Inclusive Fitness Theory (IFT) is of considerable importance to Darwinian evolutionists. The theory is concerned with the phenomena of altruistic behaviour and eusocial societies, both of which involve the willingness of some animals to sacrifice themselves for the benefit of the group. Darwin struggled to provide a rationale, and so did those who followed him. It was Bill Hamilton who put together a coherent theory and Richard Dawkins who popularised it in The Selfish Gene. For many, IFT has achieved the status of orthodoxy. It was an intellectual and an emotional shock, therefore, when a paper appeared in Nature (August 2010) from three prominent evolutionary biologists saying that the IFT paradigm is unproductive. Responses were immediate and much of it was hostile. Science journalist Roger Highfield provided an overview of the controversy:
"The mainstream media often like to portray the scientific community as regularly riven by blazing rows. Scientists, understandably, complain: after all, if you go to any mainstream academic conference, you won't find any hint of controversy about the MMR jab or the reality of climate change, let alone argy-bargy over the basic facts of evolution. But in the past few weeks, I have witnessed a bare-knuckle brouhaha that would make an uninformed outsider gasp at how bloody a battle over a seemingly arcane issue can be. The row was triggered by a paper in Nature by Martin Nowak, Edward Wilson and Corina Tarnita of Harvard University. While some hailed it as "revolutionary" and a "return to rigour", others condemned it as "sad", "baffling", "irritating" and "unscholarly"."
Honeybees and ants are eusocial insects. In the colonies, individuals cooperate and share tasks to increase productivity. (Source here)
Readers of this blog might be interested in the reactions of Jerry Coyne, who was mystified by Ed Wilson's participation in the paper.
"I don't know what's gotten into E. O. Wilson. He's certainly the world's most famous evolutionary biologist, and has gone from strength to strength over the years, winning two Pulitzer Prizes, writing great general books on not only ants but conservation and social behavior. [. . .] But now Wilson, along with some collaborators like David Sloan Wilson and Martin Nowak, is definitely heading off on the wrong track. They're attacking kin selection, maintaining not only that it has nothing to do with the evolution of social insects, but that's it's also a bad way to look at evolution in general. And they're wrong - dead wrong."
Coyne is also outraged by the publication of this paper in a prestigious journal: "Finally, a big raspberry to the folks at Nature who decided to publish such a strange paper in the interest of stirring up controversy. If they'd gotten decent reviewers, and followed their advice, it never would have seen print." Richard Dawkins was also quick to repudiate the paper and say that the authors display numerous misunderstandings:
"This is no surprise. Edward Wilson was misunderstanding kin selection as far back as Sociobiology, where he treated it as a subset of group selection [. . .] Inclusive fitness theory is not some kind of supernumerary excrescence, to be 'resorted to' only if 'standard natural selection theory' is found wanting (Misunderstanding One). On the contrary, inclusive fitness theory is one way of expressing what was logically inherent in the synthesis ever since Fisher and Haldane, but had been largely overlooked because people (with the exception of those two geniuses) didn't think about collateral kin. [. . .]"
A welcome, reflective, commentary on the controversy is provided by David Hughes in Trends in Ecology and Evolution. As a young researcher, Hughes had Hamilton as a supervisor; then became a member of the socio-biology research community; then worked at Harvard University where he "engaged extensively with all three of the authors" as they developed their ideas on IFT. He made a point of talking to both sides of the controversy. His short essay does not grapple with the technical issues, but presents some analysis of "the manner in which we carry out evolutionary biology today" and offers some reflective guidelines for constructive debate.
The first problem identified is that "we currently give far too much weight to celebrity status, impact factors and instant commentary". It would appear that far too many have not had the time to engage with the arguments, but are swayed by which experts they trust. In response to this, Hughes first advocates thorough preparation by reading the relevant literature, followed by a ruthless quest for precise language on the part of active participants in the debate. Only then can hard, but relevant, questions be asked.
"Having publically (at conferences) and privately questioned both the authors and opponents, I know that these celebrities on both sides are willing to patiently explain their position. However, because it is difficult to raise hard questions with high profile celebrities we are typically left to discuss the validity and significance of the suggested new approach amongst ourselves. Earnest discussions on difficult topics in biology departments and on conference circuits are oftentimes less than ideal."
Secondly, the science community has imbibed the zeitgest of our age that "lacks a culture of respectful debate". This means that close questioning is often perceived as aggression.
"It seems gladiatorial to ask a difficult question, whether at a conference, seminar or a journal club, but how else can we resolve important issues? Leaving it to comment boxes on blogs is not the way forward. Respectful debating is especially needed now that biology is so large and diverse: we are all ignorant, only on different topics. Eroding this ignorance requires the respectful exchange of information."
