Preprint Review Version 2 Preserved in Portico This version is not peer-reviewed

The Non-Tragedy of the Non-Linear Commons

Version 1 : Received: 12 April 2020 / Approved: 15 April 2020 / Online: 15 April 2020 (02:51:41 CEST)
Version 2 : Received: 14 June 2020 / Approved: 16 June 2020 / Online: 16 June 2020 (12:35:40 CEST)

How to cite: Archetti, M.; Scheuring, I.; Yu, D. The Non-Tragedy of the Non-Linear Commons. Preprints 2020, 2020040226. Archetti, M.; Scheuring, I.; Yu, D. The Non-Tragedy of the Non-Linear Commons. Preprints 2020, 2020040226.


Public goods are produced at all levels of the biological hierarchy, from the secretion of diffusible molecules by cells to social interactions in humans. However, the cooperation needed to produce public goods is vulnerable to exploitation by free-riders — the Tragedy of the Commons. The dominant solutions to this problem of collective action are that some form of positive assortment (due to kinship or spatial structure) or enforcement (reward and punishment) is necessary for public-goods cooperation to evolve and be maintained. However, these solutions are only needed when individual contributions to the public good accrue linearly, and the assumption of linearity is never true in biology. We explain how cooperation for nonlinear public goods is maintained endogenously and does not require positive-assortment or enforcement mechanisms, and we review the considerable empirical evidence for the existence and maintenance of nonlinear public goods in microbiology, cancer biology, and behavioral ecology. We argue that it is time to move beyond discussions about assortment and enforcement in the study of cooperation in biology.


positive assortment; public goods; collective action; common-pool resources; cooperation; game theory; group selection; kin selection; social dilemma


Biology and Life Sciences, Other

Comments (1)

Comment 1
Received: 16 June 2020
Commenter: Douglas Yu
Commenter's Conflict of Interests: Author
Comment: Extensive edits for clarity
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