I do not agree with the cynical adage “science progresses–funeral by funeral”, but I fear that it might be true in your case for the subject of group selection.
In your response to the recent article in Nature by Martin Nowak, Corina Tarnita, and E.O. Wilson, which critiques inclusive fitness theory, you say this about group selection:
Edward Wilson was misunderstanding kin selection as far back as Sociobiology, where he treated it as a subset of group selection … Kin selection is not a subset of group selection, it is a logical consequence of gene selection. And gene selection is (everything that Nowak et al ought to mean by) ‘standard natural selection’ theory: has been ever since the neo-Darwinian synthesis of the 1930s.
I do not agree with the Nowak et al. article in every respect and will articulate some of my disagreements in subsequent posts. For the moment, I want to stress how alone you are in your statement about group selection. Your view is essentially pre-1975, a date that is notable not only for the publication of Sociobiology but also a paper by W.D. Hamilton, one of your heroes, who correctly saw the relationship between kin selection and group selection thanks to the work of George Price. Ever since, knowledgeable theoretical biologists have known that inclusive fitness theory includes the logic of multilevel selection, which means that altruism is selectively disadvantageous within kin groups and evolves only by virtue of groups with more altruists contributing more to the gene pool than groups with fewer altruists. The significance of relatedness is that it clusters the genes coding for altruistic and selfish behaviors into different groups.
Even the contemporary theoretical biologists most critical of multilevel selection, such as Stuart West and Andy Gardner, acknowledge what you still deny. In an earlier feature on group selection published in Nature, Andy Gardner is quoted as saying “Everyone agrees that group selection occurs”–everyone except you, that is.
You demonstrate more ignorance about group selection when you contrast it with gene selection. You correctly say that gene selection is standard natural selection theory. Essentially, it is a popularization of the concept of average effects in population genetics theory, which averages the fitness of alternative genes across all contexts to calculate what evolves in the total population. For that reason, it is an elementary mistake to regard gene selection as an alternative to group selection. Whenever a gene evolves in the total population on the strength of group selection, despite being selectively disadvantageous within groups, it has the highest average effect compared to the genes that it replaced. Please consult the installment of my “Truth and Reconciliation for Group Selection” series titled “Naïve Gene Selectionism” for a refresher course. While you’re at it, check out the installment titled “Dawkins Protests–Too Much“.
The Nowak et al. article includes several critiques of inclusive fitness theory that need to be distinguished from each other. One issue is whether inclusive fitness theory is truly equivalent to explicit models of evolution in multi-group populations, or whether it makes so many simplifying assumptions that it restricts itself to a small region of the parameter space. A second issue is whether benefiting collateral kin is required for the evolution of eusociality and other forms of prosociality. A third issue is whether inclusive fitness theory, as understood by the average evolutionary biologist and the general public, bears any resemblance to inclusive fitness theory as understood by the cognoscenti.
In an e-mail discussion among theorists on both sides of the debate that I organized several years ago, I conducted a citation analysis comparing Hamilton’s original 1964 papers on group selection with his 1975 reformulation in the light of the Price equation. These papers are cited in a ratio of approximately 15:1 with no tendency for citations of the 1975 paper to increase in frequency over the decades. In other words, most evolutionary biologists still have a pre-1975 conception of inclusive fitness theory, no matter how much it has been elaborated by the cognoscenti. This degree of illiteracy about foundational issues is an embarrassment for the field of evolutionary biology. The Nowak article is a wake-up call for the average evolutionary biologist and the general public to reconsider the conventional wisdom about inclusive fitness theory, in addition to the debates that will take place among the cognoscenti.
Richard, if you wish to join the debates among the cognoscenti, you will need to abandon the priestly way that you make pronouncements, expecting what you say to be taken on faith, and rejoin the ranks of scientists who hold each other accountable for what they say.