tag:blogger.com,1999:blog-61375156833161676032024-03-15T08:39:09.631-07:00Kin selectionsTim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.comBlogger67125tag:blogger.com,1999:blog-6137515683316167603.post-47233543063401770042017-06-22T06:01:00.001-07:002017-06-22T17:23:23.040-07:00Nowak and Wilson are at it again in 2017Nowak and Wilson are at it again, with a 2017 paper:<P>
"<a href="http://www.pnas.org/content/114/22/5665.full">The general form of Hamilton’s rule makes no predictions and cannot be tested empirically</a>"<P>
It's the old claim that "survival of the fittest" is a tautology (since the fittest are those that survive) repackaged for kin selection.<P>
Evolutionary biologists have long had a stock answer to the
claim that "survival of the fittest" is a tautology. This hinges on
the difference between <em>expected fitness</em> and <em>observed fitness</em>.<P>
"Survival of those with the highest observed fittesses" would indeed be a kind of circular argument.
However, "survival of those with the highest expected fittesses" is making a falsifiable statement
about the model with which the expected fitnesses are calculated.<P>
Nowak and Wilson say:
<BLOCKQUOTE>The mathematical investigation of HRG reveals three astonishing facts.
First, HRG is logically incapable of making any prediction about any
situation because the benefit, BB, and the cost, CC, cannot be known
in advance. They depend on the data that are to be predicted. At the
outset of an experiment, BB and CC are unknown, and so there is no
way to say what Hamilton’s rule would predict.</BLOCKQUOTE><P>
Those are observed fitnesses (measured after the fact). Rather obviously, if you
want to make predictions, you need to be using <em>expected</em> fitnesses
when determining B and C. Nowak and Wilson are simply mistaken
in claiming that there's no way in which these could possibly
be calculated in advance. You could predict these values by using a mathematical model
of the situation, or by using a computer simulation - for example.<P>
IMHO, it is pretty amazing that some scientists bothered to write a whole
paper about this issue, without first understanding it.<P>
Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com0tag:blogger.com,1999:blog-6137515683316167603.post-54156768485680035802016-09-28T17:58:00.001-07:002016-09-28T18:18:16.369-07:00Group selection vs kin selection: evidence from examples<img src="https://nyamiledotcom.files.wordpress.com/2014/09/nuer-dinka.png" align=right width=240>Modern versions of kin selection and group selection have turned out to be <A HREF="http://kinselections.blogspot.com/2012/09/group-selection-and-kin-selection.html">essentially equivalent</A>. However, historical examples of his selection and group selection have covered some rather different territory. Group selection enthusiasts have chosen phenomena such as senescence and warning signals. From the perspective of kin selection, some of the most obvious examples include breast feeding, parental care and nepotism. The kin selection examples are good, but the group selection ones are more dubious. I've previously argued that the reason group selection was associated with dodgy examples is because advocates sought to distinguish their theory from kin selection - and so avoided examples where it was obvious that kin selection was responsible. The details are in my 2104 article "<a href="http://kinselections.blogspot.com/2014/08/how-kin-selection-pushed-group.html">How kin selection pushed group selection into the scientific fringes</a>".<P>
I think we are seeing a similar phenomenon with cultural group selection and <A HREF="http://on-memetics.blogspot.com/2014/08/cultural-kin-selection.html">cultural kin selection</A>. My articles on cultural kin selection are illustrated with examples of where kin selection works on shared memes, rather than shared genes. These examples include uniforms, money and religious brotherhoods and sisterhoods. I think these are real examples of cases where Hamilton's rule applies to memes. However, the cultural group selection examples look rather different. One example often given involves Dinka and Nuer populations in the Sudan. Another example involves monogamous marriage customs. In these cases, a simple explanation is that some memes involved are fitter than other ones. Evidence that the memes involved are deleterious to individuals yet spread through group level reproduction or extinction seems to be completely missing. Maybe these are just bad memes. Cultural group selection seems largely unnecessary.<P>
The tendency of group selection enthusiasts to pick bad examples seems common to both the organic realm and the cultural realm. However the explanation I have previously given for the bad examples in the organic realm does not seem to be applicable to the cultural realm. In the case of cultural evolution, cultural group selection seems to be much better known than cultural kin selection is. So, the idea that cultural group selection was bent out of shape to avoid cultural kin selection does not seem very plausible. Another explanation would seem to be required.<P>
My perception is that the cultural group selection enthusiasts are choosing examples of cases where one group does better than another one for cultural reasons. However, that's not a particularly good way of finding examples of where Hamilton's rule applies. The native Americans did badly as a group recently - but that wasn't really due to kin selection or group selection. Their environment changed rapidly and they failed to adapt quickly. Hamilton's rule isn't terribly relevant in cases like these - even though one group is exterminating another one. This is part of the historical problem with group selection. People see groups apparently exterminating each other and then leap to group selectionist explanations without bothering to properly test them to see if they are plausible.<P>
One possible reply by group selection advocates is that kin selection and group selection don't cover the same ground after all. However, lots of people have held that position over the years, but the evidence for it doesn't seem to be there. If you accept equivalence, I think is is hard to deny the prevalence of bad quality examples of group selection. When was the last time you heard breastfeeding or parental care given as an example of group selection? Never, right? Group selection advocates are not thinking about mother-offspring pairs as "groups" - even though that's where the maths says the effect is strongest. Inferior examples associated with group selection arise in both the cultural and organic realms. My previous argument about <a href="http://kinselections.blogspot.com/2014/08/how-kin-selection-pushed-group.html">how kin selection pushed group selection into the scientific fringes</a> doesn't look as though it is the whole explanation. Maybe group selection is just misleading or confusing.<P>
Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com0tag:blogger.com,1999:blog-6137515683316167603.post-86046114967531999672016-09-10T20:17:00.000-07:002016-09-10T20:17:54.080-07:00The Meaning of Human Existence (not)<img src="https://www.timeshighereducation.com/sites/default/files/styles/the_breaking_news_image_style/public/Pictures/web/f/a/p/book-review-the-meaning-of-human-existence-by-edward-o-wilson.jpg?itok=qM3MkHLu" width=200 align=right>I read chapter 6 of Edward O Wilson's "The Meaning of Human Existence" from 2014.<P>
This is the chapter on 'inclusive fitness theory'. It was a jaw-dropping
experience. If we are to believe Edward's account, he helped to lead a
valiant crusade against a long-established theory and succeeded in
overturning it in 2010. Of course, the reality is that the 2010
