulations, and meta-analyses. Although ecology and its branches possess robust theoretical frameworks, they lack their own technological systems. Consequently, the development of core ecological technologies is essential to promote continued vitality and progression of the ecology discipline.

Key words: definition of ecology, ecological principles, ecological methodology, ecological theories, ecological hierarchy, branches of ecology, trends in ecology, ecological civilization

方精云, 石岳, 朱江玲, 沈海花, 王志恒. 生态学的再思考: 历史、理论和实践. 植物生态学报, 2026, 50(1): 1-23. DOI: 10.17521/cjpe.2025.0220

FANG Jing-Yun, SHI Yue, ZHU Jiang-Ling, SHEN Hai-Hua, WANG Zhi-Heng. Rethinking about ecology: development, theories, and applications. Chinese Journal of Plant Ecology, 2026, 50(1): 1-23. DOI: 10.17521/cjpe.2025.0220

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https://www.plant-ecology.com/CN/Y2026/V50/I1/1

图/表 8

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序号 No.	概念 Concept	定义或简介 Definition or description	提出者及年代 Proposer(s) and year(s)
1	种群 Population	在一定时间内占据一定空间的同种生物的所有个体 All individuals of the same species occupying a certain space within a certain time period	乔治•居维叶(法国) (1812) George Cuvier (France) (1812)
2	群落 Community	一定时间和一定空间上由不同种群构成的集合体 An assemblage of different populations within a certain space and time	卡尔•莫比乌斯(德国) (1877) Karl Möbius (Germany) (1877)
3	生态系统 Ecosystem	生物群落与非生物环境通过能量流动、物质循环和信息传递形成的动态功能整体 A dynamic functional whole formed by biological communities and abiotic environment through energy flow, material cycling, and information transfer	亚瑟•坦斯利(英国) (1935) Arthur Tansley (UK) (1935)
4	景观 Landscape	由相互作用的、具有不同生态特性的空间单元构成的空间镶嵌体 A spatial mosaic composed of interacting spatial units with different ecological characteristics	亚历山大•洪堡(德国)拓展这一概念(1845), 卡尔•特罗尔(德国)创造“景观生态学”一词(1939) Concept expanded by Alexander von Humboldt (Germany) (1845); term “landscape ecology” coined by Carl Troll (Germany) (1939)
5	生物圈 Biosphere	地球上生物及其生存环境的总称, 或地球生态系统的总称, 由大气圈、水圈和土壤岩石圈及生活其中的生物共同组成 The sum of all life on Earth and their living environments, or the global sum of all ecosystems on Earth, consisting of the atmosphere, hydrosphere, lithosphere, and organisms living within	爱德华•苏斯(奥地利)首次提出这一术语(1875), 弗拉基米尔•维尔纳德斯基(苏联)给出严格科学定义(1920) Term first proposed by Eduard Suess (Austria) (1875); scientifically defined by Vladimir Vernadsky (Soviet Union) (1920)
6	竞争 Competition	生物个体在种内或种间相互争夺某类资源的现象, 是达尔文进化论核心机制 The phenomenon of individuals competing within or between species for certain resources; a core mechanism of Darwinian evolutionary theory	查尔斯•达尔文(英国) (1859) Charles Darwin (UK) (1859)
7	自然选择 Natural selection	适应环境的个体更易生存并传递基因, 驱动物种进化 Individuals better adapted to their environment are more likely to survive and pass on genes, driving species evolution	查尔斯•达尔文(英国) (1859) Charles Darwin (UK) (1859)
8	协同进化 Coevolution	两个或多个物种通过相互作用驱动彼此进化(如传粉昆虫与开花植物) The process by which two or more species reciprocally influence each other’s evolution (e.g., pollinating insects and flowering plants)	保罗•埃利希(美国)和彼得•雷文(美国)共同提出(1964) Proposed jointly by Paul Ehrlich (USA) and Peter Raven (USA) (1964)
