Tag Archives: E.O. Wilson

Got questions about inclusive fitness?

Over at his blog, Andrew Gelman briefly mentions the recent profile of E. O. Wilson in the Atlantic, and along the way mentions the dustup over inclusive fitness that I may have mentioned here before  (did I? It’s hard to remember).   At the end, he makes a throw-away comment which drove me nuts:

The article also discusses Wilson’s recent crusade against selfish-gene-style simplifications of human and animal nature. I’m with Wilson 100% on this one. “Two brothers or eight cousins” is a cute line but it doesn’t seem to come close to describing how species or societies work, and it’s always seemed a bit silly to me when people try to loop everything back to a selfish-gene story.

I’ve been trying to think of a similarly aggravating comment to make about statistics in return;  maybe “lies, damned lies, and statistics”?  “You can prove anything with statistics”?  “Bayesian statistics suck because I don’t understand where priors come from?”  It bugged me enough that I left this comment:

It doesn’t seem like you know much about inclusive fitness, a theory has been massively successful in evolutionary biology. Despite the odd and unsupported comments made by Nowak et al., it stands firm as a well-supported and useful body of theory. Here’s a link to the letter published in response to Nowak et al.’s original article, signed by 137 authors including most of the field’s brightest minds:


The appeal to authority doesn’t mean that they’re right, of course, but extraordinary claims require extraordinary evidence; Nowak et al. have done nothing but ignore the entire published literature on inclusive fitness spanning decades and comprised of hundreds if not thousands of studies, while proposing a mathematical model that adds nothing to our understanding beyond what current theory already provides.

I respect your work on statistics, have always enjoyed reading your blog, and your book (BDA) is sitting on my shelf right now, but your offhand comment above is uninformed and very aggravating; I’d like to deal with that aggravation by offering to assist you in understanding one of the most powerful explanatory mechanisms in evolutionary biology. The letter above provides a succinct summary of the evidence that Nowak et al. ignore, but it might be a bit much for a non-technical audience; I haven’t published directly in this field, but I do work in evolutionary biology and I should be able to answer any specific questions you may have if you would like to pose them. If I can’t answer them myself, I will find people who can.

I’m not going to go into a full blown recapitulation of inclusive fitness theory and then defend it, because I’d have to write several inconveniently long books to do so.  But since I made the offer over there, I’ll make it here too for any interested readers:  if you have questions burning you up about this whole “inclusive fitness” thing, ask them here in the comments and I will do my best to answer them for you.  And if I don’t know what the answer is, I’ll find it.  No question is too small, though I make no promises on how long or short my answers will be!

I’ll leave off with a quotation from a fantastic book by Andrew Bourke that I’m reading right now, Principles of Social Evolution:

Like any large and active field of investigation, the theoretical study of social evolution is not free from disagreements and unresolved issues (e.g. Taylor and Nowak, 2007; West et al. 2007a).  Paradoxically, while the potential richness of inclusive fitness theory as a general theory of social evolution is still underappreciated, the theory is sometimes perceived as an entrenched orthodoxy. A tendency therefore exists for iconoclastically-minded theoreticians to derive models of cooperation in novel ways and then announce them to be fundamental additions to existing theory (e.g. Killingback et al. 2006; Nowak 2006; Ohtsuki et al. 2006; Traulsen and Nowak 2006).  It is healthy for orthodoxies to be continually challenged by new theories and new data.  However, to date, these models have fallen short of true novelty, as other authors have shown that their results are capable of being derived from inclusive fitness theory (e.g. Grafen 2007a, 2007b; Lehmann et al. 2007a, 2007b; West et al. 2007a).  Indeed, inclusive fitness theory has a long history of successfully assimilating apparent challenges and alternatives (Grafen 1974; Queller 1992; Lehmann and Keller 2006a).  This is not surprising when one considers its deep foundations in the theory of natural selection.  Although it is premature to declare a consensus, a substantial body of opinion therefore holds that claims of fundamental extensions to inclusive fitness theory will have to be radically innovative, as well as robust, to be accepted as such (e.g. Lehmann and Keller 2006a; West et al. 2007a).  For all these reasons, Hamilton’s (1964) inclusive fitness theory will underpin the conceptual reasoning employed throughout this book (pp. 22-23).


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Nowak et al. take it on the chin.

Not pretty...photo from hunterseakerhk

At the end of September last year, I was attending the opening of the ISBE conference (International Society for Behavioral Ecology) in Perth, Australia when Stuart West got up to give a keynote address.  One of the first things he said was something along the lines of “I’ve been getting a lot of questions about this, but I’m not going to address the eusociality paper”, which was met by a chorus of hoots and cries from the audience.  It was a great first-hand experience of the kind of reaction that the paper by Martin Nowak, Corina Tarnita, and E. O. Wilson, published in Nature last summer, was getting at the time.  (If you don’t know what I’m talking about, Carl Zimmer has a good write-up about it).  I’d read the paper at the time and dismissed it for largely (as far as I can tell) the same reasons as everyone else did:  it stinks.  And then, I forgot about it.

I was recently reminded, however, by the buzz that is going around right now about the responses to this paper that have just been published by Nature, some seven(!) months later.  Jerry Coyne has blogged about it, and it’s a good read for many reasons not least of which is that he’s an author on one of the critiques along with 136 people, among them some of the biggest names in evolutionary biology and social evolution.  I’d point out some of the people on that list and their accomplishments, but I’d get the same result if I wrote a book on the subject of Nowak et al.’s paper – they’d all be in it.  Honestly, the only question I have about that is how did they organize it?  The critique itself is worth reading as well, if only for the concise beat-down that they apply to the subjects of their wrath.  (D. S. Wilson disagrees, but I think that he’s managed to completely miss the point.  Read and decide for yourself, though.  Full disclosure:  I don’t agree with much of what D. S. Wilson says.)

And the icing on the cake has to be this video, from Jon Wilkins.  I’d love to embed it, but it’s on Xtranormal and WordPress.com apparently doesn’t support embedding from there, so click through and have a good gander.  If you have any interest in this issue, it’s a great video.

Do I have anything to add to this?  Honestly, only a vague sorrow, especially about Nowak.  When I was starting my master’s, I came across the work his lab was doing on evolutionary graph theory and loved it;  it was a great inspiration to me at the time, and it remains so to this day.  But when I read things like this eusociality paper, I see another Nowak, and it’s not a Nowak that I’m particularly enamored of.

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