C. Athena Aktipis, Amy M. Boddy, Gunther Jansen, Urszula Hibner, Michael E. Hochberg, Carlo C. Maley & Gerald S. Wilkinson (2015), Philosophical Transactions of the Royal Society B 370(1673):20140219 — DOI-10.1098/rstb.2014.0219

Key takeaways

  • The framing (Introduction). “Multicellularity requires the suppression of cell-level fitness in order to promote organism level fitness”, and effective multicellularity needs mechanisms for suppressing conflict as well as cooperation. Cheating is defined functionally, as breaking shared rules for a cell-level advantage, not as an evolved strategy.
  • Five foundations. Proliferation inhibition, controlled cell death, division of labour, resource allocation and transport, and maintenance of the extracellular environment. Their breakdown gives uncontrolled proliferation, inappropriate survival, deregulated differentiation, resource monopolization and a degraded environment.
  • The survey. Cancer-like phenomena (abnormal proliferation or differentiation, a deliberately broad notion) are reported across the lineages with complex multicellularity, which evolved independently at least seven times. All or nearly all bilaterian animals look susceptible, and cnidarians suggest the problem is ancient. Gaps, such as sponges and Ulvophyceae, may be lack of study.
  • What generalises (Discussion). Cheating on division of labour is present in all forms of multicellularity; resource and environment cheating are documented only in animals and plants so far. Every report surveyed showed both “demographic” and “economic” cheating.
  • Suppression. Complex multicellularity “represents a highly sophisticated cheater detection and suppression system”, working cell-autonomously, through tissue architecture and systemically. Proliferative units such as intestinal crypts may be an intermediate level of selection that makes suppression modular.

Standing in the record

Filed on 2026-10-05 at the owner’s request, as a supplement to the essay on organizational will. It is the biological case of the essay’s §6.3 “capture from below”: lower-level units gaining at the collective’s expense, and the collective as a system for suppressing them. Okasha’s overview of the major transitions (LIT-749) supplies the general principle, and cautions against applying the language of cheating where no collective action problem has been shown. The account is THEORY-135.