paper in question was an embarrassing scientific farce, which had no useful
new results and just paraded the authors' own misconceptions.<P>
I think that the lesson here is that the egos of famous elderly scientists
can get out of control, leading to delusions of grandeur. Other famous
elderly scientists should take note: don't let this happen to you.<P>
Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com0tag:blogger.com,1999:blog-6137515683316167603.post-33729273263236043132016-02-16T17:59:00.002-08:002016-02-16T17:59:22.244-08:00Google NGRAM for kin selectionHere's the Google NGRAM for group selection, kin selection, inclusive fitness, multilevel selection. It is interesting, I think.<P>
<iframe name="ngram_chart" src="https://books.google.com/ngrams/interactive_chart?content=group+selection%2Ckin+selection%2Cinclusive+fitness%2Cmultilevel+selection&year_start=1900&year_end=2000&corpus=15&smoothing=3&share=&direct_url=t1%3B%2Cgroup%20selection%3B%2Cc0%3B.t1%3B%2Ckin%20selection%3B%2Cc0%3B.t1%3B%2Cinclusive%20fitness%3B%2Cc0%3B.t1%3B%2Cmultilevel%20selection%3B%2Cc0" width=900 height=500 marginwidth=0 marginheight=0 hspace=0 vspace=0 frameborder=0 scrolling=no></iframe><P>
Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com0tag:blogger.com,1999:blog-6137515683316167603.post-82624869134973117472015-12-28T16:16:00.002-08:002015-12-28T16:34:20.049-08:00Answers for Doug HoxworthDoug Hoxworth had some questions which he <a href="https://evolution-institute.org/article/lets-find-a-compromise-between-group-selection-and-the-selfish-gene/">aired in public recently</A>:<P>
<UL>
<LI>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?<P>
<UL>
<LI>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.<P>
</UL>
<LI>How can altruism to non-relatives be explained?<P>
<UL>
<LI>For humans, the most significant mechanisms are probably <a href="http://on-memetics.blogspot.com/2014/08/cultural-kin-selection.html">cultural kin selection</a>, virtue signaling, <a href="http://on-memetics.blogspot.com/2014/11/tim-tyler-manipulation.html">manipulation</a>, over-generalization and environmental mismatch - in roughly that order.<P>
</UL>
<LI>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)?<P>
<UL>
<LI>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.<P>
</UL>
</UL>
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.<P>
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.<P>
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 <EM>actually</EM> work.<P>
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 <a href="https://www.youtube.com/watch?v=zHYsTSmD84w">ridicule</a>. At this stage, most of the facts seem to be in and the group selection controversy resembles a mopping up operation.<P>Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com6tag:blogger.com,1999:blog-6137515683316167603.post-60557666292046896742015-11-14T11:32:00.001-08:002015-11-14T11:32:11.969-08:00Stuart West on kin vs group selectionHere's Stuart West classic video discussion of kin vs group selection:<P>
<iframe src="https://player.vimeo.com/video/8202768?color=e50090&byline=0" width="800" height="600" frameborder="0" webkitallowfullscreen mozallowfullscreen allowfullscreen></iframe> <p><a href="https://vimeo.com/8202768">4. Group selection debate - Stuart West</a> from <a href="https://vimeo.com/londonevolution">LERN: London Evolution</a> on <a href="https://vimeo.com">Vimeo</a>.</p>Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com0tag:blogger.com,1999:blog-6137515683316167603.post-92195423529716731842015-08-23T05:59:00.000-07:002015-08-23T06:00:56.362-07:00Prospective kin<img src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEibpc-4wACRaSIUa0zbMKMM9_kN0CSc6dyZ6DzwnJE2cZO9KfC6Q_w1ikb3trm2vpwARJ7cN_uIDFoTeBbxl799Wxb77hhiNCX-ZnMvnvPfiUvVVQrJveoMedwIuTInN4WrqHrkiR7iM_Y/s1600/TheMAN.JPG" width=290 align=right>Organisms treat their mates better than average conspecifics - even before they have had
any offspring with them.<P>
It isn't hard to think of reasons for this, but many of of those reasons invoke the idea
of <em>prospective kin</em> - unborn children.<P>
Most children have a probability of becoming an ancestor - and it is possible to extend this
idea to children that haven't been born - and to children that haven't been even conceived.<P>
Pretty standard kin selection models can thus be applied to courtship behaviour, bower construction,
nuptual gifts - and so forth. These are cases where cooperative behaviour without relatedness occurs.
In fact, there <em>is</em> relatedness - relatedness to unborn children.<P>Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com0tag:blogger.com,1999:blog-6137515683316167603.post-24456182743579174292015-03-28T19:23:00.002-07:002015-05-17T15:54:15.462-07:00David Queller on the evolution of eusociality<img src="https://wubio.wustl.edu/files/biology/styles/person_node/public/people/images/queller-headshot_0.jpg?itok=CBOWo2in" align=right>Liao, Rong and Queller just weighed in on the <a href="http://www.nature.com/nature/journal/v466/n7310/full/nature09205.html">Nowak/Tarnita/Wilson 2010 model of the evolution of eusociality</a>.<P>
That paper was <EM>mostly</EM> smacked down by critics because of its <EM>delusional</EM> depiction of kin selection, but few bothered to criticize its model of the evolution of eusociality. Now, <a href="http://journals.plos.org/plosbiology/article?id=10.1371/journal.pbio.1002098">David Queller's gone through the model</a>, and the results are not too pretty.<P>
<BLOCKQUOTE>They had a modeling strategy that should work and should be fine, but they weren’t careful enough when they made claims about their models’ novel results</BLOCKQUOTE>
...David Queller is quoted as saying.<P>
From the abstract of <a href="http://journals.plos.org/plosbiology/article?id=10.1371/journal.pbio.1002098">the article</a>:<P>
<BLOCKQUOTE>The claim of these authors was bolstered by a new model of the evolution of eusociality with novel conclusions that appeared to overturn some major results from inclusive fitness. Here we report an expanded examination of this kind of model for the evolution of eusociality and show that all three of its apparently novel conclusions are essentially false. Contrary to their claims, genetic relatedness is important and causal, workers are agents that can evolve to be in conflict with the queen, and eusociality is not so difficult to evolve. The misleading conclusions all resulted not from incorrect math but from overgeneralizing from narrow assumptions or parameter values. For example, all of their models implicitly assumed high relatedness, but modifying the model to allow lower relatedness shows that relatedness is essential and causal in the evolution of eusociality. Their modeling strategy, properly applied, actually confirms major insights of inclusive fitness studies of kin selection. This broad agreement of different models shows that social evolution theory, rather than being in turmoil, is supported by multiple theoretical approaches. It also suggests that extensive prior work using inclusive fitness, from microbial interactions to human evolution, should be considered robust unless shown otherwise.</BLOCKQUOTE>
Queller's paper certainly makes entertaining reading. However I can't help thinking that it takes the Nowak/Tarnita/Wilson paper too seriously. The most obvious response to that paper is, I think, ridicule. Jon Wilkins hit roughly the right note with <a href="https://www.youtube.com/watch?v=zHYsTSmD84w">Important Harvard Scientists Attack Kin Selection</a>.<P>
News coverage:<P>
<UL>
<LI><a href="http://journals.plos.org/plosbiology/article?id=10.1371/journal.pbio.1002098">
