食物网稳定性机制研究进展:一个基于等级系统的框架

食物网理论沟通了群落生态学和生态系统生态学，将生物多样性和生态系统功能的研究统一起来，是理解生态系统运作机制的关键。自从1973年Robert May的经典研究引发著名的"复杂性-稳定性"论辩之后，人们认识到食物网的稳定性是其结构维持、功能发挥和动态演化的一个重要前提，并开始了对食物网稳定性机制的探索。早期研究主要关注只包含拓扑关系的定性食物网，但后来人们逐渐认识到相互作用强度的重要性，并提出了诸如自限性、弱相互作用、适应性捕食等一系列机制。本文系统梳理了过往研究中模块层面的各类稳定性机制和全网层面对各模块的整合机制，从而清晰地展示了"模块-全网"双层框架的全貌。通过在其基础上的扩展，进而提出了一个基于等级系统的食物网稳定性框架，并从动力学和能量学角度，对各层级内部的稳定性机制以及层级之间的关系进行了探讨，以期为建立普适的食物网稳定性理论提供一些思路。未来的研究方向包括：①将稳定性机制的研究从食物网扩展到更一般的生态网络；②综合考虑生物物理要素、动力学稳定性、系统对能流功率的追求、环境的平稳程度、演化历史等影响因素，从而得到关于食物网结构和动态的更为深刻的认识。

Research progress on stability mechanisms of food webs: a hierarchy-based framework

Food web theory, which bridges community ecology and ecosystem ecology and unifies the research of biodiversity and ecosystem function, holds the key for understanding the operation mechanism of ecosystems. Since the famous debate on "complexity-stability" invoked by Robert May''s classical work in 1973, researchers began to realize that stability is an important premise of food webs for their structure maintenance, function exertion and dynamic evolution. During the past half century, the exploration of food web stability has transitioned from qualitative food webs containing only topological relations to quantitative food webs that also containing interaction strength. A series of mechanisms such as self-limiting, weak interaction and adaptive predation have been proposed, and a "module-web" double-layer framework has been gradually established. Based on previous studies, we figured out the various stability mechanisms at the module level and the integration mechanisms of each module at the whole web level, which further enhanced this framework. To overcome its shortcomings in the face of complex food webs, we further proposed a food web stability framework based on a hierarchical structure. From the perspective of dynamics and energetics, the internal stability mechanism of each level and the relationship between the levels were discussed in order to provide some ideas for building a more universal food web stability theory. Future research directions include:(1) expanding the research on stability mechanism from food webs to more general ecological networks; (2) obtaining a deeper understanding of the structure and dynamics of food webs through comprehensively considering influencing factors such as biophysical elements, dynamic stability, the pursuit of energy flow power by the system, the stability of the environment, and evolution history.