整体大于部分之和: 生态自组织斑图及其涌现属性

doi:10.17520/biods.2020225

摘要/Abstract

摘要：

近30年来, 自组织理论已经发展成为解释生态系统呈现规则空间格局的有效理论。伴随着生态系统自发有序空间格局的生成, 自组织过程产生一系列的涌现属性, 这些特征对生态系统功能至关重要。在此, 我们将介绍这一正蓬勃发展的研究领域的主要理论进展。首先, 叙述了自组织这一概念的发展历程与定义, 详细阐述了自组织理论的两个经典理论框架: 图灵原理与相分离原理。然后, 根据几个典型的生态自组织研究案例, 描述了图灵原理与相分离原理在不同生态系统中的具体数学模型表达形式。接着, 分别阐述了图灵原理的涌现属性对生态系统功能以及相分离原理的涌现属性对细胞功能的作用。最后, 从多尺度自组织斑图、瞬态斑图和生物个体行为自组织3个方面对未来生态自组织理论发展方向进行了探讨。自组织研究在生态学与生物学研究中方兴未艾, 希望更多的学者在未来关注与参与该领域的发展。

关键词: 空间自组织, 规则空间格局, 图灵原理, 相分离原理, 生态模型, 生态系统功能

Abstract

Over the past 30 years, the self-organization theory has effectively explained the regular spatial patterning of ecosystems and has led to a proliferation of studies investigating spatial patterns in ecology and biology. Indeed, the emergent properties generated by this self-organization process are now recognized as critical to ecosystem functioning. Here, we review this important theoretical framework by assessing the definition and development of the concept of self-organization and by evaluating two fundamental theoretical principles of self-organization theory, the Turing principle and the phase separation principle. We further describe the mathematical models of each principle in the context of different, unique ecosystems, and explain the emergent properties of the Turing principle on ecosystem functioning and the phase separation principle on cell functions, respectively. Finally, we propose three promising future developments for ecological self-organization theory: multi-scale self-organization patterns, transient patterns, and individual behavioral self-organization. Our review provides an assessment of this fundamental ecological theory and offers exciting new research directions and applications.

Key words: spatial self-organization, regular spatial patterning, Turing principle, phase separation principle, ecological model, ecosystem functioning

葛振鹏, 刘权兴 (2020) 整体大于部分之和: 生态自组织斑图及其涌现属性. 生物多样性, 28, 1431-1443. DOI: 10.17520/biods.2020225.

Zhenpeng Ge, Quanxing Liu (2020) More than the sum of its parts: Self-organized patterns and emergent properties of ecosystems. Biodiversity Science, 28, 1431-1443. DOI: 10.17520/biods.2020225.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2020225

https://www.biodiversity-science.net/CN/Y2020/V28/I11/1431

图/表 4

图1 贻贝(Mytilus edulis)与“细胞”内的规则斑图以及相应的尺度依赖的反馈与密度依赖的运动的机制。(a)荷兰瓦登海潮间带贻贝床, 图片来自于本文的通讯作者; (b)基于液体光影技术对细胞内结构的重现, 图片来自https://liquidlightlab.com/artwork/3839057. html/, 版权属于Oil art: Steve Pavlovsky/Liquid Light Lab.; (c)尺度依赖的反馈示意图; (d)密度依赖的运动示意图, 其中主图中的两条曲线对应插图中的两条曲线, (c)和(d)修改自Liu等(2016)。

Fig. 1 The regular patterns of mussel and “cell”, and the scale-dependent feedback and density-dependent feedback. (a) Mussel bed in the intertidal zone of Wadden Sea, the Netherlands, the image come from the corresponding author of this paper; (b) The recurrence of a cell based on liquid light technology, the image is modified from https://liquidlightlab.com/home.html Copyright Oil art: Steve Pavlovsky/Liquid Light Lab.; (c) Schematic representation of scale-dependent feedback; (d) Schematic representation of density-dependent movement, (c) and (d) are modified from Liu et al (2016).

图2 方程组(7)和(8)所表示的系统相图与数值模拟。上方子图中红色实线为均相区与双节区的分界线, 蓝色实线为双节区与旋节线分解区的分界线。上方子图中“1”至“5”对应下方5张子图, 这些子图为相应的数值模拟结果, “1”与“5”位于均质区, 系统不存在斑图; “2”与“4”位于双节区, 系统存在点状斑图; “3”位于旋节线分解区, 系统存在迷宫状斑图。数值模拟采用周期边界条件, 数值模拟的matlab代码可在https://github.com/liufengyinxue/Liu_Mussel_PNAS下载。

Fig. 2 Phase diagram and numerical simulations of the equations (7) and (8). In the upper figure, the red solid line is the boundary between homogeneous zone and binodal zone, and the solid blue line is the boundary between binodal zone and spinodal decomposition zone. ‘1’ to ‘5’ in the upper figure correspond to the below five figures which are the numerical simulation results, ‘1’ and ‘5’ are located in homogeneous zone where the system doesn’t have pattern; ‘2’ and ‘4’ are located in binodal zone where the system has spot pattern, ‘3’ is located in the spinodal decomposition zone where the system has labyrinth pattern. The numerical simulation adopts periodic boundary condition, and the corresponding matlab code can be downloaded at https://github.com/liufengyinxue/Liu_Mussel_ PNAS.

图3 贻贝聚集的涌现属性之一: 降低被海鸟捕食的可能性

Fig. 3 One of the emergent properties of mussel aggregation: reducing the likelihood of predation by seabirds

图4 自组织在生态学中的发展趋势展望(英文版见附录1)。经过多年发展的图灵原理被发现并不能完全解释各种生态系统的多尺度斑图, 新兴的相分离原理亦然, 因而未来需要发展新的理论框架来理解生态系统的多尺度斑图; 瞬态斑图由于可能具备独特的生态功能在未来将备受关注; 生物对环境变化产生响应导致生物行为发生改变, 这种响应如何涌现至生态系统层次对理解跨尺度的生态系统自组织至关重要。

Fig. 4 The predominant research ideas of self-organization ecology in the future (The English version see appendix 1). After years of development, Turing's principle was found to be unable to fully explain the multi-scale patterns of various ecosystems, as well as the phase separation principle. Therefore, a new theoretical framework needs to be developed in the future to understand the multi-scale patterns of ecosystems. Transient pattern will be paid more attention in the future due to its unique ecological functioning. The response of organisms to environmental change leads to changes in biological behavior, and how this response emerges to the ecosystem level is critical to understanding the self-organization of ecosystems across scales.

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