Relatedness, Conflict, and the Evolution of Eusociality</a> by Xiaoyun Liao, Stephen Rong and David C. Queller
<LI><a href="http://www.futurity.org/inclusive-fitness-eusocial-883242/">Did a scientific battle about altruism just end?</a> by Diana Lutz (<a href="https://news.wustl.edu/news/Pages/modeling-supports-inclusive-fitness.aspx">source</a>)
<LI><a href="https://whyevolutionistrue.wordpress.com/2015/03/27/new-paper-shows-that-nowak-et-al-were-wrong-kin-selection-remains-a-valuable-tool-in-evolutionary-biology/">New paper shows that Nowak et al. were wrong: kin selection remains a valuable concept in evolutionary biology</a> by Jerry Coyne
<LI><a href="http://www.unz.com/gnxp/much-ado-about-inclusive-fitness/">Much Ado About Inclusive Fitness</a> by Razib Khan
</UL>
<hr>Update 2015-05-09:<p>
Nowak and Allen reply in <a href='http://journals.plos.org/plosbiology/article?id=10.1371/journal.pbio.1002134'>Inclusive Fitness Theorizing Invokes Phenomena That Are Not Relevant for the Evolution of Eusociality</a><p>
Liao, Rong and Queller reply: <a href=http://journals.plos.org/plosbiology/article?id=10.1371/journal.pbio.1002133">Some Agreement on Kin Selection and Eusociality?</a><p>
Nowak's papers on the topic are <a href="http://ped.fas.harvard.edu/inclusive-fitness">here</a>.<P>
Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com0tag:blogger.com,1999:blog-6137515683316167603.post-21909007844078311352015-03-01T10:14:00.000-08:002015-03-01T10:22:57.213-08:00Popular internet dictionary parrots kin selection FUDI visited the <a href="http://dictionary.reference.com/browse/inclusive+fitness">reference.com entry for inclusive fitness</a> today and was surprised to see the following <a href="http://en.wikipedia.org/wiki/Fear,_uncertainty_and_doubt">F.U.D.</a>:<P>
<BLOCKQUOTE>Inclusive fitness turns out to be a phantom measure that cannot be obtained.</BLOCKQUOTE>
...and...<P>
<BLOCKQUOTE>Inclusive fitness theory is neither useful nor necessary to explain the evolution of eusociality or other phenomena.</BLOCKQUOTE>
Further, the second quote was mirrored by Google at the top of its search results for "inclusive fitness" - as though it was a definition of the term! You might still be able to see this for yourself <a href="https://www.google.com/#safe=off&q=inclusive+fitness">here</a>.<P>
Low levels of machine intelligence can cause this sort of problem, it seems.<P>
Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com0tag:blogger.com,1999:blog-6137515683316167603.post-91972742566102214582015-02-06T18:43:00.003-08:002015-02-09T13:50:42.743-08:00David Sloane Wilson has doubts; asks for help<amg src="http://www.otago.ac.nz/philosophy/gfx/USR_DSW.jpg" align=right>People have been trying for decades to find something that group selection
predicts that kin selection does not. Now the results are in. The most
coherent forms of group selection make no new predictions. They are an
<EM>alternative accounting technique</EM> that makes the same set of predictions that kin
selection makes. This is <EM>fairly</EM> widely acknowledged by most of the parties
involved these days. Of course, kin selection has been part of the standard orthodoxy in biology for decades.<P>
However, it seems that David Sloane Wilson still has doubts about this. He doesn't
see how to apply kin selection in some cases - and he's <a href="https://evolution-institute.org/article/challenge-to-kin-selectionists-explain-this/">publicly asking for help</a>. As far as I can see, David has nothing. One of his three
examples is <EM>cultural group selection</EM>. I replied
<a href="http://on-memetics.blogspot.com/2015/02/david-sloane-wilson-on-cultural-kin.html">here</a>.<P>
I'm not sure how much help David will get from other kin selection enthusiasts.
Most have been unimpressed with David's attempts to rechristen and take
credit for a well-established existing theory. Group selection has a
long association with junk science. Kin selection depends on close relatedness
to produce adaptations. If you emphasize group membership instead of close
relatedness - you include a bunch of cases where there's no close relatedness,
and adaptations are not produced.
<a href="http://kinselections.blogspot.com/2014/08/how-kin-selection-pushed-group.html">Group
selection was pushed by kin selection into the scientific fringes</a> - where it focused on
cases not obviously explicable by kin selection - which were mostly cases where the theory
didn't actually work. In short, group selection is kin selection's evil twin.<P>
David engages in a bit of a straw man attack on kin selection in his article. He focuses
on Hamilton's rule. Hamilton's rule is one of the findings by kin selection theorists -
but it isn't the same thing as kin selection. Here's how Hamilton explained the topic:<P>
<BLOCKQUOTE>The existence of altruism in nature can be explained by thinking about the replication of genes. We need to descend to the level of the gene, rather than the individual, in order to see that the gene exists surrounded by copies of identical genes that exist in all its relatives - in particular in its close relatives, its siblings, who have a half chance of carrying a copy of that particular gene, its offspring, which also have a half chance, parents: a half-chance, cousins: one eighth, etc. Seeing this swarm of genes that exists around a particular one, we can then ask what is the behavior caused by this gene that is most likely to cause the propagation of this set of copies in the relatives around it.</BLOCKQUOTE>
That is more like kin selection in a nutshell. Of course, these days, we have to clarify that it's the <a href="http://on-memetics.blogspot.com/2015/02/the-evolutionary-gene.html">evolutionary gene</a> that we mean here. Hamilton's rule is the product of kin selection and a bunch of assumptions.<P>
Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com0tag:blogger.com,1999:blog-6137515683316167603.post-6018814144143875042015-01-04T15:09:00.001-08:002015-01-04T15:15:22.328-08:00Herbert Gintis vs inclusive fitness theory<img src="http://edge.org/sites/default/files/styles/member-photo/public/member-pictures/bk_826_herbert_gintis.JPG?itok=JQGxs3nF" align=right>Herbert Gintis has an elaborate attack on inclusive fitness theory on his web site. It's titled: <a href="http://www.umass.edu/preferen/gintis/GeneralizedHR.pdf">Inclusive Fitness and the Sociobiology of the Genome</a>. It says, among other things:<P>
<BLOCKQUOTE>The general point is that if there is a conflict among loci concerning fitness maximization, and if the frequency of alleles at one locus affect the fitness costs and payoffs at other loci, then it is logically impossible that the allele at each locus maximize its inclusive fitness. Rather, the proper setting is evolutionary game theory [...]</BLOCKQUOTE><P>
I think this is a case of expecting too much from the concept of maximization. The idea of maximization does not imply that all parties reach the maximum they are seeking. It doesn't even imply that they increase the value they are trying to maximize. A hill-climber might be climbing a hill on a mountain that is sinking into the sea. Their elevation may decrease until it reaches zero and they die. This is quite consistent with the idea of maximization.<P>
Gintis apparently claims that frequency-dependence invalidates the idea of maximization of inclusive fitness:<P>
<BLOCKQUOTE>If the genome’s success is based on a pattern of cooperation, promotion, and suppression across loci, which will occur, for instance, if the