9	生态位 Ecological niche	物种在生态系统中, 在时间、空间上所占据的位置及与其他种群之间相互关系的总和 The total sum of a species’ position in space and time within an ecosystem and its interactions with other populations	约瑟夫•格林尼尔(美国) (1913)、查尔斯•埃尔顿(英国) (1927)分别独立阐述这一概念 Proposed independently by Joseph Grinnell (USA) (1913) and Charles Elton (UK) (1927)
10	演替 Succession	群落的物种组成和结构随时间推移而发生演化的过程 The process by which species composition and structure of a community change over time	让-安德烈•德吕克(英国)首次使用这一术语(1806), 弗雷德里克•克莱门茨(美国)发展这一概念(1916) Term first used by Jean-André Deluc (UK) (1806); developed by Frederic Clements (USA) (1916)
11	关键种	在群落中起重要作用, 且重要程度与其数量或生物量不成比例的物种 Species that play a critical role in a community whose importance is disproportionately large compared to their abundance or biomass	罗伯特•佩因(美国) (1969) Robert Paine (USA) (1969)
12	食物链/食物网 Food chain/food web	物种间通过取食与被取食关系形成的能量传递链/网络 Chain/networks of energy transfer formed by feeding and being fed upon relationships among species	查尔斯•埃尔顿(英国) (1927) Charles Elton (UK) (1927)
13	营养级 Trophic level	有机体在食物链中所处的层级 The level occupied by an organism within a food chain	雷蒙德•林德曼(美国) (1942) Raymond Lindeman (USA) (1942)
14	生物地球化学循环 Biogeochemical cycle	碳、氮、磷等元素在生物与非生物环境间的循环(如碳循环、氮循环) Cycles of elements such as carbon, nitrogen, and phosphorus between biotic and abiotic environments (e.g., carbon cycle, nitrogen cycle)	弗拉基米尔•维尔纳德斯基(苏联) (1926) Vladimir Vernadsky (Soviet Union) (1926)
15	生物多样性 Biodiversity	一定时空范围内所有生物及其变异的总和, 包括基因、物种和生态系统多样性 The total variety of life and its variation within a given time and space, including genetic, species, and ecosystem diversity	爱德华•威尔逊(美国)推广这一概念(1988) The concept popularized by Edward Wilson (USA) (1988)

序号 No.	概念 Concept	定义或简介 Definition or description	提出者及年代 Proposer(s) and year(s)
1	种群 Population	在一定时间内占据一定空间的同种生物的所有个体 All individuals of the same species occupying a certain space within a certain time period	乔治•居维叶(法国) (1812) George Cuvier (France) (1812)
2	群落 Community	一定时间和一定空间上由不同种群构成的集合体 An assemblage of different populations within a certain space and time	卡尔•莫比乌斯(德国) (1877) Karl Möbius (Germany) (1877)
3	生态系统 Ecosystem	生物群落与非生物环境通过能量流动、物质循环和信息传递形成的动态功能整体 A dynamic functional whole formed by biological communities and abiotic environment through energy flow, material cycling, and information transfer	亚瑟•坦斯利(英国) (1935) Arthur Tansley (UK) (1935)
4	景观 Landscape	由相互作用的、具有不同生态特性的空间单元构成的空间镶嵌体 A spatial mosaic composed of interacting spatial units with different ecological characteristics	亚历山大•洪堡(德国)拓展这一概念(1845), 卡尔•特罗尔(德国)创造“景观生态学”一词(1939) Concept expanded by Alexander von Humboldt (Germany) (1845); term “landscape ecology” coined by Carl Troll (Germany) (1939)
5	生物圈 Biosphere	地球上生物及其生存环境的总称, 或地球生态系统的总称, 由大气圈、水圈和土壤岩石圈及生活其中的生物共同组成 The sum of all life on Earth and their living environments, or the global sum of all ecosystems on Earth, consisting of the atmosphere, hydrosphere, lithosphere, and organisms living within	爱德华•苏斯(奥地利)首次提出这一术语(1875), 弗拉基米尔•维尔纳德斯基(苏联)给出严格科学定义(1920) Term first proposed by Eduard Suess (Austria) (1875); scientifically defined by Vladimir Vernadsky (Soviet Union) (1920)
