Evolution of multicellularity, bet hedging and
Publications (Google Scholar link)
Submitted / In Revision
Herron, M. and W.C. Ratcliff. Trait heritability in major transitions. BioRxiv link.
Libby, E., and W.C. Ratcliff. Time-dependent fitness effects can drive bet-hedging populations extinct. BioRxiv link.
Libby, E., W.W. Driscoll and W.C. Ratcliff. Programmed cell death can improve the efficacy of microbial bet hedging.
Rebolleda-Gomez, M., W.C. Ratcliff, J. Fankhauser, and M. Travisano. Multicellularity as a key trait facilitating adaptive
radiation. BioRxiv link.
McNally, L., E. Bernardy, J. Thomas, A. Kalziqi, J.T. Pentz, S. Brown, B. Hammer, P. Yunker and W.C. Ratcliff. 2017.
Killing by Type VI secretion drives clonal phase separation and correlates with cooperation. Nature Communications
14371. Open access link.
Weitz, J.S., S.P. Brown, C. Eskin, K. Paarporn, and W.C. Ratcliff. 2016. Replicator dynamics with feedback-evolving
games: towards a co-evolutionary game theory. PNAS. 11: 47. E7518–E7525. Link.
Libby, E., P. Conlin, B. Kerr and W.C. Ratcliff. 2016. Stabilizing multicellularity through ratcheting. Philosophical
Transactions of the Royal Society B. 331:20150444. Open access link.
Conlin, P. and W.C. Ratcliff. 2016. Trade-offs drive the evolution of increased complexity in nascent multicellular digital
organisms. In Multicellularity: Origins and Evolution. Vienna Series in Theoretical Biology. Download.
Pentz, J., B. Taylor and W.C. Ratcliff. 2016. Apoptosis in snowflake yeast: novel trait, or side effect of toxic waste? Journal
of the Royal Society Interface. 118: 20160121. Link.
Pentz, J.T., T. Limberg, N. Beerman, and W.C. Ratcliff. 2015. Predator escape: an ecologically realistic scenario for the
evolutionary origins of multicellularity. Evolution: Education and Outreach, 8:13. Open access link.
Ratcliff, W.C., Fankhauser, J.F., Rogers, D., Greig, D. and Travisano, M. 2015. Origins of multicellular evolvability in
snowflake yeast. Nature Communications 6: 6102. Open access link.
Ratcliff, W.C., P. Hawthorne, and E. Libby. 2015. Courting disaster: how diversification mode affects fitness under risk.
Evolution 69: 126-135. Download.
Libby, E. and W.C. Ratcliff. 2014. Ratcheting the evolution of multicellularity. Science 346 (6208), 426-427. Download.
Libby, E., W.C. Ratcliff, M. Travisano and B. Kerr. 2014. Geometry shapes evolution of early multicellularity. PLoS
Computational Biology 10 (9:e1003803), 1-12. Download.
Pentz, J.T., M. Travisano and W.C. Ratcliff. 2014. Clonal development is evolutionarily superior to aggregation in wild-
collected Saccharomyces cerevisiae. Proceedings of the Fourteenth International Conference on the Simulation and
Synthesis of Living Systems, 550-554. Download.
Ratcliff, W.C. and M. Travisano. 2014. Experimental evolution of multicellular complexity in Saccharomyces cerevisiae.
BioScience, 64 (5), 383-393. Download.
Ratcliff, W.C., A. Raney, S. Westreich, and S. Cotner. 2014. A novel laboratory activity for teaching about the evolution of
multicellularity. The American Biology Teacher 76 (2), 81-87. Download.
Ratcliff, W.C., M. Herron, K. Howell, J. Pentz, F. Rosenzweig, and M. Travisano. 2013. Experimental evolution of an
alternating uni- and multicellular life cycle in Chlamydomonas reinhardtii. Nature Communications, 4: 2742. Open
Ratcliff, W.C., M. Hoverman, M. Travisano. and R. F. Denison. 2013. Disentangling direct and indirect fitness effects of
microbial dormancy. The American Naturalist, 182:147-156. Download.
Ratcliff, W.C., J.T. Pentz, and M. Travisano. 2013. Tempo and mode of multicellular adaptation in experimentally-evolved
Saccharomyces cerevisiae. Evolution, 67: 1573–1581. doi: 10.1111/evo.12101. Download.
E.T. Kiers, W.C. Ratcliff and R.F. Denison. 2012. Single strain inoculation may creates spurious correlations between
legume fitness and rhizobial fitness. New Phytologist, 198: 4-6. DOI: 10.1111/nph.12015 Download.
Rebolleda-Gomez, M, W.C. Ratcliff and M. Travisano. 2012. Adaptation and divergence during experimental evolution of
multicellular Saccharomyces cerevisiae. Proceedings of the Thirteenth International Conference on the Simulation and
Synthesis of Living Systems, 99-104. Download.
Ratcliff, W.C., R. F. Denison, M. Borrello and M. Travisano. 2012. Experimental evolution of multicellularity. PNAS, 109:5,
1595-1600. Open Access Link.
Ratcliff, W.C., K. Underbakke and R. F. Denison. 2011. Measuring the fitness of symbiotic rhizobia. Symbiosis, 55:2, 85-
Ratcliff W.C. and R.F. Denison. 2011. Alternative actions for antibiotics. Science, 332:6029, 547-548. Download
Ratcliff W.C. and R.F. Denison. 2010. Individual-level bet hedging in the bacterium Sinorhizobium meliloti. Current Biology
20:1740-1744. Open Access link. See commentary by Zhang X.X. and P.B. Rainey. 2010: “Bet hedging in the
Ratcliff W.C. and R.F. Denison. 2010. Bacterial persistence and bet hedging in Sinorhizobium meliloti. Communicative &
Integrative Biology 4:1, 1-3. Open Access link.
Ratcliff W. C., P. Hawthorne, M. Travisano, and R. F. Denison. 2009. When stress predicts a shrinking gene pool, trading
early reproduction for longevity can increase fitness, even with lower fecundity. PLoS One, 4:e6055. Open Access link
Ratcliff W. C. and R. F. Denison. 2009. Rhizobitoxine producers gain more poly-3-hydroxybutyrate in symbiosis than do
competing rhizobia, but reduce plant growth. The ISME Journal, 3:870-872. Download
Ratcliff W. C., S. V. Kadam, and R. F. Denison. 2008. Polyhydroxybutyrate supports survival and reproduction in starving
rhizobia. FEMS Microbiology Ecology, 65:391-399. Download
We are extremely grateful to the National Science Foundation, NASA Exobiology, the Packard Foundation,
the Keck Foundation, and the Simons Foundation for supporting our research!