production of a protein, RNA sequence, or social behavior requires the collaborative activity of many genes (Noble 2011), or if there are frequency dependent social interactions among individuals in a social species (Maynard Smith 1982), then neither genes nor individuals can be characterized as maximizing inclusive fitness. The conditions under which a population genetics model of gene flow implies fitness maximization at the gene or individual level has been carefully explored (Grafen 1999, 2002, 2006; Metz et al. 2008; Gardner and Wild 2011; Gardner West and Wild 2011). With frequency independence, they affirm the maximization hypothesis. With frequency dependence, the hypothesis it is in general false, and no careful researcher has ever claimed otherwise.</BLOCKQUOTE><P>
Frequency dependence typically means that genes are spending time in environments that differ from the average of the environments they evolved in. It's a case of adaptive lag - where organisms are adapted not to their current environment, but to a weighted average of ancestral environments. This is a classic case where organisms sometimes behave sub-optimally in ways that don't further their own interests.<P>
In such cases, organisms don't always act to maximize their inclusive fitness - but rather act to maximize their inclusive fitness under the hypothesis that they are in an environment that is like the one that their ancestors evolved in. Of course, that hypothesis <EM>might</EM> be mistaken.<P>
In fact, sufficiently sophisticated creatures might be able to evolve a superior strategy - where they sample the frequency involved and change their behaviour adaptively, based on the observed frequency. So, what frequency dependence often boils down to is that organisms can sometimes fail to maximise their inclusive fitness - because they have cognitive limitations and make mistakes.<P>
In this case, I feel that Gintis is expecting too much from inclusive fitness theory. It doesn't claim that all creatures maximize inclusive fitness <EM>perfectly</EM>. Creatures have all kinds of imperfections and limitations that prevent them from acting as <EM>perfect</EM> maximizers.<P>
Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com0tag:blogger.com,1999:blog-6137515683316167603.post-16472129412457251902014-12-29T18:58:00.001-08:002014-12-29T19:28:31.403-08:00More fallout from the 2010 Nowak, Tarnita and Wilson kin selection meltdown<img src="http://www.scientificamerican.com/sciam/cache/file/FD4C7E63-86E1-4CC2-A6BC887D2B6CB9AE.jpg" align=right width=240>Fallout from the 2010 Nowak, Tarnita and Wilson kin selection meltdown continues to rain down. Earlier this year we had <a href="http://www.pnas.org/content/111/35/12585.full">this</a> (from Wilson and Nowak):<P>
<BLOCKQUOTE>
<STRONG>Inadequacy of Inclusive Fitness and Beyond</STRONG><P>
The evolution of social insects often is presented as a testing ground for inclusive fitness theory. It has been claimed that inclusive fitness can explain sex allocation, worker policing, conflict resolution, and evolution of eusociality (14), but precise calculations of inclusive fitness do not exist for any of these phenomena. Relatedness-based arguments, such as the monogamy window hypothesis, are not necessarily wrong but rarely provide a complete picture; moreover, one cannot rely on inclusive fitness to determine when they are correct. The failure of inclusive fitness theory to provide exact calculations is not surprising, because a mathematically meaningful approach to inclusive fitness (72) cannot be performed for the majority of evolutionary processes (5), and the linear regression method (73⇓–75) does not provide meaningful insights and cannot make empirical predictions (76). In general it is not possible to study social evolution from the perspective of an individual by evoking the virtual quantity of inclusive fitness. Instead we should focus on how natural selection acts on alleles that modify social behavior. On the level of genes or alleles, there is no inclusive fitness: Mathematical descriptions of the evolutionary dynamics of genetic mutations do not require a partition of fitness effects (which usually is impossible anyway) or any other aspect of inclusive fitness theory.</BLOCKQUOTE><P>
These folk have a bee in their bonnett. I - and many other scientists - think it is a stupid one. For homework, I think these authors should write an article explaining - at undergraduate level - why kin selection has been as successful as it has been - including when and why it is useful. At the moment, it doesn't look as though they are clear on these topics. The <a href="http://kinselections.blogspot.com/2012/09/enter-george-price.html">Price equation</a> can't be used to make empirical predictions? It might be funny if it wasn't so silly and sad. If you don't have a sympathetic understanding of a topic, you are often in a poor position to criticise it. You wind up attacking straw men of your own making.<P>
IMO, probably the main lesson here for other scientists is the value of humility in science. If you are overconfident, nail your flag to the mast and then dig in then it is easy to wind up making a fool out of yourself.<P>
Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com0tag:blogger.com,1999:blog-6137515683316167603.post-90538392044806834462014-12-27T20:18:00.002-08:002014-12-27T20:19:49.134-08:00Kin or group selection: which is more confusing<img src="http://vignette4.wikia.nocookie.net/uncyclopedia/images/3/3f/So_Confusing.jpg" align=right width=266>The last decade has seen a bit of a shake out in the domain of kin selection and
group selection.<P>
Until recently it was possible to argue that the group selection advocates were
<EM>consistently</EM> more confused about social evolution than users of kin
selection. However as group selection advocates learned more about their
topic some of them gradually started making sense - and some of them now
hold fairly reasonable positions.<P>
Also, it has become clear that <EM>some</EM> of the <EM>opponents</EM> of group
selection are very confused about the whole topic. Steven Pinker wrote a
<a href="http://edge.org/conversation/the-false-allure-of-group-selection">fairly
embarrassing article on the topic</a> in 2012 - and some of that article's
commentators made similarly embarrassing follow-ups.<P>
Popular blogger <a href="http://whyevolutionistrue.wordpress.com/">Jerry Coyne</a>
has written a string of articles about group selection. He doesn't seem to have
made much effort to understand what advocates of group selection are saying - and
so produces mostly straw man attacks.<P>
Richard Dawkins isn't exactly helping either. For example, he
<a href="http://edge.org/conversation/the-false-allure-of-group-selection">writes</a>:<P>
<BLOCKQUOTE>Is a group a replicator? No. We do not have a 'group pool', a metapopulation
in which some groups are more successful than others at making replicas of
themselves, replicas that persist through geological time.</BLOCKQUOTE>
This seems like classic <a href="http://on-memetics.blogspot.com/2011/12/replicator-rot.html">replicator rot</a>
to me. Try talking about copying and heredity instead, and we do indeed have collections of groups,
some of which are better at making copies of themselves than others. Maybe none of them are
<EM>especially</EM> brilliant at making copies of themselves: but so what?<P>
The broad equivalence between kin and group selection makes the issue of which
framework causes more confusion into a significant issue. One of the main problems
with group selection historically has not been that it's wrong, but that it is
confusing and easy to mis-apply.<P>
The confusion by the group selection <EM>opponents</EM> is unfortunate. It doesn't help
to make the case that kin selection is less confusing and less subject to abuse.<P>
However, I think it is still dwarfed by the confusion related to group selection.