6	竞争 Competition	生物个体在种内或种间相互争夺某类资源的现象, 是达尔文进化论核心机制 The phenomenon of individuals competing within or between species for certain resources; a core mechanism of Darwinian evolutionary theory	查尔斯•达尔文(英国) (1859) Charles Darwin (UK) (1859)
7	自然选择 Natural selection	适应环境的个体更易生存并传递基因, 驱动物种进化 Individuals better adapted to their environment are more likely to survive and pass on genes, driving species evolution	查尔斯•达尔文(英国) (1859) Charles Darwin (UK) (1859)
8	协同进化 Coevolution	两个或多个物种通过相互作用驱动彼此进化(如传粉昆虫与开花植物) The process by which two or more species reciprocally influence each other’s evolution (e.g., pollinating insects and flowering plants)	保罗•埃利希(美国)和彼得•雷文(美国)共同提出(1964) Proposed jointly by Paul Ehrlich (USA) and Peter Raven (USA) (1964)
9	生态位 Ecological niche	物种在生态系统中, 在时间、空间上所占据的位置及与其他种群之间相互关系的总和 The total sum of a species’ position in space and time within an ecosystem and its interactions with other populations	约瑟夫•格林尼尔(美国) (1913)、查尔斯•埃尔顿(英国) (1927)分别独立阐述这一概念 Proposed independently by Joseph Grinnell (USA) (1913) and Charles Elton (UK) (1927)
10	演替 Succession	群落的物种组成和结构随时间推移而发生演化的过程 The process by which species composition and structure of a community change over time	让-安德烈•德吕克(英国)首次使用这一术语(1806), 弗雷德里克•克莱门茨(美国)发展这一概念(1916) Term first used by Jean-André Deluc (UK) (1806); developed by Frederic Clements (USA) (1916)
11	关键种	在群落中起重要作用, 且重要程度与其数量或生物量不成比例的物种 Species that play a critical role in a community whose importance is disproportionately large compared to their abundance or biomass	罗伯特•佩因(美国) (1969) Robert Paine (USA) (1969)
12	食物链/食物网 Food chain/food web	物种间通过取食与被取食关系形成的能量传递链/网络 Chain/networks of energy transfer formed by feeding and being fed upon relationships among species	查尔斯•埃尔顿(英国) (1927) Charles Elton (UK) (1927)
13	营养级 Trophic level	有机体在食物链中所处的层级 The level occupied by an organism within a food chain	雷蒙德•林德曼(美国) (1942) Raymond Lindeman (USA) (1942)
14	生物地球化学循环 Biogeochemical cycle	碳、氮、磷等元素在生物与非生物环境间的循环(如碳循环、氮循环) Cycles of elements such as carbon, nitrogen, and phosphorus between biotic and abiotic environments (e.g., carbon cycle, nitrogen cycle)	弗拉基米尔•维尔纳德斯基(苏联) (1926) Vladimir Vernadsky (Soviet Union) (1926)
15	生物多样性 Biodiversity	一定时空范围内所有生物及其变异的总和, 包括基因、物种和生态系统多样性 The total variety of life and its variation within a given time and space, including genetic, species, and ecosystem diversity	爱德华•威尔逊(美国)推广这一概念(1988) The concept popularized by Edward Wilson (USA) (1988)

序号 No.	理论 Theory	核心内容 Core content		科学意义及影响 Scientific significance	提出者及年代 Proposer(s) and year(s)
1	贝格曼法则与阿伦法则 Bergmann’s Rule and Allen’s Rule	贝格曼法则指同种或近缘种恒温动物, 在低温地区有更大体型的现象。阿伦法则是贝格曼法则的补充, 指同种及近缘种之间, 生活在寒冷地区的个体呈现浑圆型体型(即器官突出的部分较短) Bergmann’s rule states that within the same or closely related species of endotherms, individuals in colder regions tend to have larger body sizes. Allen’s rule complements this, indicating that individuals in colder areas have more compact body shapes (i.e., shorter extremities)		揭示了生物对环境的适应进化特征 Revealed evolutionary adaptations of organisms to their environments	卡尔•贝格曼(德国) (1847)、乔伊•阿伦(美国) (1877)分别提出 Proposed respectively by Carl Bergmann (Germany) (1847) and Joel Allen (USA) (1877)
