Monday 28 December 2015

Answers for Doug Hoxworth

Doug Hoxworth had some questions which he aired in public recently:

  • If transmission only happens based on the “selfish” gene theory, how do behavioral or personality traits that are advantageous for the group but disadvantageous for the individuals within the group ever get transmitted to the next generation and become dominant/ubiquitous? If it is heritable, how can this phenomenon be explained?

    • Kin selection. For example, worker sterility is disadvantageous to the worker as an individual, but beneficial to the group they are part of. The reason for this is because the worker and the reproductives (queens and drones) share genes. It is possible to explain this in terms of group selection as well. However group selection explanations often turn into simplistic group-level functionalism - whereas kin selection quantifies relatedness (using 'r') and so is a better tool for making quantitative predictions with - in those cases where relatedness is less than 1.

  • How can altruism to non-relatives be explained?

    • For humans, the most significant mechanisms are probably cultural kin selection, virtue signaling, manipulation, over-generalization and environmental mismatch - in roughly that order.

  • If it is through enculturation and imitation, how are these behaviors so ubiquitous at a very young age, if not at birth (even applicable to animals/organisms other than humans that presumably do not emit pheromones indicating that they are unambiguously related, e.g., ants)?

    • Cultural transmission is often advantageous to individuals. Using socially-transmitted information from others can give a short-cut to learning - if maladaptive traditions can be avoided. There are various ways of avoiding maladaptive traditions.

Doug's article contrasts group selection with the selfish gene. In fact these ideas are compatible. The selfish gene is compatible with kin selection and it is compatible with group selection. Richard Dawkins might not agree - but we don't need to heed him here. The disagreement in this area mostly lies elsewhere.

The problem with group selection is not so much that it's wrong, but that it causes confusion among its practitioners. For decades group selection advocates held out hope that their theory would make novel predictions. In the last decade, this hope has mostly collapsed and most now recognize that group selection and kin selection are broadly equivalent. It is a matter of different accounting techniques, so to speak.

The main problem is that we know that kin selection is strongest between close relatives. Group selection advocates often want to apply the theories to whole tribes and to warfare. This sort of group selection isn't equivalent to kin selection - and, for the most part, it doesn't actually work.

What is left of this whole debate? Not too much. There's some noise surrounding Hamilton's inclusive fitness concept, but this is mostly coming from Martin Nowak and friends - and his papers have been met with ridicule. At this stage, most of the facts seem to be in and the group selection controversy resembles a mopping up operation.

6 comments:

  1. http://kinselections.blogspot.com/2015/12/answers-for-doug-hoxworth.html

    Q1: If it (e.g., pure altruism) is heritable, how can this phenomenon be explained?
    A1: Kin selection
    Q1A: This does not provide adequate explanation for social behaviors like pure/disinterested altruism that is often directed toward non-kin (even sometimes extending to the out-group) and with no expectation of reciprocation in more advanced organisms like dolphins, elephants, and especially primates (including humans). Viewing this as some sort of biological misfiring or parasite does not seem to bear itself out when looking at the research.

    Q2: How can altruism to non-relatives be explained?
    A2: Cultural kin selection, virtue signaling, manipulation, over-generalization and environmental mismatch
    Q2A: I'm not familiar with these and will have to do more research. Can you direct me to articles/books that elaborate on this. For me, the attempts that I've seen on this are much too complex and involved and I frankly get lost through the circuitous development of the model(s).

    Q3: If it is through enculturation and imitation, how are these behaviors so ubiquitous at a very young age, if not at birth?
    A3: Cultural transmission is often advantageous to individuals...
    Q3A: I did not see a response to this nor did I see it even acknowledged neither in the body or the endnotes of Dawkins's "The Selfish Gene."

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    1. Regarding: "Kin selection does not provide adequate explanation for social behaviors like pure/disinterested altruism that is often directed toward non-kin (even sometimes extending to the out-group) and with no expectation of reciprocation in more advanced organisms like dolphins, elephants, and especially primates (including humans)"...

      For humans, there's cultural kin selection based on shared memes - rather than shared genes. The other cases are outside the scope of your original question. You asked how behavioral/personality traits that are advantageous for the group but disadvantageous for the individuals within the group get transmitted to the next generation. The answer to that is kin selection - including cultural kin selection. Modern defensible versions of group selection are not an alternative explanations to kin selection, they are reformulations of kin selection which make all the same predictions. If there exists disinterested altruism that isn't explicable by kin selection (in elephants and dolphins) then it isn't explicable by group selection either - and it doesn't fit into the category you described.

      My post linked to resources on cultural kin selection and manipulation. Virtue signalling is sometimes referred to by academics as the "heart on your sleeve" hypothesis. This, manipulation and environmental mismatch are covered unsympathetically by Boyd and Richerson in "Solving the Puzzle of Human Cooperation". While their treatment is critical, that article does offer a summary and has reasonable references.

      Sorry that my reply to the third question appears unresponsive to your main point. It looks as though I was answering another question. Essentially, I deny the premises here. Young uncultured humans are typically very self-centred and don't have a mental model which allows them to empathize with others. They are 'born selfish' - as Richard Dawkins put it.

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    4. After reading Alan Grafen's essay 'Natural Selection, Kin Selection and Group Selection' as recommended by Dawkins in the footnote on page 110 and his new chapter 12 "Nice Guys Finish First" it does seem quite clear that the answer to my question is found in Evolutionarily Stable Strategies (ESS) like tit-for-tat. I think that I do agree that group selection doesn't seem to work, most of all that it seems to require competition between groups (and so cannot work in the absence of this pressure) and the replicator or physical mechanism for transmission is not clear to me.