Martin Nowak and E.O. Wilson are perhaps the most prominent examples - but it seems
to me that they represent only the tip of a pretty substantial iceberg.<P>
Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com0tag:blogger.com,1999:blog-6137515683316167603.post-49400237534465378192014-12-27T17:08:00.000-08:002014-12-28T15:32:36.977-08:00Kin selection and its critics<img src="http://www.ebi.ac.uk/sites/ebi.ac.uk/files/shared/images/Events/OKASHA_Samir.png" align=right width=250>Here's an interesting new paper: <a href="http://bioscience.oxfordjournals.org/content/early/2014/12/12/biosci.biu196.short?rss=1">Kin Selection and Its Critics</a> - Jonathan Birch and Samir Okasha.<P>
Samir Okasha wrote a rather flawed book on group selection some years ago. However, this article shows that he is doing a good job of keeping up with developments in the field - and it isn't so easy to find significant mistakes in this large recent article.<P>
I think one problem is that it takes the work of Nowak and Wilson a bit too seriously.<P>
In one place the authors argue <EM>against</EM> <a href="http://kinselections.blogspot.com/2012/09/group-selection-and-kin-selection.html">equivalence</a>, saying:<P>
<BLOCKQUOTE>In one respect, the kin selection approach is arguably more general than the multilevel approach, because the latter requires that individuals be nested into nonoverlapping groups, as in figure 4; this is necessary for the decomposition technique in box 2 to apply (Hamilton 1975, Okasha 2006, Frank 2013). Groups of this sort exist in some taxa (e.g., the colonies of many social insect species). But in other cases, individuals engage in social interactions with their conspecifics, but there are no well-defined, discrete groups. The kin selection approach can handle such cases easily; indicative of this is that in deriving equation 4 above (box 1), we did not make use of the fact that the individuals were nested into nonoverlapping groups. Therfore, the claim that kin and multilevel selection are formally equivalent requires at least this qualification. </BLOCKQUOTE><P>
This doesn't seem like <EM>too</EM> much of a stumbling block to me. The modern "group selection" approaches depend critically on defining a "group" to include any collection of organisms - no matter how fleeting or ephemeral. You have to buy into this conception of a "group" for the approach to be worth considering in the first place.<P>
The authors say:<P>
<BLOCKQUOTE>The widespread preference for kin selection may be partly due to multilevel selection's association with the flawed good-of-the-group tradition of the 1950s and 1960s and the associated superorganism concept, of which many biologists remain suspicious. It is undeniable that the careless appeal to group-level advantage as a way of explaining a trait's evolution led to serious errors in the past, so biologists’ wariness of this mode of explanation is understandable.</BLOCKQUOTE><P>
That's about the size of it. However, this paints group selection's problems as being in the past. I think that this is inaccurate. A fairly cursory look at the evolutionary social sciences shows that misapplication of group selection is still widespread.<P>
The essay closes with a plea for "causal aptness": use kin selection when you have relatives, use group selection when you have interacting groups. This proposal <EM>sounds</EM> reasonable - but I think it would do little to stem the existing misuse of group selection. The problem is that people see differential group reproduction, reach for group selection, and produce just-so stories about how group traits are the product of differential group extinction or reproduction. This is a systematically bad methodology that use of group selection directly encourages. Using "causal aptness" would probably boost usage of group selection. That seems as though it is likely to cause a range of negative outcomes associated with the misuse of group selection - and so I regard the proposal as suspect.<P>
"Causal aptness" is one proposal. A big health warning relating to the misuses of group selection is another. I think that, if you adopt the first proposal, you should also adopt the second one.<P>
Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com0tag:blogger.com,1999:blog-6137515683316167603.post-22108811249719384502014-10-31T03:27:00.001-07:002014-12-27T06:42:26.738-08:00Disagree with Steve FrankI read Steve Frank's paper <a href="http://stevefrank.org/reprints-pdf/13RDA.pdf">A new theory of cooperation</a> recently. Steve Frank is an expert, but I thought that this paper was <EM>mostly</EM> wrong. The theme of the paper is that suppression of competition within groups represents a new theory of cooperation - that beyond kin selection and reciprocity. He credits the development of the idea to Richard Alexander - in <a href="http://smile.amazon.com/Biology-Moral-Systems-Foundations-Behavior/dp/0202011747/">The Biology of Moral Systems</a>.<P>
Many of the examples of suppression of competition within groups Steve gives are due to kin selection. Steve disagrees with this, writing:<P>
<BLOCKQUOTE>[...] the main weakness of the theory was also apparent. Extensive cooperation occurs between nonrelatives. Different genes in genomes are functionally integrated but not related. Larger human societies often have many highly cooperative but distantly related individuals. Some of this cooperation between nonkin can be explained by extensions of reciprocity to a general notion of mutual benefit for interacting partners (West Eberhard 1975).<P>
In the early 1980s, kin selection plus these extended notions of reciprocity were the main conceptual tools. Those limited conceptual tools led to blind spots about unsolved problems. Only rather forced theories of mutualism could work for the nearly complete integration of genes into cooperative genomes. Only a very enthusiastic belief in the scope of reciprocity could explain the broad social integration in larger groups of weakly related human</BLOCKQUOTE><P>
However, cooperation between groups of "unrelated humans" that is not due to reciprocity has turned out to be <EM>largely</EM> associated with <a href="http://on-memetics.blogspot.com/2014/08/cultural-kin-selection.html">cultural kin selection</a>. This is kin selection applied to memes - not genes. Many of Steve Frank's examples fit onto the familiar kin selection / group selection axis - though he apparently doesn't fully realise this.<P>
There is indeed <EM>another</EM> force that produces cooperation besides reciprocity and kin selection - and that is "manipulation". Manipulation is where agents impose their wills on other agents. Teams sometimes cooperate because they are cooerced into doing so by supervisor figures. Manipulation helps to explain suppression of competition in cooperative genomes. The efforts of individual genes to bypass meiosis is thwarted by the "parliament of genes" in the genome. Manipulation isn't the same as kin selection or reciprocity - so Steve Frank's paper is <EM>partly</EM> correct.<P>
However, I think that there's a good reason to call this "manipulation" and not "suppression of competition". "Suppression of competition" is just another way of saying "cooperation". Explaining cooperation in terms of "suppression of competition" seems pretty circular to me.<P>
Manipulation is responsible for <a href="http://on-memetics.blogspot.com/2014/07/tim-tyler-eusociality-symbiont.html">the symbiont hypothesis of eusociality</a>. This was proposed in 1934. The "parliament of genes" phrase comes from Leigh (1971). Since manipulation is such an obvious and well-known phenomenon, there may well be earlier examples of it being involved to explain cooperative behaviour. As a theory of cooperation, the idea is not <EM>exactly</EM> new - and I'm pretty sure that Richard Alexander wasn't responsible for it.<P>
