The Evolution of Multicellularity – Mathew D. Herron, Peter L. Conlin, and William C. Ratcliff

This edited volume is an excellent overview of the field, ideal for newcomers or as a reference point for established researchers. You can order it here, or read the unformatted chapters for free here. [Twitter thread]

Here, we list papers that are currently in press/ preprints/ submitted and papers that were published in the last five years. For a full list, please refer to the Google Scholar profile of Prof. Ratcliff.


  • Zhang, C., Ratcliff, W. C. & Hammer, B. K. Constitutive expression of the Type VI secretion system carries no measurable fitness cost in Vibrio cholerae. (bioRxiv) (2023). [Twitter thread]
  • Pineau, R. M. et al. Emergence and maintenance of stable coexistence during a long-term multicellular evolution experiment. (bioRxiv) (2023). In revision for Nature Ecology and Evolution. [Twitter thread]
  • Peterson, A., Baskett, C., Ratcliff, W. C. & Burnetti, A. Using light for energy: examining the evolution of phototrophic metabolism through synthetic construction. (bioRxiv) (2023). In revision for Current Biology. [Twitter thread]
  • Dahaj, S. A. Z. et al. Spontaneous emergence of multicellular heritability. (bioRxiv) (2023). Revised MS submitted to The American Naturalist.
  • Conlin, P. L. et al. Division of labor promotes the entrenchment of multicellularity. (bioRxiv) (2023). [Twitter thread]
  • Pentz, J. T. et al. Evolutionary consequences of nascent multicellular life cycles. (bioRxiv) (2022). In revision for eLife. [Twitter thread]
  • MacGillivray, K. A. et al. Trade-offs constrain adaptive pathways to T6 survival. (bioRxiv) (2022). In revision for iScience. [Twitter thread]
  • Burnetti, A. & Ratcliff, W. The Origin of Phototrophy Reveals the Importance of Priority Effects for Evolutionary Innovation. (Preprints) (2022).

Published / in press


  • Bozdag, G. O. et al. De novo evolution of macroscopic multicellularity. Nature 1–8 (2023). [Twitter thread]
  • Ratcliff, W.C. The David Attenborough Style of Scientific Presentation. In press at Trends in Molecular Medicine.
  • Conlin, P.L. and W.C. Ratcliff. Evolution: Understanding the origins of facultative multicellular life cycles. Current Biology. 33, R356–R358.


  • Tong, K., Bozdag, G. O. & Ratcliff, W. C. Selective drivers of simple multicellularity. Current Opinion in Microbiology 67, 102141 (2022). [Twitter thread]
  • Ratcliff, W. C. The Biological Robots Are Coming! But Note They Have Been Here for ∼3.5 Billion Years. GEN Biotechnology 1, 26–27 (2022).
  • Day, T. C. et al. Varied solutions to multicellularity: The biophysical and evolutionary consequences of diverse intercellular bonds. Biophysics Reviews 3, 021305 (2022).
  • Day, T. C. et al. Cellular organization in lab-evolved and extant multicellular species obeys a maximum entropy law. eLife 11, e72707 (2022). [Twitter thread]
  • Datta, S. & Ratcliff, W. C. Illuminating a new path to multicellularity. eLife 11, e83296 (2022).
  • Burnetti, A. & Ratcliff, W. C. Experimental evolution is not just for model organisms. PLOS Biology 20, e3001587 (2022).


  • Márquez-Zacarías, P. et al. Evolution of Cellular Differentiation: From Hypotheses to Models. Trends in Ecology & Evolution 36, 49–60 (2021).
  • Márquez-Zacarías, P., Conlin, P. L., Tong, K., Pentz, J. T. & Ratcliff, W. C. Why have aggregative multicellular organisms stayed simple? Curr Genet 67, 871–876 (2021).
  • Libby, E. & Ratcliff, W. C. Lichens and microbial syntrophies offer models for an interdependent route to multicellularity. The Lichenologist 53, 283–290 (2021).
  • Isaksson, H., Conlin, P. L., Kerr, B., Ratcliff, W. C. & Libby, E. The Consequences of Budding versus Binary Fission on Adaptation and Aging in Primitive Multicellularity. Genes 12, 661 (2021).
  • Bozdag, G. O., Libby, E., Pineau, R., Reinhard, C. T. & Ratcliff, W. C. Oxygen suppression of macroscopic multicellularity. Nat Commun 12, 2838 (2021). [Twitter thread]


  • Yanni, D. et al. Topological constraints in early multicellularity favor reproductive division of labor. eLife 9, e54348 (2020).
  • Pentz, J. T. et al. Ecological Advantages and Evolutionary Limitations of Aggregative Multicellular Development. Current Biology 30, 4155-4164.e6 (2020).


  • Yanni, D., Márquez-Zacarías, P., Yunker, P. J. & Ratcliff, W. C. Drivers of Spatial Structure in Social Microbial Communities. Current Biology 29, R545–R550 (2019).
  • Tan, J. et al. Copper oxide nanoparticles promote the evolution of multicellularity in yeast. Nanotoxicology 13, 597–605 (2019).
  • Libby, E. & Ratcliff, W. C. Shortsighted Evolution Constrains the Efficacy of Long-Term Bet Hedging. The American Naturalist 193, 409–423 (2019).
  • Herron, M. D. et al. De novo origins of multicellularity in response to predation. Sci Rep 9, 2328 (2019).
  • Gulli, J. G., Herron, M. D. & Ratcliff, W. C. Evolution of altruistic cooperation among nascent multicellular organisms. Evolution 73, 1012–1024 (2019).


  • Mai, T. et al. Cycles in Zero-Sum Differential Games and Biological Diversity. in Proceedings of the 2018 ACM Conference on Economics and Computation 339–350 (Association for Computing Machinery, 2018).
  • Libby, E., Driscoll, W. W. & Ratcliff, W. C. Programmed cell death can increase the efficacy of microbial bet hedging. Sci Rep 8, 1120 (2018).
  • Jacobeen, S. et al. Cellular packing, mechanical stress and the evolution of multicellularity. Nature Phys 14, 286–290 (2018).
  • Jacobeen, S. et al. Geometry, packing, and evolutionary paths to increased multicellular size. Phys. Rev. E 97, 050401 (2018).
  • Herron, M. D., Zamani-Dahaj, S. A. & Ratcliff, W. C. Trait heritability in major transitions. BMC Biology 16, 145 (2018).
  • Herron, M. D., Ratcliff, W. C., Boswell, J. & Rosenzweig, F. Genetics of a de novo origin of undifferentiated multicellularity. Royal Society Open Science 5, 180912 (2018).