Eusociality and a blow to kin selection

A new paper hit the internet today. "The Evolution of Eusociality" by Martin Nowak, Corina Tarnita, and E.O. Wilson re-frames an old evolutionary question and strikes a blow in an increasingly heated debate.

Eusociality is when individual organisms act as a collective reproducing unit. The best-known examples are ants and honeybees, but recently discovered examples include certain beetles, shrimp, and mole rats. Typically all reproduction is done by a single queen, and the rest of the colony exists only to support and protect the queen. Eusociality represents the highest degree of social organization found in nature.

The evolutionary origins of eusociality are something of a puzzle. To transition to eusociality, individuals must give up their own reproductive potential to support that of the queen. This is the ultimate sacrifice, as far as evolution is concerned. If evolution favors those who produce the most offspring, how can it select for actually giving up the chance to reproduce?

The classical answer to this question is kin selection: the idea that cooperative acts can occur between close relatives. Dawkins explained this using the concept of "selfish genes" that promote cooperation with others who have the same gene. One proponent, J.B.S. Haldane, famously said he would jump into a river to save two brothers, or eight cousins.

Ants and honeybees, the two oldest-known examples of eusocial animals, have a special genetic structure in which siblings share 3/4 of their genes, as compared to 1/2 in most sexual reproducers. It seemed reasonable that these close genetic relationships made possible such large-scale organization and extreme altruism.

However, as more eusocial species were discovered, including mammals, this association fell apart. There no longer appears to be any significant relationship between eusociality and relatedness of siblings.

Nowak, Tarnita, and Wilson provide a new model which focuses on the competition between reproductive units, which can be individual or collective. But perhaps more importantly, they thoroughly deconstruct the mathematics underlying kin selection theory.

The big debate in evolutionary theory right now is between those who believe all cooperation can be explained by kin selection (in its more mathematical guise of inclusive fitness theory), and those who believe that the more standard natural selection concept has more explanatory power. This debate has become increasingly heated in recent years.

Backed by rigorous mathematics, the authors argue that

Inclusive fitness theory is not a simplification over the standard approach. It is an alternative accounting method, but one that works only in a very limited domain. Whenever inclusive fitness does work, the results are identical to those of the standard approach. Inclusive fitness theory is an unnecessary detour, which does not provide additional insight or information.

The import of this argument might not be apparent to those not immersed in the field, but this paper could be a turning point in how the evolution of cooperation is understood. Social behavior cannot all be reduced to selfish genes. There are in fact many mechanisms allowing cooperation to evolve. Understanding these mechanisms will continue to be a fascinating question in evolutionary theory.