Solving the freeloaders paradox: Genetic associations and frequency-dependent selection in the evolution of cooperation among nonrelatives

Proceedings of the National Academy of Sciences of the United States of America
Leticia Avilés


One of the enduring problems in the study of social evolution has been to understand how cooperation can be maintained in the presence of freeloaders, individuals that take advantage of the more cooperative members of groups they are eager to join. The freeloader problem has been particularly troublesome when groups consist of nonrelatives, and no inclusive fitness benefits accrue to individuals that contribute more heavily to communal activities. These theoretical difficulties, however, are not mirrored by the numerous examples of cooperative or even altruistic behaviors exhibited by groups of nonrelatives in nature (e.g., many human groups, communally nesting bees, multiple queen-founding ants, cellular slime molds, and social bacteria). Using a model in which cooperation and grouping tendencies are modeled as coevolving dynamical variables, I show that the freeloader problem can be addressed when group-size effects on fitness are considered explicitly. I show that freeloaders, whose presence is reflected in the development of linkage disequilibrium between grouping and cooperation, increase in frequency when rare, but are selected against when common due to the reduced productivity of the groups they overburden with their pr...Continue Reading


Jun 7, 1988·Journal of Theoretical Biology·Robert Boyd, Peter J Richerson
Apr 1, 1972·Annals of Human Genetics·G R Price
Jul 1, 1964·Journal of Theoretical Biology·W D Hamilton
Jun 1, 1980·Theoretical Population Biology·M Uyenoyama, M W Feldman
Mar 27, 1981·Science·R Axelrod, W D Hamilton
Nov 5, 1999·Trends in Ecology & Evolution·G Bernasconi, J E Strassmann
Nov 7, 1980·Science·M J Wade
Aug 19, 2008·The American Naturalist·Leticia AvilésAsher D Cutter
Feb 1, 1995·Trends in Ecology & Evolution·R C Connor


May 12, 2004·Journal of Theoretical Biology·J A Fletcher, Martin Zwick
May 17, 2006·Trends in Ecology & Evolution·David Sloan Wilson
Dec 29, 2011·Proceedings of the National Academy of Sciences of the United States of America·Margaret Chatham Crofoot, Ian C Gilby
Sep 30, 2005·Proceedings. Biological Sciences·Ian M Hamilton, Michael Taborsky
Sep 9, 2006·Proceedings. Biological Sciences·Christoph HauertMichael Doebeli
Oct 2, 2008·Proceedings. Biological Sciences·Mikael Puurtinen, Tapio Mappes
Mar 4, 2014·PLoS Computational Biology·Thomas GarciaSilvia De Monte
Aug 31, 2013·PloS One·Roberto H SchonmannNestor Caticha
Jul 25, 2014·Theoretical Population Biology·Hilla BeharYoram Louzoun
Jul 28, 2004·The American Naturalist·Leticia AvilésAsher D Cutter
Sep 20, 2007·The American Naturalist·Adin Ross-GillespieAshleigh S Griffin
Jul 29, 2006·The American Naturalist·J A Fletcher, Martin Zwick
Nov 28, 2015·ELife·Thomas GarciaSilvia De Monte
Jan 26, 2008·The Quarterly Review of Biology·David Sloan Wilson, Edward O Wilson
Jun 15, 2011·Evolution and Human Behavior : Official Journal of the Human Behavior and Evolution Society·C Athena Aktipis
Oct 13, 2015·Journal of Theoretical Biology·Yuriy PichuginPaul B Rainey
Sep 7, 2007·Behavioural Processes·Michael Taborsky
Aug 18, 2004·Journal of Evolutionary Biology·K R Foster
Jul 22, 2009·Evolution; International Journal of Organic Evolution·Jonathan N Pruitt, Susan E Riechert
Aug 17, 2006·Journal of Evolutionary Biology·Laurent Lehmann, L Keller
Aug 17, 2006·Journal of Evolutionary Biology·J A Fletcher, M Doebeli
Sep 12, 2007·Molecular Ecology·Patrick Abbot, Vikram Chhatre
Mar 3, 2012·Evolution; International Journal of Organic Evolution·Jorge Peña
Jun 8, 2011·Evolution; International Journal of Organic Evolution·Simon T PowersRichard A Watson
Sep 29, 2009·Evolution; International Journal of Organic Evolution·Andy Gardner, Stuart A West
Nov 8, 2006·Journal of Theoretical Biology·J A Fletcher, Martin Zwick
May 17, 2015·Physics of Life Reviews·Zhen WangJun Tanimoto
Jun 18, 2015·Frontiers in Psychology·M Paula CacaultMathias Thoenig
Jul 31, 2012·Journal of Theoretical Biology·Jessica PurcellLeticia Avilés
Jan 30, 2015·PloS One·Daniel J van der PostCharlotte K Hemelrijk
Mar 17, 2010·Journal of Theoretical Biology·Matthijs van VeelenLeticia Avilés
Jan 7, 2017·The Journal of Experimental Biology·Corina E Tarnita
Sep 9, 2017·PLoS Computational Biology·Jaideep JoshiVishwesha Guttal
Apr 7, 2004·Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences·M V OlsonR Kaul
Apr 25, 2018·Oecologia·Brian R Haney, Jennifer H Fewell
Jan 8, 2013·Evolution; International Journal of Organic Evolution·Thomas Garcia, Silvia De Monte
Mar 19, 2016·Biological Reviews of the Cambridge Philosophical Society·Simon T Powers, Laurent Lehmann

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