2	李比希最小因子定律与谢尔福德耐受性定律 Liebig’s Law of the Minimum and Shelford’s Law of Tolerance	前者指有机体的生长发育过程由其最稀缺必需因子所决定; 后者拓展了前者, 指有机体的存活受限于环境因子的临界范围 Liebig’s law states that an organism’s growth is limited by the scarcest essential factor; the law of tolerance extends this by stating that an organism’s survival is limited by the critical ranges of environmental factors		量化物种分布与环境适应的关系 Quantified the relationship between species distribution and environmental adaptation	尤斯图斯•李比希(德国) (1855)、维克托•谢尔福德(美国) (1913)分别提出 Proposed respectively by Justus Liebig (Germany) (1855) and Victor Shelford (USA) (1913)
3	自然选择理论 Theory of Natural Selection	适应环境的个体更易生存并传递基因, 驱动物种进化 Individuals better adapted to the environment are more likely to survive and pass on genes, driving species evolution		奠定生态学与进化生物学的共同根基 Laid the foundational basis for both ecology and evolutionary biology	查尔斯•达尔文(英国) (1859) Charles Darwin (UK) (1859)
4	r/K选择理论 r/K Selection Theory	生物演化出两种资源分配策略: r策略适应不稳定环境, K策略适应稳定环境 Organisms evolve two resource allocation strategies: r-strategy for unstable environments and K-strategy for stable ones		链接种群生态学与进化理论 Linked population ecology with evolutionary theory	罗伯特•麦克阿瑟(美国)和爱德华•威尔逊(美国)共同提出(1967) Proposed jointly by Robert MacArthur (USA) and Edward Wilson (USA) (1967)
5	洛特卡-沃尔泰拉模型 Lotka-Volterra Model	描述捕食者与猎物之间相互作用的方程, 分别用Logistic方程刻画两者种群数量的变化 Equations describing predator-prey interactions, using Logistic equations to depict population changes of both species		奠定了种间竞争关系研究的理论基础 Established the theoretical basis for studying interspecific competition	阿尔弗雷德•洛特卡(美国) (1925)、维托里奥•沃尔泰拉(意大利) (1926)分别独立提出 Proposed independently by Alfred Lotka (USA) (1925) and Vito Volterra (Italy) (1926)
6	竞争排除原理 Competitive Exclusion Principle	生态位完全重叠的物种无法长期共存, 必有一方被淘汰 Species with completely overlapping ecological niches cannot coexist long-term; one will be excluded		解释物种共存的生态位分化机制 Explained the mechanism of niche differentiation enabling species coexistence	格奥尔基•高斯(苏联) (1934) Georgii Gause (Soviet Union) (1934)
7	生态位理论 Ecological Niche Theory	物种的生存由其资源需求、环境耐受范围及种间关系共同定义 A species’ existing is determined by its resource requirements, environmental tolerance, and interspecific relationships		构建群落生态学理论框架 Constructed the theoretical framework for community ecology	伊夫林•哈钦森(美国)发展(1957) Developed by G. Evelyn Hutchinson (USA) (1957)
8	-3/2自然稀疏法则 The -3/2 Law of Self-thinning	由竞争导致种群自然稀疏的过程, 可用幂指数为-3/2的幂函数表示 The natural thinning of populations due to competition follows a power law with an exponent of -3/2		为达尔文进化理论提供数量基础 Provided a quantitative basis for Darwinian evolutionary theory	依田恭二(日本)和吉良龙夫(日本)共同提出(1963) Proposed jointly by Yoda Kyoji (Japan) and Kira Tatsuo (Japan) (1963)
9	竞争-密度理论 Competition- density Theories	由种群密度不同引起的密度-生长关系变化的一系列理论和法则, 如最终产量恒定法则和竞争-密度倒数式等 A series of theories and laws explaining changes in density-growth relationships due to population density, such as the constant final yield law and inverse density-competition law		量化了达尔文生存竞争的理论和观点 Quantified Darwinian theory on survival competition	吉良龙夫(日本)、篠崎吉郎(日本)等共同提出(1953, 1956) Proposed jointly by Kira Tatsuo (Japan), Shinozaki Kichiro (Japan), etc. (1953, 1956)
