Iki Sakumi
Center for the Evolutionary Origins of Human Behavior, Kyoto University, Inuyama, Aichi, 484-8506, Japan.
Biol Rev Camb Philos Soc. 2025 Aug;100(4):1467-1483. doi: 10.1111/brv.70009. Epub 2025 Mar 5.
The origins of play remain a profound puzzle in animal evolution. Play is often characterised as a seemingly non-functional behaviour that confers little survival or reproductive benefit. This characteristic makes the evolution of play appear paradoxical under Darwinian principles, which posit that traits must be beneficial to be selected. Given that the adaptive benefits are unclear even for the well-established forms of play in extant animals, it seems improbable that an incipient form of play in the earliest stages of evolution emerged due to a decisive selective advantage. The conventional view that has gained traction suggests that play evolved not as an adaptive trait but as a by-product of energetic, ontogenetic, ecological, and psychological facilitating factors. Building upon previous empirical and theoretical studies, this review discusses the evolutionary relationship between play and exploration. More specifically, it argues that relying on the classification dividing exploration into intrinsic and extrinsic types can help us articulate both the evolutionary and mechanistic continuities and discontinuities between play and exploration. Based on this distinction, this article proposes the following hypothesis: play originally evolved as a by-product of curiosity-motivated intrinsic exploration. This hypothesis is supported by recent empirical evidence indicating that play may have evolved by co-opting some of the mechanisms of curiosity, including (i) the commonality between stimuli that elicit curiosity and those that elicit play, and (ii) the shared neural basis of curiosity and play involving the reward and executive control systems. I also discuss new testable predictions derived from this hypothesis and outline future research directions, including comparative phylogenetic studies, eco-sociological analyses, and psycho-behavioural approaches.
玩耍的起源在动物进化中仍然是一个深刻的谜题。玩耍通常被描述为一种看似无功能的行为,几乎没有生存或繁殖益处。这种特性使得玩耍的进化在达尔文主义原则下显得自相矛盾,达尔文主义原则认为特征必须有益才能被选择。鉴于即使对于现存动物中已确立的玩耍形式,其适应性益处也不明确,在进化早期阶段出现的初始玩耍形式似乎不太可能是由于决定性的选择优势。已获得认可的传统观点认为,玩耍并非作为一种适应性特征进化而来,而是作为能量、个体发育、生态和心理促进因素的副产品。基于先前的实证和理论研究,本综述讨论了玩耍与探索之间的进化关系。更具体地说,它认为依靠将探索分为内在和外在类型的分类方法,可以帮助我们阐明玩耍与探索之间在进化和机制上的连续性与非连续性。基于这一区分,本文提出以下假设:玩耍最初是作为由好奇心驱动的内在探索的副产品进化而来的。这一假设得到了最近实证证据的支持,这些证据表明玩耍可能是通过利用一些好奇心机制进化而来的,包括:(i)引发好奇心的刺激与引发玩耍的刺激之间的共性,以及(ii)好奇心和玩耍涉及奖励和执行控制系统的共享神经基础。我还讨论了从这一假设得出的新的可测试预测,并概述了未来的研究方向,包括比较系统发育研究、生态社会学分析和心理行为方法。