Thirdly, there is a confusion of evidence-based science and expert opinion. Hughes finds that much of the debate actually boils down to the expression of opinion, and this cannot be a good thing. The response should be to build arguments on data.
"Because science is evidentiary we do ourselves a disservice by not pushing the data forward. I know that it has become difficult to grasp the meaning of certain data as the standard of both data collection and analysis has risen very sharply. That is why thoughtful and earnest discussions are required. The goal of both parties should be to clearly explain the facts and the ways in which those facts have been collected, however torturous that may be."
Fourthly, Hughes refers to the problem of publishing. As well as the cult of the expert, we have to realise that there is a cult of the high impact journal. These two cults feed off each other. The science community needs to "recognise the playing field we are on". But judgments have to be made.
"However, to decide this you, dear colleague, need to read the paper and question, question, question. We as a community need to experiment, experiment, experiment so as to make available the data that a forward-moving, integrative biology needs. (This of course requires that we increase the rate at which we publish negative results as well). The decisions on such great issues of the day cannot be derived from sound bites in on-line magazines manipulated by sensationalist journalists whose industry thrives on polarization. Nor should our view be the recycled opinion of a celebrity. As idealistic as it sounds, do please be part of a movement to build a society where what is important is the content of the research, the quality of the evidence and the originality and validity of the ideas and not the venue of publication or the celebrity status of the researchers."
All this is good advice and it is also transferable advice. Shift the focus to the issue of intelligent design in nature and all is relevant. Darwinians often play the card of "celebrity status, impact factors and instant commentary". There is typically a lack of respectful debate, with ad hominem arguments and the dark shadow of being expelled hanging over those who step out of line. The point about confusion between evidence-based science and expert opinion is directly relevant: it comes up again and again. The cult of the high impact journal affects the ID debate, as one often comes across examples of people appealing to a journal critique of ID as though it has said the last word and debate is over. There is no acknowledgement that ID scholars have responded to the challenge and demonstrated that the debate is far from over.
A point not made by Hughes but nevertheless relevant is that evolutionary biology is, for many, more than a science. It forms part of their world view. Their opinions take on an identity of their own, because they have persuaded themselves that they have reached the position where they can deliver truth. This explains something of the passion and the polarisation of debates.
"Theologians worry away at the 'problem of evil' and a related 'problem of suffering.' ... On the contrary, if the universe were just electrons and selfish genes, meaningless tragedies like the crashing of this bus are exactly what we should expect, along with equally meaningless good fortune. Such a universe would be neither evil nor good in intention. It would manifest no intentions of any kind. In a universe of blind physical forces and genetic replication, some people are going to get hurt, other people are going to get lucky, and you won't find any rhyme or reason in it, nor any justice. The universe we observe has precisely the properties we should expect if there is, at bottom, no design, no purpose, no evil and no good, nothing but blind, pitiless indifference. As that unhappy poet A.E. Housman put it: 'For Nature, heartless, witless Nature Will neither care nor know.' DNA neither cares nor knows. DNA just is. And we dance to its music."
(Dawkins R., River out of Eden: A Darwinian View of Life, Phoenix: London, 1996, p.155.)
Recent developments in sociobiology and the scientific method
David P. Hughes
Trends in Ecology and Evolution, Volume 26, Issue 2, February 2011, Pages 57-58 | doi:10.1016/j.tree.2010.12.002
You might have noticed that a recent high profile paper on Inclusive Fitness Theory (IFT), presented in the journal Nature, has lead to an enormous and emotional response in sociobiology. The authors, Martin Nowak, Corina Tarnita and Ed Wilson, state that Bill Hamilton's (and the majority of subsequent researchers') paradigm of how altruistic behaviors and eusocial societies evolve is unproductive. The reaction to the publication has been vocal and polarizing, leading to camps of opinions. [snip]
Eusociality, in which some individuals reduce their own lifetime reproductive potential to raise the offspring of others, underlies the most advanced forms of social organization and the ecologically dominant role of social insects and humans. For the past four decades kin selection theory, based on the concept of inclusive fitness, has been the major theoretical attempt to explain the evolution of eusociality. Here we show the limitations of this approach. We argue that standard natural selection theory in the context of precise models of population structure represents a simpler and superior approach, allows the evaluation of multiple competing hypotheses, and provides an exact framework for interpreting empirical observations.
Gilbert, N., Altruism can be explained by natural selection. Evolutionary biologists overturn long-held kin-selection theory. Nature News, (25 August 2010) | doi:10.1038/news.2010.427
Highfield, R. The evolution of altruism: selflessness in ants? That's fighting talk (The Daily Telegraph, 12 October 2010).
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