10	相关生长原理 Principles of Allometric Growth	描述生物个体量与部分量之间的数量关系, 又称异速生长原理 Describes quantitative relationships between organismal size and parts, also known as the principles of allometric scaling		为整体量与部分量之间关系的研究提供方法 Provided methods to study the relationship between whole and parts	朱利安•赫胥黎(英国)提出, 并创造“allometry”一词(1936) Julian Huxley (UK), who coined the term “allometry” (1936)
11	岛屿生物地理学理论 Island Biogeography Theory	岛屿物种多样性由迁入率与灭绝率的动态平衡决定 Species diversity on islands is determined by a dynamic balance between immigration and extinction rates	指导保护生物学与生境碎片化研究 Guided conservation biology and habitat fragmentation researches		罗伯特•麦克阿瑟(美国)和爱德华•威尔逊(美国)共同提出(1967) Proposed jointly by Robert MacArthur (USA) and Edward Wilson (USA) (1967)
12	环境梯度理论 Theory of Environmental Gradients	沿环境梯度的连续性变化, 群落及其环境共同构成的生态系统, 在结构与功能上也呈现相一致的格局 Along continuous environmental gradients, communities and the ecosystems formed by these communities and their environments exhibit structural and functional patterns corresponding to those gradients	为生态系统结构、功能及其机制研究提供理论构架 Provided a theoretical framework for studying ecosystem structure, function, and mechanisms		罗伯特•惠特克(美国) (1948) Robert Whittaker (USA) (1948)
13	群落演替理论 Community Succession Theory	认识群落演替性质、解释物种更替过程以及顶极群落决定因素的一系列学说和理论 A series of theories explaining the nature of community succession, species replacement processes, and determinants of climax communities	指导生态恢复与灾害管理 Guided ecological restoration and disaster management		弗雷德雷克•克莱门茨(美国)早期提出(1905, 1916), 亨利•格里森(美国) (1917)、弗兰克•埃格勒(美国) (1954)、罗伯特•惠特克(美国) (1960)等人发展 Proposed early by Frederic Clements (USA) (1905, 1916), and further developed by Henry Gleason (USA) (1917), Frank Egler (USA) (1954), Robert Whittaker (USA) (1960), and others
14	中性理论 Neutral Theory	群落中物种多度分布可由随机过程(如扩散、出生死亡)解释, 无需生态位分化 Species abundance distributions in communities can be explained by stochastic processes (e.g., dispersal, birth, death) without niche differentiation	挑战传统生态位理论, 推动群落生态学范式革新 Challenged traditional niche theory and promoted paradigm shifts in community ecology		斯蒂芬•哈贝尔(美国) (2001) Stephen Hubbell (USA) (2001)
15	生态系统理论 Ecosystem Theory	生物群落与非生物环境通过能量流动、物质循环和信息传递形成的功能整体 Biological communities and abiotic environments form a functional whole through energy flow, material cycling, and information transfer	确立生态学系统研究的范式 Established the paradigm for ecosystem-level ecological research		亚瑟•坦斯利(英国)提出“生态系统”概念(1935); 多人发展了生态系统理论, 如雷蒙德•林德曼(美国)提出营养级关系(1942), 查尔斯•埃尔顿(英国)提出食物链/食物网概念(1927), 奥德姆兄弟(美国)提出“能流模型” (1956, 1968)、“系统生态学” (1983)等 Arthur Tansley (UK) proposed the concept of “ecosystem” (1935). The theory of ecosystems was further developed by many scholars, including Raymond Lindeman (USA), who introduced the concept of trophic levels (1942); Charles Elton (UK), who proposed the concepts of food chains/food webs (1927); and Eugene Odum and Howard Odum (USA), who developed the “energy flow model” (1956, 1968) and “systems ecology” (1983)

序号 No.	理论 Theory	核心内容 Core content		科学意义及影响 Scientific significance	提出者及年代 Proposer(s) and year(s)
