Abstract Ecosystems are complex adaptive systems (CAS) by nature, which means that macroscopic patterns and properties emerge from, and feed back to affect, the interactions among adaptive individual ecological agents. These agents then further adapt (genetically) to the outcomes of those interactions. The concept of self-organization has become increasingly important for understanding ecosystem spatial heterogeneity and its consequences. It is well accepted that ecosystems can self-organize, and that resulting spatial structures carry functional consequences. Feedbacks from the outcome of spatial pattern to the individual agents from which patterns emerge, are an essential component of the definition of CAS but have been rarely examined for ecosystems. We explore whether spatial self-organization provides a mechanism for such feedback for ecosystems as CAS, that is, whether ecosystem-level outcomes of self-organized patterning could feed back to affect or even reinforce local pattern-forming processes at the agent level. Diffuse feedbacks of ecological and evolutionary significance ensue as a result of spatial heterogeneity and regular patterning, whether this spatial heterogeneity results from an underlying template effect or from self-organization. However, feedbacks directed specifically at pattern-forming agents to enhance pattern formation—reinforcing feedback—depend upon the level of organization of agents. Reinforcing evolutionary feedbacks occur at the individual level or below. At the ecosystem level, evidence for mechanisms of feedback from outcomes to patterning to agents forming the patterning remain tenuous. Spatial self-organization is a powerful dynamic in ecosystem and landscape science but feedbacks have been only loosely integrated so far. Self-organized patterns influencing dynamics at the ecosystem level represent “order for free”. Whether or not this free order generated at the ecosystem level carries evolutionary function or is merely epiphenomenal is a fundamental question that we address here.

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生态系统空间自组织：无所事事的自由秩序？

摘要 生态系统本质上是复杂的适应系统（CAS），这意味着宏观模式和特性从适应性个体生态代理之间的相互作用中产生并反馈到影响。然后这些代理进一步适应（遗传）这些相互作用的结果。自组织的概念对于理解生态系统空间异质性及其后果变得越来越重要。众所周知，生态系统可以自组织，由此产生的空间结构会产生功能性后果。从空间模式的结果到模式出现的个体代理的反馈是 CAS 定义的重要组成部分，但很少对生态系统进行检查。我们探索空间自组织是否为 CAS 等生态系统提供了一种反馈机制，即自组织模式的生态系统层面的结果是否可以反馈影响甚至加强代理层面的局部模式形成过程。作为空间异质性和规则模式的结果，生态和进化意义的扩散反馈随之而来，无论这种空间异质性是由潜在的模板效应还是自组织造成的。然而，专门针对模式形成代理以增强模式形成的反馈——强化反馈——取决于代理的组织水平。强化进化反馈发生在个体层面或更低层面。在生态系统层面，从结果到模式到形成模式的代理的反馈机制的证据仍然很薄弱。空间自组织是生态系统和景观科学中的强大动力，但到目前为止，反馈只是松散地整合。在生态系统层面影响动态的自组织模式代表“免费秩序”。这种在生态系统层面产生的自由秩序是否具有进化功能或仅仅是附带现象是我们在这里解决的一个基本问题。