In <EM>The Biology of Moral Systems</EM>, Richard Alexander wrote:<P>
<BLOCKQUOTE>It is a common error to suppose that something additional to nepotism and reciprocity is required to account for the structure of society. (p.153)</BLOCKQUOTE>
Here it sounds as though he repudiates this particular revolution.<P>
Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com0tag:blogger.com,1999:blog-6137515683316167603.post-75577505092560441882014-10-19T14:18:00.000-07:002014-10-19T14:18:39.983-07:00West and Gardner on kin vs group selectionWest and Gardner have not been shy about pointing out the virtues of kin selection over group selection. Here they are with <a href="http://www.zoo.ox.ac.uk/group/gardner/publications/WestGardner_2013.pdf">a summary in 2013</a>:<P>
<BLOCKQUOTE>The most frequently used methods are neighbour-modulated and inclusive fitness. In particular, modern neighbour-modulated fitness methods allow the modeller to go from the underlying biology to an expression or fitness, in a way that facilitates the development of relatively general models [8,27,28,35,37]. In contrast, the group selection approach is used relatively little for modelling specific traits partly because as soon as one moves away from the simplest, most abstract models, and wants to add in real world biology, it often becomes analytically intractable — for example, when populations are structured into different classes of individual, according to sex, age, caste or ploidy [38–40].</BLOCKQUOTE>
I know these folk also have plenty of other objections to group selection. Here, forced to pick one, they went for analytic tractablility. It seems like an odd choice to me. As a veteran computer modeller, analytic tractablility comes relatively low on my list of virtues. I would probably list group selection's association with junk science as my number one complaint.<P>
Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com0tag:blogger.com,1999:blog-6137515683316167603.post-14647399078733386662014-10-18T04:09:00.001-07:002014-10-18T04:32:27.613-07:00Identity by descent: a confusing concept<IMG src="http://upload.wikimedia.org/wikipedia/commons/thumb/0/0c/Pedigree%2C_recombination_and_resulting_IBD_segments%2C_schematic_representation.png/440px-Pedigree%2C_recombination_and_resulting_IBD_segments%2C_schematic_representation.png" align=right width=266>There's a lot of discussion of kin selection using the term "identical by descent" and "identical by state". The idea
of "identical by descent" is that genes are shared as a result of direct descent from a common ancestor - without recombination or mutation. "Identical by state" just means that the DNA sequence is shared. It is said that 50% of their genes with their daughters "IBD".<P>
I think the "identical by descent" terminology is confusing and not useful. In biology, if genes are identical, they are practically always identical through being copied from one (or more) shared ancestors. Mothers share more than 50% of their genes with their daughters - due to genes that have reached fixation, inbreeding and so on. However they still share these gene sequences due to descent from shared ancestors. As to a gene mutating into another form and then mutating back again. If you do the sums, for a gene of any reasonable size this rapidly becomes ridiculously unlikely. There's too much scope for neutral mutations elsewhere. In practice, when genes are identical, the odds are enormously in favour of this being due to shared ancestry. The idea that recombination with an identical gene makes genes not "identical by descent" is an awful one. They are still "identical by descent" - just descent from various different ancestors.<P>
You can't say that mothers share 50% of their genes with their daughters "IBD". It is confusing and mistaken. If you want to use the 50% figure, you have to find another reason for doing so.<P>
Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com0tag:blogger.com,1999:blog-6137515683316167603.post-67954338411006961442014-10-17T19:01:00.000-07:002014-10-18T03:43:31.748-07:00Analysis of "group selection and inclusive fitness are not equivalent"<img src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEixS8GFFIMsT3_POcJc4LSWUCWEpkFtmTlAlj4-I9iCPvIzGxi7ekHVzb2Ml3Mxw_l0INoz1A7ft0tYGxgPTBrdbNX9tDAhts1rAXRclK7iTOJoBEEHDwXAiCc84Xf0lfD25e8reHQkYBQp/s1600/Slide1.jpg" align=right width=222>I briefly analyzed the paper:
<BLOCKQUOTE><a href="http://garciajulian.com/preprints/08.pdf">Group selection and inclusive fitness are
not equivalent; the Price equation vs. models and statistics</a> by Matthijs van Veelen, Julian Garcia, Maurice W. Sabelis, Martijn Egas.</BLOCKQUOTE><P>
This is one of the papers mentioned on my <a href="http://kinselections.blogspot.com/2012/09/equivalence-naysayers.html">equivalence naysayers</a> page.<P>
The paper claims that group selection and kin selection are not equivalent. It argues that inclusive fitness requires fitnesses to be "additive". <EM>Additive fitness</EM> is a common assumption when deriving Hamilton's rule - and is <EM>indeed</EM> associated with inclusive fitness. However, inclusive fitness is a simplified model of kin selection. Kin selection enthusiasts are not too impressed with such critiques - the limitations of inclusive fitness are well known. The paper uses "kin selection" and "inclusive fitness" as though these concepts are interchangeable. I think this is not all that useful an approach.<P>
Kin selection doesn't depend on fitness being additive. That idea is associated with inclusive fitness and Hamilton's rule. These are concepts associated with <EM>simplified</EM> models of kin selection.<P>
Hamilton advocated using "inclusive fitness" instead of "kin selection". Hamilton (1975) "Innate social aptitudes of man" says:<P>
<BLOCKQUOTE>The usefulness of the ‘inclusive fitness’ approach to social behaviour (i.e. an approach using criteria like (b K-k) > 0) is more general than the ‘group selection’, ‘kin selection’, or ‘reciprocal altruism’ approaches.</BLOCKQUOTE><P>
However, I think the pendulum has swung away from "inclusive fitness" and back towards "kin selection" as the term of choice. That's what Gardner and West use tend to use, for example. I'm with them.<P>
Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com0tag:blogger.com,1999:blog-6137515683316167603.post-88895927373671227222014-10-12T07:26:00.001-07:002014-10-12T10:53:55.192-07:00Steven Frank on why kin selection beats group selection<img src="http://www.faculty.uci.edu/img/faculty/2115.jpg" align=right>Here's Steven Frank <a href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3258362/">writing <EM>fairly</EM> recently</a> - on why kin selection is better than group selection:<P>
<BLOCKQUOTE>In more complicated biological problems, it often becomes difficult to express all of the selective forces in terms of relative variances among groups. The problem is that patterns of interaction may differ with respect to different processes, such as mating, competition between certain individuals such as males, and competition between other individuals such as females. In that sort of realistic scenario, it is far easier to trace pathways of causation through a series of partial correlations that can be interpreted as an extended form of kin selection analysis (Frank, 1986, 1998). In practice, it is rarely sensible to express such multiple pathways of causation by expressions of relative amounts of variance among groups, although such expressions may be possible mathematically. For that reason, kin selection often becomes a more natural form of analysis for realistic biological problems, leading to a generalized path analysis framework.</BLOCKQUOTE><P>