1	贝格曼法则与阿伦法则 Bergmann’s Rule and Allen’s Rule	贝格曼法则指同种或近缘种恒温动物, 在低温地区有更大体型的现象。阿伦法则是贝格曼法则的补充, 指同种及近缘种之间, 生活在寒冷地区的个体呈现浑圆型体型(即器官突出的部分较短) Bergmann’s rule states that within the same or closely related species of endotherms, individuals in colder regions tend to have larger body sizes. Allen’s rule complements this, indicating that individuals in colder areas have more compact body shapes (i.e., shorter extremities)		揭示了生物对环境的适应进化特征 Revealed evolutionary adaptations of organisms to their environments	卡尔•贝格曼(德国) (1847)、乔伊•阿伦(美国) (1877)分别提出 Proposed respectively by Carl Bergmann (Germany) (1847) and Joel Allen (USA) (1877)
2	李比希最小因子定律与谢尔福德耐受性定律 Liebig’s Law of the Minimum and Shelford’s Law of Tolerance	前者指有机体的生长发育过程由其最稀缺必需因子所决定; 后者拓展了前者, 指有机体的存活受限于环境因子的临界范围 Liebig’s law states that an organism’s growth is limited by the scarcest essential factor; the law of tolerance extends this by stating that an organism’s survival is limited by the critical ranges of environmental factors		量化物种分布与环境适应的关系 Quantified the relationship between species distribution and environmental adaptation	尤斯图斯•李比希(德国) (1855)、维克托•谢尔福德(美国) (1913)分别提出 Proposed respectively by Justus Liebig (Germany) (1855) and Victor Shelford (USA) (1913)
3	自然选择理论 Theory of Natural Selection	适应环境的个体更易生存并传递基因, 驱动物种进化 Individuals better adapted to the environment are more likely to survive and pass on genes, driving species evolution		奠定生态学与进化生物学的共同根基 Laid the foundational basis for both ecology and evolutionary biology	查尔斯•达尔文(英国) (1859) Charles Darwin (UK) (1859)
4	r/K选择理论 r/K Selection Theory	生物演化出两种资源分配策略: r策略适应不稳定环境, K策略适应稳定环境 Organisms evolve two resource allocation strategies: r-strategy for unstable environments and K-strategy for stable ones		链接种群生态学与进化理论 Linked population ecology with evolutionary theory	罗伯特•麦克阿瑟(美国)和爱德华•威尔逊(美国)共同提出(1967) Proposed jointly by Robert MacArthur (USA) and Edward Wilson (USA) (1967)
5	洛特卡-沃尔泰拉模型 Lotka-Volterra Model	描述捕食者与猎物之间相互作用的方程, 分别用Logistic方程刻画两者种群数量的变化 Equations describing predator-prey interactions, using Logistic equations to depict population changes of both species		奠定了种间竞争关系研究的理论基础 Established the theoretical basis for studying interspecific competition	阿尔弗雷德•洛特卡(美国) (1925)、维托里奥•沃尔泰拉(意大利) (1926)分别独立提出 Proposed independently by Alfred Lotka (USA) (1925) and Vito Volterra (Italy) (1926)
6	竞争排除原理 Competitive Exclusion Principle	生态位完全重叠的物种无法长期共存, 必有一方被淘汰 Species with completely overlapping ecological niches cannot coexist long-term; one will be excluded		解释物种共存的生态位分化机制 Explained the mechanism of niche differentiation enabling species coexistence	格奥尔基•高斯(苏联) (1934) Georgii Gause (Soviet Union) (1934)
7	生态位理论 Ecological Niche Theory	物种的生存由其资源需求、环境耐受范围及种间关系共同定义 A species’ existing is determined by its resource requirements, environmental tolerance, and interspecific relationships		构建群落生态学理论框架 Constructed the theoretical framework for community ecology	伊夫林•哈钦森(美国)发展(1957) Developed by G. Evelyn Hutchinson (USA) (1957)
8	-3/2自然稀疏法则 The -3/2 Law of Self-thinning	由竞争导致种群自然稀疏的过程, 可用幂指数为-3/2的幂函数表示 The natural thinning of populations due to competition follows a power law with an exponent of -3/2		为达尔文进化理论提供数量基础 Provided a quantitative basis for Darwinian evolutionary theory	依田恭二(日本)和吉良龙夫(日本)共同提出(1963) Proposed jointly by Yoda Kyoji (Japan) and Kira Tatsuo (Japan) (1963)
