生物间高阶相互作用研究进展

doi:10.17520/biods.2020217

摘要/Abstract

摘要：

生物间的相互作用是物种共存和生物多样性维持的关键。传统的物种共存研究主要关注配对物种之间的直接相互作用, 而忽略了更为复杂的间接相互作用。本文首先介绍了两种间接相互作用: 链式相互作用(本质上仍是两两物种之间的相互作用)和高阶相互作用。在此基础上, 我们回顾了高阶相互作用定义的演变历史(包括狭义的高阶相互作用和广义的高阶相互作用)及其检验方法, 并介绍了高阶相互作用在多营养级之间和同一营养级内的研究概况。目前, 生态学家主要对多营养级之间(如食物网)的高阶相互作用的特征、发生机制、作用途径及实验证据等方面进行了详尽的研究。近年来, 同一营养级内的高阶相互作用也开始受到关注, 因此我们进一步介绍了同一营养级内个体水平高阶相互作用的重要意义和度量方法。从个体水平上研究高阶相互作用, 既能统一狭义和广义高阶相互作用在定义上的争议, 又可以将个体间的差异(如个体大小、个体的空间分布等信息)考虑进来。最后, 本文对高阶相互作用一些可能的重要研究方向进行了展望: 在自然群落中(尤其同一营养级内)检验高阶相互作用的普遍性与相对重要性, 探讨高阶相互作用的发生机制以及如何将高阶相互作用整合到现有的理论体系中等。高阶相互作用的研究有助于我们全面深刻地理解物种共存和生物多样性的维持机制, 丰富和完善群落生态学的理论框架, 为人类世背景下的生物多样性保护和生态系统功能维持与提升提供基础。

关键词: 密度介导的间接相互作用, 性状介导的间接相互作用, 相互作用的调节, 个体水平高阶相互作用, 非线性密度制约, 营养级, 生态网络

Abstract

It is well known that interactions between organisms are the key to species coexistence and biodiversity maintenance. Traditional studies focused overwhelmingly on direct interactions between species pairs, ignoring the more complex indirect interactions. In this review, we first distinguished two types of indirect interactions, i.e. interaction chains and higher-order interactions (HOIs). Then we reviewed the definition of higher-order interactions including the hard-HOIs and soft-HOIs, and the studies of HOIs among multiple trophic levels and within a single trophic level. In the food-web literature (among multiple trophic levels), ecologists widely studied the properties, mechanisms, pathways and experimental evidence of HOIs. Recently, there is an increasing interest in HOIs within a single trophic level. Therefore, we further introduced the significance and quantification of individual-level HOIs within a single trophic level. Not only can individual-level HOIs reconcile the hard-HOIs and soft-HOIs, but also allow us to consider variatons between individuals (e.g. individual size and spatial distribution). Finally, we proposed some promising research directions of HOIs including but not limited to: testing the prevalence and relative importance of HOIs in natural communities, exploring the mechanisms of HOIs and integrating HOIs to existing theories of community ecology. Inclusion of HOIs will help us understand the mechanisms of species coexistence and biodiversity maintenance profoundly and comprehensively, enrich and refine the theoretical framework of community ecology, and lay the foundation for biodiversity conservation and management of ecosystems in the Anthropocene.

Key words: density-mediated indirect interactions, trait-mediated indirect interactions, interaction modifications, individual-level higher-order interactions, nonlinear density dependence, trophic levels, ecological network

李远智, 肖俊丽, 刘翰伦, 王酉石, 储诚进 (2020) 生物间高阶相互作用研究进展. 生物多样性, 28, 1333-1344. DOI: 10.17520/biods.2020217.

Yuanzhi Li, Junli Xiao, Hanlun Liu, Youshi Wang, Chengjin Chu (2020) Advances in higher-order interactions between organisms. Biodiversity Science, 28, 1333-1344. DOI: 10.17520/biods.2020217.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2020217

https://www.biodiversity-science.net/CN/Y2020/V28/I11/1333

图/表 4

图1 包含直接和间接相互作用的生态网络。灰色箭头为直接配对相互作用(箭头1-3), 黑色箭头为间接相互作用(箭头4-5)。在间接相互作用中, 箭头4表示高阶相互作用, 即物种k影响的是物种j和i之间的相互作用, 箭头5表示链式相互作用, 即物种k先影响物种j的密度进而影响物种i。可见, 物种k对物种i既存在直接相互作用(箭头1), 也存在间接相互作用(箭头4和5)。箭头表示作用方向, 为简单起见, 只绘出了单向作用。

Fig. 1 The ecological network including direct (arrows 1-3) and indirect interactions (arrows 4-5) between species. Arrow 4 indicates that species k may indirectly affect species i by modifying the per capita effect of species j on species i (higher-order interactions, HOIs). Arrow 5 indicates that species k may indirectly affect species i by changing population density of species j (interaction chains). Therefore, species k may have both direct (arrow 1) and indirect (arrows 4-5) effects on species i. For simplicity, we only display direct and indirect effects of species j and k on species i.