I think this is part of it. The idea of a "trait group" represents an attempt to overcome this issue. However there are other reasons too:<P>
<UL>
<LI>Kin selection is more quantification-friendly - with its r, b and c;
<LI>In practice, use of group selection often seems to lead to <EM>naive group selection</EM>;
<LI>Group selection <a href="http://kinselections.blogspot.com/2014/08/how-kin-selection-pushed-group.html">avoids problems where kin selection is obviously responsible</a>;
</UL>
These problems have also led to issues associated with scientific status. Group selection is promoted by scientific rebels who try and pick holes in established views. That appeals to some - while repulsing others.<P>
Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com0tag:blogger.com,1999:blog-6137515683316167603.post-82558382778807457322014-10-03T04:32:00.001-07:002014-10-29T03:18:37.842-07:00Confusion about the topic in the popular press<img src="https://farm2.staticflickr.com/1208/621255143_5477c30f7f.jpg" align=right width=300>The popular presentation of the kin selection vs group selection affair still seems to be very confused. For example, consider the coverage of a recent study by some group selection advocates: "<a href="http://www.simonsfoundation.org/quanta/20141002-in-social-spiders-evidence-that-groups-evolve/">Elusive Form of Evolution Seen in Spiders</a>". The article says:<P>
<BLOCKQUOTE>According to one model, known as kin selection, highly related organisms such as bees and ants can develop altruistic behavior — for example, many females forgo reproduction in order to raise the queen’s brood — because they will still pass down their genes indirectly, through the queen. But despite its altruistic appearance, kin selection is selfish — it helps an individual’s genes to survive. Can natural selection promote truly unselfish traits, behaviors that are good for the group, but not necessarily to the benefit of individuals (or their immediate kin)? Some evolutionary models predict that it can [...]</BLOCKQUOTE><P>
The rest of the article is all about group selection. Can kin selection be selfish while group selection is altruistic? Not according to the modern scientific consensus on the topic - that has <EM>kin selection</EM> and <EM>group selection</EM> being different accounting methods that attempt to account for the way in which genes propagate in viscous populations. Modern versions of these theories make <a href="http://kinselections.blogspot.com/2012/09/group-selection-and-kin-selection.html">the same predictions</a>. <P>
<EM>Originally</EM>, group selection was <EM>widely</EM> imagined as taking place <EM>between</EM> demes. However Williams, Maynard Smith, Dawkins and many others pointed out that this sort of "group selection" didn't work very well - individual-level adaptations swamped the group-level ones. Much later, the "groups" of group selection were re-imagined as involving <EM>any</EM> kind of social cohesion, including - critically - family groups consisting of mothers and their offspring - or groups of siblings. Individuals are then modeled as being part of an enormous number of partly-overlapping groups - including, critically, close family groups. With this radical readjustment, this new form of group selection - if properly applied - reproduces the predictions of kin selection exactly.<P>
Most group selection advocates have now (finally) come on board with the scientific consensus on kin selection that was established in the 1970s. They recognize they these two theories - if properly applied - produce the same predictions and results. However, you wouldn't guess at this from this popular article. It's promoting the idea of group selection as a novel, revolutionary theory. This conception of group selection is not scientifically accurate.<P>
<a href="http://guardianlv.com/2014/10/group-selection-in-social-spiders/">Another article</a> promoting the study has a different take on it: group selection vs gene selection. Their champions of "gene selection" are Pinker and Dawkins. It should <EM>really</EM> be group selection vs kin selection. Genes (in the broad sense of heritable information) underlie both.<P>
More coverage:
<UL>
<LI><a href="http://www.techtimes.com/articles/16958/20141003/is-group-selection-a-thing-spiders-support-an-evolutionary-revolutionary-idea.htm">Is 'group selection' a thing? Spiders support an evolutionary, revolutionary idea</a> - Jim Algar
<LI><a href="http://guardianlv.com/2014/10/the-silken-reef-social-spiders-and-evolution-by-group-selection/">The Silken Reef: Social Spiders and Evolution by Group Selection</a> - Michael Schultheiss
</UL>
<HR>
Update 2014-10-29:
Goodnight has <a href="https://blog.uvm.edu/cgoodnig/2014/10/22/spider-group-selection/">weighed in on his spider study on his blog</a>. <a href="http://en.wikipedia.org/wiki/Kin_selection">Kin selection</a> and <a href="http://kinselections.blogspot.com/2012/09/group-selection-and-kin-selection.html">equivalence</a> are apparently unmentionable topics. To me this seems like a parallel universe.<P>
Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com1tag:blogger.com,1999:blog-6137515683316167603.post-53017932269176677972014-09-27T03:09:00.001-07:002014-10-03T18:00:35.469-07:00Cultural group selection - bibliographyI have previously posted bibliographies of <a href="http://kinselections.blogspot.com/2013/02/cultural-kin-selection-bibliography.html">cultural kin selection</a> and the closely-related topic of <a href="http://kinselections.blogspot.com/2012/09/tag-based-cooperation.html">tag-based cooperation</a>. However some academics have been studying the same topic under a different name and using different models and approaches.<P>
Most bibliographies of <a href="http://en.wikipedia.org/wiki/Cultural_group_selection">cultural group selection</a> seem to be in pre-internet document formats: devoid of links. Here's an HTML <EM>linkified</EM> one:<P>
<UL>
<LI>Andre, J.B. and Morin, O. (2011) <a href="http://jb.homepage.free.fr/HomepageJB/publications_files/Andre_Morin_2011.pdf">Questioning the cultural evolution of altruism</a>.
<LI>Bell, Adrianm Richerson, Peter J. and McElreath, Richard (2009) <a href="http://www.pnas.org/content/106/42/17671.abstract">Culture rather than genes provides greater scope for the evolution of large-scale human prosociality</a>.
<LI>Boyd, R. and Richerson, Peter J. (2010) <a href="http://rstb.royalsocietypublishing.org/content/365/1559/3787.full.pdf">Transmission coupling mechanisms: cultural group selection</a>.
<LI>Boyd, R., Richerson, Peter J. (2002) <a href="http://www.sscnet.ucla.edu/anthro/faculty/boyd/BoydRichersonJTB2002_Clean.pdf">Group beneficial norms can spread rapidly in a structured population</a>.
<LI>Boyd, R., Richerson, Peter J. (2009) <a href="http://rstb.royalsocietypublishing.org/content/364/1533/3281.full.pdf">Culture and the evolution of human cooperation</a>.
<LI>Boyd, R. & Richerson, Peter J. (1985) <a href="http://smile.amazon.com/Culture-Evolutionary-Process-Robert-Boyd/dp/0226069338/">Culture and the Evolutionary Process</a>.
<LI>Cullen, Ben R. S. (1995) <a href="http://www.academicroom.com/article/cultural-group-selection">On Cultural Group Selection</a>.
<LI>De Smedt, Johan and De Cruz, Helen (2012) <a href="http://users.ox.ac.uk/~some3056/docs/DeSmedt_DeCruz_PBB.pdf">Human Artistic Behaviour: Adaptation, Byproduct, or Cultural Group Selection?</a>
<LI>Efferson, Charles, Rafael Lalive, and Ernst Fehr (2008) <a href="http://www.sciencemag.org/content/321/5897/1844.short">The coevolution of cultural groups and ingroup favoritism</a>.