9	竞争-密度理论 Competition- density Theories	由种群密度不同引起的密度-生长关系变化的一系列理论和法则, 如最终产量恒定法则和竞争-密度倒数式等 A series of theories and laws explaining changes in density-growth relationships due to population density, such as the constant final yield law and inverse density-competition law		量化了达尔文生存竞争的理论和观点 Quantified Darwinian theory on survival competition	吉良龙夫(日本)、篠崎吉郎(日本)等共同提出(1953, 1956) Proposed jointly by Kira Tatsuo (Japan), Shinozaki Kichiro (Japan), etc. (1953, 1956)
10	相关生长原理 Principles of Allometric Growth	描述生物个体量与部分量之间的数量关系, 又称异速生长原理 Describes quantitative relationships between organismal size and parts, also known as the principles of allometric scaling		为整体量与部分量之间关系的研究提供方法 Provided methods to study the relationship between whole and parts	朱利安•赫胥黎(英国)提出, 并创造“allometry”一词(1936) Julian Huxley (UK), who coined the term “allometry” (1936)
11	岛屿生物地理学理论 Island Biogeography Theory	岛屿物种多样性由迁入率与灭绝率的动态平衡决定 Species diversity on islands is determined by a dynamic balance between immigration and extinction rates	指导保护生物学与生境碎片化研究 Guided conservation biology and habitat fragmentation researches		罗伯特•麦克阿瑟(美国)和爱德华•威尔逊(美国)共同提出(1967) Proposed jointly by Robert MacArthur (USA) and Edward Wilson (USA) (1967)
12	环境梯度理论 Theory of Environmental Gradients	沿环境梯度的连续性变化, 群落及其环境共同构成的生态系统, 在结构与功能上也呈现相一致的格局 Along continuous environmental gradients, communities and the ecosystems formed by these communities and their environments exhibit structural and functional patterns corresponding to those gradients	为生态系统结构、功能及其机制研究提供理论构架 Provided a theoretical framework for studying ecosystem structure, function, and mechanisms		罗伯特•惠特克(美国) (1948) Robert Whittaker (USA) (1948)
13	群落演替理论 Community Succession Theory	认识群落演替性质、解释物种更替过程以及顶极群落决定因素的一系列学说和理论 A series of theories explaining the nature of community succession, species replacement processes, and determinants of climax communities	指导生态恢复与灾害管理 Guided ecological restoration and disaster management		弗雷德雷克•克莱门茨(美国)早期提出(1905, 1916), 亨利•格里森(美国) (1917)、弗兰克•埃格勒(美国) (1954)、罗伯特•惠特克(美国) (1960)等人发展 Proposed early by Frederic Clements (USA) (1905, 1916), and further developed by Henry Gleason (USA) (1917), Frank Egler (USA) (1954), Robert Whittaker (USA) (1960), and others
14	中性理论 Neutral Theory	群落中物种多度分布可由随机过程(如扩散、出生死亡)解释, 无需生态位分化 Species abundance distributions in communities can be explained by stochastic processes (e.g., dispersal, birth, death) without niche differentiation	挑战传统生态位理论, 推动群落生态学范式革新 Challenged traditional niche theory and promoted paradigm shifts in community ecology		斯蒂芬•哈贝尔(美国) (2001) Stephen Hubbell (USA) (2001)
15	生态系统理论 Ecosystem Theory	生物群落与非生物环境通过能量流动、物质循环和信息传递形成的功能整体 Biological communities and abiotic environments form a functional whole through energy flow, material cycling, and information transfer	确立生态学系统研究的范式 Established the paradigm for ecosystem-level ecological research		亚瑟•坦斯利(英国)提出“生态系统”概念(1935); 多人发展了生态系统理论, 如雷蒙德•林德曼(美国)提出营养级关系(1942), 查尔斯•埃尔顿(英国)提出食物链/食物网概念(1927), 奥德姆兄弟(美国)提出“能流模型” (1956, 1968)、“系统生态学” (1983)等 Arthur Tansley (UK) proposed the concept of “ecosystem” (1935). The theory of ecosystems was further developed by many scholars, including Raymond Lindeman (USA), who introduced the concept of trophic levels (1942); Charles Elton (UK), who proposed the concepts of food chains/food webs (1927); and Eugene Odum and Howard Odum (USA), who developed the “energy flow model” (1956, 1968) and “systems ecology” (1983)