图2 生物间相互作用的类型和关系。同一方框内的不同术语为不同角度描述的同一类型的相互作用, 虚线部分为作者见解, 尚无相关文献明确说明。

Fig. 2 The types of biotic interactions. The different terms in the same box were used to describe the same type of interaction in different studies. The part in dashed line is our own opinion.

表1 检验模型是否包含狭义高阶相互作用的方法及部分常见模型检验结果。√表示模型中包含狭义高阶相互作用(模型不满足方法中等式), ×表示模型不含狭义高阶相互作用(模型满足方法中等式)。这些方法旨在将狭义高阶相互作用从广义高阶相互作用中区分出来, 其中(1)、(3)和(5)用于检验两物种或两物种以上系统是否包含狭义高阶相互作用, (2)和(4)分别是(1)和(3)用于将狭义高阶相互作用严格定义在三物种或三物种以上系统中时的情况, 因而方法(1)和(2), (3)和(4)在三个以上物种组成的系统中等效。Fi表示物种i的单位种群增长率是其自身及竞争者密度的函数, 这里给出几个常见模型的Fi函数表达式。如果函数Fi对Nj的偏导数?Fi/?Nj能表达成Nj的函数Gij(Nj) (方法1), 或是Ni和Nj的函数Gij(Ni, Nj) (方法2), 或是Nj和Fi自身的函数Gij(Fi, Nj) (方法3), 或是Ni和Nj以及Fi的函数Gij(Fi, Ni, Nj) (方法4), 则模型没有狭义高阶相互作用。方法(5)中, Qi表示函数Fi中所有参数的集合, jij表示除物种j外所有物种的密度均为0时函数Fi(0, …, Nj, …, 0)中的参数, Fi则是jij (j = 1, …, S)的并集。若Qi = Fi, 则模型没有狭义高阶相互作用。

Table 1 The methods of detecting whether a species interaction model contains hard higher-order interactions (hard-HOIs) and the outcomes of some well known models. √ and × indicate the model contains (the equation in a method is violated) and does not contain (the equation in a mothed is satisfied) hard-HOIs, respectively. Methods (1), (3) and (5) are used in the case of HOIs defined in systems of two or more species, and methods (2) and (4) are special cases of (1) and (3) where HOIs are strictedly defined in systems of three or more species. Fi indicates the per capita growth rate of species i as a function the densities of itself and its competitiors. If the partial derivative of Fi to Nj (?Fi/?Nj) can be expressed as only a function of Nj: Gij(Nj) (method 1), or a function of Ni and Nj: Gij(Ni, Nj) (method 2), or function of Nj and Fi: Gij(Fi, Nj) (method 3), or a function of Ni, Nj and Fi: Gij(Fi, Ni, Nj) (method 4), then the model Fi does not contain hard-HOIs according to methods 1-4, respectively. In method (5), Qi indicates the set of paramters in function Fi; jij indicates the set of parameters in function Fi(0, …, Nj, …, 0) when densities of all species are zero except species j; Fi is the union of jij (j = 1, …, S). If Qi = Fi, then the model Fi does not contain hard-HOIs.

图3 邻体对目标个体的直接(直线箭头)与高阶相互作用(曲线箭头)。参数$\alpha_{i_{m}j_{p}}$表示的是邻体jp对目标个体im的直接相互作用, 参数$\beta_{i_{m}j_{p},k_{q}}$表示的是邻体kq通过个体jp对目标个体im的高阶相互作用。森林群落研究中一般假定直接相互作用发生于邻体jp在目标个体im半径为R的邻域内(实直线箭头), 因而高阶相互作用发生于当邻体jp在目标个体im半径为R的邻域内且邻体的邻体kq在邻体jp的邻域内(实曲线箭头)。虚线箭头表示邻域外不需要考虑的直接与高阶相互作用。

Fig. 3 Direct (straight arrows) and higher-order interactions (curve arrows) of neighbouring trees on a focal tree. The parameter $\alpha_{i_{m}j_{p}}$ quantifies the direct effect of a neighbour (individual p of species j) on the focal tree (individual m of species i). The direct interaction occurs only when a neighbour (jp) is located within a maximum radius (R) of im (solid straight arrows). The parameter $\beta_{i_{m}j_{p},k_{q}}$ quantifies the higher-order effect of a neighbour (individual q of species k) on the focal tree through another neighbour (individual p of species j). Higher-order interaction occurs only jp is located within the maximum radius (R) of im and kq is located within the maximum radius (R) of jp (solid curve arrows). Dashed arrows indicate direct interactions and higher-order interactions that are not considered when a neighbour is located outside the maximum radius (R) of the focal tree or its neighbour(s).

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