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COOPERATION, ORGANISMALITY, AND
MAJOR TRANSITIONS IN EVOLUTION

The second reason for expanded interest in cooperation is a growing appreciation that it is important for how organisms came to be. Cooperative major transitions in life alter the raw material for natural selection in fundamental ways (Buss, 1987; Maynard Smith and Szathmáry, 1995). One of the earliest transitions brought molecules together into cells in which the fates of all were intertwined in a cooperative network. Eukaryotes themselves represent a major transition resulting from the capture of a bacterium that becomes the mitochondrion (Margulis, 1970). The level of cooperation between these partners is profound but not complete. Mitochondria are maternally inherited and do not go through meiosis, and thus will favor daughter production and have no interest in son production.

Another major transition resulted in multicellularity (Queller, 1997, 2000; Grosberg and Strathmann, 1998; Herron and Michod, 2008). Multicellularity has evolved multiple times in both bacterial and eukaryote lineages. Animals and plants have elaborated multicellularity into a plethora of diverse types. There are also a number of comparatively simple multicellular forms, like some single-species biofilms, the algal group Volvocales, or Dictyostelium (Herron and Michod, 2008; Strassmann and Queller, 2010). The transition to multicellularity is different from the transition to eukaryotes because the former involves an aggregate of like entities, whereas the latter binds different elements. The major transitions can thus be categorized as fraternal, with like cooperating with like, or egalitarian, where the cooperating units bring different things to the collaboration (Queller, 1997). Either kind of collaborative organism will usually retain conflicts, but these conflicts must be controlled if the partnership is to survive. How these controls operate is a major research topic under this view of life.

The selective factors that favored a past transition are not easy to study because they have already completed their work. There are living systems that could be considered to be more representative of transitional stages, however. These, we believe, may be the most productive for investigation into the advantages of cooperation and how conflict is controlled. We have argued elsewhere that organisms themselves can be defined as adapted bundles of cooperative elements, wherein actual conflict is at a minimum (Queller and Strassmann, 2009; Strassmann and Queller, 2010). In a 2D space, with one axis being cooperation and the other being conflict, organisms are those collaborative living units at the high end of cooperation and the low end of conflict. There is variation in the level of organismality, however, and those lacking complete coop-



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