序号 No.	姓名及生卒年 Name and birth-death years	主要贡献 Major contribution
1	亚历山大•洪堡(1769-1859) Alexander von Humboldt (1769-1859)	提出“自然整体性”概念, 开创生物地理学 Proposed the concept of “the unity of nature” and pioneered biogeography
2	查尔斯•达尔文(1809-1882) Charles Darwin (1809-1882)	创建自然选择理论, 推动种内和种间竞争研究 Proposed the theory of natural selection and advanced research on intra- and interspecific competition
3	恩斯特•海克尔(1834-1919) Ernst Haeckel (1834-1919)	首次定义“生态学”, 确立学科独立性 First defined “ecology” and established it as an independent discipline
4	欧根纽斯•瓦尔明(1841-1924) Eugenius Warming (1841-1924)	提出植被分布的环境解释, 出版第一本生态学教科书, 推动生态学体系化 Provided environmental explanations for vegetation distribution, published the first ecology textbook, and promoted the systematization of ecology
5	弗拉基米尔•维尔纳德斯基(1863-1945) Vladimir Vernadsky (1863-1945)	提出“生物圈”概念的严格科学释义, 定义生物地球化学循环 Gave a rigorous scientific definition of the “biosphere” concept and defined biogeochemical cycles
6	亚瑟•坦斯利(1871-1955) Arthur Tansley (1871-1955)	提出“生态系统”概念, 整合生物与非生物组分纳入统一研究框架 Proposed the concept of the “ecosystem”, integrating biotic and abiotic components into a unified research framework
7	弗雷德里克•克莱门茨(1874-1945) Frederic Clements (1874-1945)	提出群落演替“单元顶极理论”, 主张群落为超级有机体 Proposed “the monoclimax theory” of community succession and advocated that communities function as superorganisms
8	查尔斯•埃尔顿(1900-1991) Charles Elton (1900-1991)	动物生态学先驱, 定义食物链、生态金字塔概念 Pioneer of animal ecology, defined the concepts of the food chain and the ecological pyramid
9	伊夫林•哈钦森(1903-1991) G. Evelyn Hutchinson (1903-1991)	现代生态学理论奠基人, 拓展生态位概念, 提出湖泊生态系统模型 Founder of modern ecological theory, expanded the concept of ecological niche, and proposed the lake ecosystem models
10	尤金•奥德姆(1913-2002) Eugene Odum (1913-2002)	出版《生态学基础》, 生态系统生态学的重要创立者 Published Fundamentals of Ecology, a key founder of ecosystem ecology
11	雷蒙德•林德曼(1915-1942) Raymond Lindeman (1915-1942)	提出能量流动“十分之一定律” Proposed the “ten percent law” of energy flows in ecosystems
12	吉良龙夫(1919-2011) Kira Tatsuo (1919-2011)	提出竞争-密度效应模型和-3/2自疏法则 Proposed the competition-density model and the -3/2 law of self-thinning
13	罗伯特•惠特克(1920-1980) Robert Whittaker (1920-1980)	发展环境梯度理论, 提出多样性概念和测度 Developed the theory of environmental gradients, proposed concepts and metrics for diversity
14	爱德华•威尔逊(1929-2021) Edward Wilson (1929-2021)	共同提出岛屿生物地理学理论, 生物多样性保护理论与“半地球计划”倡导者 Co-proposed the island biogeography theory, advocated biodiversity conservation and the Half-Earth Project
15	罗伯特•麦克阿瑟(1930-1972) Robert MacArthur (1930-1972)	共同提出岛屿生物地理学理论 Co-proposed the island biogeography theory

生态学的再思考: 历史、理论和实践

Rethinking about ecology: development, theories, and applications

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