      What puzzles me is the following quotes from Dawkins, where he obviously struggles with humans that seem to defy expectations:

      "So far, I have not talked much about man in particular, though I have not deliberately excluded him either...Are there any good reasons for supposing our own species to be unique? I believe the answer is yes.

      Most of what is unusual about man can be summed up in one word: 'culture'."
      [p. 189 chapter 11 memes: the new replicators]

      "Man may well have spent large portions of the last several million years living in small kin groups. Kin selection and selection in favour of reciprocal altruism may have acted on human genes to produce many of our basic psychological attributes and tendencies...The argument I shall advance, surprising as it may seem coming from the author of the earlier chapters, is that, for an understanding of the evolution of modern man, we must begin by throwing out the gene as the sole basis of our ideas on evolution."
      [p. 191 chapter 11 memes: the new replicators]

      "We can even discuss ways of deliberately cultivating and nurturing pure, disinterested altruism—something that has no place in nature, something that has never existed before in the whole history of the world. We are built as gene machines and cultured as meme machines, but we have the power to turn against our creators. We, alone on earth, can rebel against the tyranny of the selfish replicators.*"
      [pp. 200-201 chapter 11 memes: the new replicators]

      His footnote to the last line above does not really answer the question but simply makes the same assertion again in other words ("perfectly possible to hold that genes exert a statistical influence on human behaviour while at the same time believing that this influence can be modified, overridden or reversed by other influences" vs. "rebel against the tyranny of the selfish replicators" then giving the example of contraception as if that settles the issue).

      What he seems to ignore is that it is now clear that all primates, but especially humans, seem to be wired for prosociality (as demonstrated in the work of people like Frans de Waal on primates and Paul Bloom and his wife, Karen Wynn on babies). I do not think that cultural kin selection and memes account for this. So I think that all of this militates against Dawkins's statement that this "has no place in nature, something that has never existed before in the whole history of the world" and your statement that "Young uncultured humans are typically very self-centred and don't have a mental model which allows them to empathize with others. They are 'born selfish' - as Richard Dawkins put it." All of the current research that is occurring with primates and babies disproves this model and assumed premise.

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  2. "What is left of this whole debate? Not too much...most of the facts seem to be in and the group selection controversy resembles a mopping up operation."
    I'm no so sure. If that is the case, I am not sure why articles and books continue to be written about this issue, especially the attempt to provide an evolutionary explanation and basis for morality and true altruism.

    In fact, Dawkins's endnote to p. 110 "Wynne-Edwards... has been mainly responsible for promulgating the idea of group selection." in Chapter 7 Family Planning on page 297 says the same:

    "...Group selection, in the sense in which we have all long understood it, is even more out of favour among biologists than it was when my first edition was published. You could be forgiven for thinking the opposite: a generation has grown up, especially in America, that scatters the name 'group selection' around like confetti. It is littered over all kinds of cases that used to be (and by the rest of us still are) *clearly and straightforwardly understood as something else, say kin selection*. I suppose it is futile to become *too annoyed* by such semantic parvenus. Nevertheless, the whole issue of group selection was *very satisfactorily settled* a decade ago by John Maynard Smith and others, and *it is irritating* to find that we are now two generations, as well as two nations, divided only by a common language. It is particularly unfortunate that philosophers, now belatedly entering the field, have started out *muddled* by this recent caprice of terminology. I recommend Alan Grafen's essay 'Natural Selection, Kin Selection and Group Selection' as a clear-thinking, and I hope now definitive, sorting out of the neo-group selection problem."

    What often bothers me about Dawkins is his tone. I really do like a lot of Dawkins's ideas but his smugness (perhaps even nationalist??) and arrogant annoyance by the mere fact that the idea is even still kicking around is very off-putting and makes it difficult to listen to anything else he has to say. His dismissiveness does not win him points in my book. Perhaps it is just not so obvious an open/shut as he thinks it is.

    I personally think that it is presumptuous and gratuitous to assume that group selectionist advocates are somehow all unsuspecting ignoramuses (apparently duped by the 'group selection' meme being scattered around like confetti in the USA...) as seems to be the implication from his statements quoted above. He groups all of them together and assumes they are under some spell and then quickly dismisses them.

    After reading nearly all of his book (I'm on chapter 12), I can say that I have not read anything that approaches what he claims. Perhaps I missed it in my reading or it's in chapter 12 or 13 but I did not think that Dawkins provided the knock-out punch that he thinks that he has. Also, I have downloaded but have yet to read Alan Grafen's essay but I will be sure to do so to see if he provides the resolution to the issue in a parsimonious way as Dawkins claims.

    That being said, I did find chapter 11 on Memes of "The Selfish Gene" very intriguing. I really like the idea but again I think it ignores the work of people like Frans de Waal on morality in primates and Paul Bloom on morality in children. Perhaps it is just because they are not biologists that Dawkins does not interact with them. I am actually not familiar with all of Dawkins's works so perhaps I am mistaken when I say that he doesn't interact with the work or type of work like the abovementioned individuals. If that is the case, I apologize in advance. I will have to admit that I am just beginning to dive into all of this and answer the questions. Unfortunately as of right now, I am not seeing the silver bullet and do not think that kin selection does what Dawkins and co are thinking that it does. For what it's worth...

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