生物多样性 ›› 2020, Vol. 28 ›› Issue (11): 1345-1361. DOI: 10.17520/biods.2020110
收稿日期:
2020-03-23
接受日期:
2020-05-27
出版日期:
2020-11-20
发布日期:
2020-06-18
通讯作者:
宋础良
作者简介:
* E-mail: clsong.ecology@gmail.comReceived:
2020-03-23
Accepted:
2020-05-27
Online:
2020-11-20
Published:
2020-06-18
Contact:
Chuliang Song
摘要:
群落内物种间相互作用的结构是高度组织化的。群落结构对多物种共存的影响机制是群落生态学的核心科学问题之一。目前生态学界在这一问题上存在多种不同的观点。一个可能的原因是, 由于环境因子的复杂性, 大部分研究忽略了环境因子对群落结构和物种共存的重要影响。在这一背景下, 近期发展起来的结构稳定性理论系统地联系了群落结构、环境因子和物种共存, 并在此基础上建立了一个和经验数据紧密结合的理论框架。本文首先简要回顾了当前关于群落结构研究的争鸣, 然后介绍了结构稳定性的理论框架和计算方法, 最后详细介绍了结构稳定性理论在不同生态群落和不同生态学问题中的应用。在全球气候变化的背景下, 结构稳定性理论提供了一种新的视角来理解群落层面的生物多样性维持机制。
宋础良 (2020) 结构稳定性: 概念、方法和应用. 生物多样性, 28, 1345-1361. DOI: 10.17520/biods.2020110.
Chuliang Song (2020) Structural stability: Concepts, methods, and applications. Biodiversity Science, 28, 1345-1361. DOI: 10.17520/biods.2020110.
图1 两种经典的群落结构。这里的矩阵展示了群落结构: 每一列代表不同植物, 而每一行代表了不同的授粉种(图A)或者草食动物(图B)。灰色的格子代表两个物种存在种间关系, 而白色的格子则代表不存在。图A展示了嵌套性结构。基于一个物种的种间关系的相对数量, 可以把物种分为泛化种(generalist)和特化种(specialist)。嵌套性结构的特征是认为泛化种会和泛化种以及特化种都相互作用, 而特化种几乎只和泛化种相互作用。嵌套性结构被认为是互利群落的一种普适结构。图B展示了模块化结构。模块化结构的特征是群落分成几个模块, 物种几乎只和同一模块的物种相互作用, 而很少和别的模块的物种相互作用。嵌套性结构被认为是食物网的一种普适结构。
Fig. 1 Two ecological community structures. Here we present an illustration of two hypothetical network structures. The community structure is represented as a matrix. The columns correspond to different plant species while the rows correspond to different pollinators (in Panel A) or herbivores (in Panel B). A species interaction is represented as a gray grid. Panel (A) illustrates the nested structure. The defining feature of a nested structure is that highly connected species interact with both highly connected and poorly connected species, while poorly connected species interact almost exclusively with highly connected species. The nested structure is widely conceived as a universal structure in mutualistic communities. Panel (B) illustrates the modular structure. The defining feature of the modular structure is that in which groups of species have many interactions among them, but few interactions with the rest of the species in the network. The modular structure is widely conceived as a universal structure in food webs.
图2 群落结构研究的争鸣。互利群落中关于嵌套性和物种多样性的研究中, 存在三种不同的派系。图A展示了第一种派系, 认为嵌套性是决定互利群落的物种多样性的原因。图B展示了第二种派系, 认为嵌套性并不是决定互利群落的物种多样性的原因。图C展示了第三种派系, 认为嵌套性并不是互利群落的普适结构, 因此嵌套性和物种多样性之间的关系无从谈起。
Fig. 2 The debates in the study of community structures. Three different schools of thought coexist in linking nestedness and species coexistence in mutualistic communities. The first school (Panel A) argues that nestedness is a key factor to support the biodiversity in mutualistic communities, the second school (Panel B) argues there is no causal relationship between the nested pattern and biodiversity, and the third school (Panel C) argues that nested patterns are not universal in observed mutualistic communities, thus makes the whole question a straw man.
图3 结构稳定性理论的基础数学框架。和经典的群落结构-物种共存理论相比, 结构稳定性理论引入了环境因子。环境因子同时会影响群落结构和物种共存。结构稳定性(SS)的计算需要两部分的信息: 共存域(CD)和环境域(ED)。共存域指的是可以使得物种共存的全部参数的空间, 图中用蓝色区域(图A)指代。共存域是由群落结构通过群落动力学而决定的。环境域指的是这一环境条件下所有可能的参数的空间, 图中用绿色区域(图B)指代。环境域是在给定环境的限制下所有可能的参数的空间。环境域是环境因子决定的。结构稳定性是共存域相对于环境域的大小。结构稳定性越大, 物种在这一环境条件下就有更大可能可以共存。
Fig. 3 The mathematical framework of the structural stability approach. To compute the structural stability (SS) of an ecological community, we need two pieces of information: the coexistence domain (CD) and the environment domain (ED). The coexistence domain (denoted in blue; region A) is the full range of parameters that are compatible with species coexistence. The coexistence domain is determined by the community structure via the ecological dynamics. The environment domain (denoted in green; region B) is the full range of parameters constrained by the given environmentally conditions. The environment domain is determined by the environmental factors. Structural stability is defined as the relative size of the coexistence domain comparing to the environment domain. The larger the structural stability is, the more likely species can coexist under the given environmental conditions.
图4 结构稳定性理论的物种共存框架。群落中有4个核心因素: 物种库、群落结构、环境因子和物种共存。这些核心因素之间都有着具体的生态过程(蓝色): 群落聚合、结构变化、群落动力学、环境梯度和环境胁迫。结构稳定性理论试图将这些生态过程整合在一个统一的框架下。对于图中的每一个生态过程, 结构稳定性理论都提出了新的假说和验证方法, 并在实验以及观测数据中得到了验证。
Fig. 4 The structural stability approach for understanding multispecies coexistence. The nodes denote the key elements in ecological dynamics: species pool, community structure, environmental factors, and species coexistence. The links represent the ecological processes (denoted in blue) that connect these elements: community assembly, structural change, ecological dynamics, environmental gradient, and environmental stress. The structural stability approach aims to integrate these ecological processes under a unified framework. For each process, the structural stability approach provides new theoretical predictions, which are validated by experimental and/or observational data.
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