<LI>Henrich, Joseph (2004) <a href="http://www2.unine.ch/files/content/sites/ethol/files/shared/documents/Henrich04.pdf">Cultural group selection, coevolutionary processes and large-scale cooperation</a> (<a href="http://www2.psych.ubc.ca/~henrich/Website/Papers/HenrichplusCommentsJebo2003.pdf">preprint</a>).
<LI>Henrich, Joseph and Boyd, R. (1998) <a href="http://www2.psych.ubc.ca/~henrich/Website/Papers/H&B98clean.pdf">The Evolution of Conformist Transmission and the Emergence of Between-Group Differences</a>
<LI>Molleman, L., Quinones, A.E. & Weissing, F.J. (2013) <a href="http://www.rug.nl/research/theoretical-biology/_pdf/mo_ea_ehb13.pdf">Cultural evolution of cooperation: the interplay between forms of social learning and group selection</a>
<LI>Richerson, Peter J. & Boyd, R. (1990) <a href="http://www.sscnet.ucla.edu/anthro/faculty/boyd/BoydRichersonJTB90.pdf">Group selection among alternative evolutionarily stable strategies</a>.
<LI>Richerson, Peter J. & Boyd, R. (1999) <a href="http://www.des.ucdavis.edu/faculty/Richerson/complex.PDF">The evolutionary dynamics of a crude super organism</a>.
<LI>Richerson, Peter J. & Boyd, R. (2008) <a href='http://xcelab.net/rm/wp-content/uploads/2008/10/boyd-richerson-human-altruism-chapter.pdf'>Solving the Puzzle of Human Cooperation</a>
<LI>Soltis, J., Boyd, R., Richerson, Peter J. (1995) <a href="http://www.sscnet.ucla.edu/anthro/faculty/boyd/SoltisBoydRichersonCA95.pdf">Can group-functional behaviors evolve by cultural group selection? An empirical test</a>.
<LI>Steele, D. R. <a href="https://mises.org/journals/jls/8_2/8_2_1.pdf">Hayek's Theory of Cultural Group Selection</a>.
<LI>Zimmerman, Matt, McElreath, Richard and Richerson, Peter J. (2012) <a href="http://www.des.ucdavis.edu/faculty/Richerson/Cooperation%20Evolution%20of.pdf">Evolution of Cooperation</a>.
</UL>
Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com0tag:blogger.com,1999:blog-6137515683316167603.post-4322065927386381902014-09-01T12:04:00.001-07:002014-09-01T12:04:48.486-07:00Group selection is surprisingly popularSome statistics follow:<P>
Google trends:<P>
<script type="text/javascript" src="//www.google.com/trends/embed.js?hl=en-US&q=group+selection,+kin+selection,+inclusive+fitness&cmpt=q&content=1&cid=TIMESERIES_GRAPH_0&export=5&w=800&h=330"></script><P>
Google NGRAM:<P>
<iframe name="ngram_chart" src="https://books.google.com/ngrams/interactive_chart?content=group+selection%2Ckin+selection%2Cinclusive+fitness&year_start=1960&year_end=2008&corpus=15&smoothing=0&share=&direct_url=t1%3B%2Cgroup%20selection%3B%2Cc0%3B.t1%3B%2Ckin%20selection%3B%2Cc0%3B.t1%3B%2Cinclusive%20fitness%3B%2Cc0" width=800 height=360 marginwidth=0 marginheight=0 hspace=0 vspace=0 frameborder=0 scrolling=no></iframe><P>
Google scholar:<P>
<UL>
<LI>"group selection": 66,000;
<LI>"kin selection": 29,300;
<LI>"inclusive fitness": 21,500;
</UL><P>
Even if you add together the "kin selection" and "inclusive fitness" volumes, "group selection" often comes out on top.<P>
Group selection doesn't look <EM>particularly</EM> down-trodden - from this data.<P>
Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com0tag:blogger.com,1999:blog-6137515683316167603.post-2354167768049686902014-08-31T17:08:00.001-07:002014-08-31T17:27:03.263-07:00How kin selection pushed group selection into the scientific fringes<img src="http://cdn.pearltrees.com/s/pic/th/science-wikipedia-encyclopedia-14148457" align=right width=280>Group selection lost out to kin selection in the 1960s and 1970s as
an explanation for cooperative behaviour. Group selection went on to
lead an underground existence - in which a few remaining proponents
sought out ways to find things which group selection explained which
kin selection did not.<P>
We now know that
<a href="http://kinselections.blogspot.com/2012/09/group-selection-and-kin-selection.html">modern
versions of these theories make the same predictions</a>. So there isn't really anything
that group selection can explain that kin selection cannot. However this didn't
stop the group selection proponents from trying to find these types of phenomena.
Indeed, <EM>if</EM> they had succeeded, fame and glory would have awaited them.<P>
This quest took group selection to the borderlands of science. Kin selection was
uncontroversially used to explain
<a href="http://kinselections.blogspot.com/2014/06/obvious-types-of-kin-selection.html">obvious adaptations</a> - such as breast feeding,
parental care and nepotism. Group selection picked more controversial targets -
senescence, the maintenance of sexual recombination and cultural evolution.
The theme of these topics seems to have been avoiding rapid refutation of:
"kin selection explains that without invoking group selection".<P>
This quest now seems to be mostly over. Most of the group selection
enthusiasts have publicly given up their original quest - and have accepted the
equivalence between kin selection and group selection.<P>
However the topics group selection is invoked to explain still
seem to be skewed away from those that kin selection is used to explain.
As a result, group selection is generally offered as an explanation in
areas where the theory doesn't function well. This
seems like a hangover which group selection is still recovering from.<P>
Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com0tag:blogger.com,1999:blog-6137515683316167603.post-44703749710159006222014-08-31T11:54:00.002-07:002014-08-31T11:56:10.524-07:00The "multi-level" terminology<img src="http://www.sequenceinc.com/fraudfiles/wp-content/uploads/2013/03/pyramid-selling-scam.jpg" align=right width=266>This article will claim that "multi-level selection" and "group selection" should not be understood as being synonyms.<P>
It does appear that "multi-level selection" is terminology that was invented to give
group selection a face lift - and allow it to escape from an unglamorous past.<P>
However, group selection is not more or less "multi-level" than kin selection is.
The concepts of "relatedness" and "kin selection" can be applied at multiple
levels too. One can meaningfully discuss the relatednes between cells in an
ant, the relatedness between an ant and its sisters and the relatedness
between entire colonies of ants. This has been understood since the 1970s and
it can be seen as a result of the generality of the Price equation.<P>
These days, group selection enthusiasts often prefer to use the "multi-level selection"
terminology. This article is my explanation of why I rarely follow them - and why I think
it is a confusing gambit. Kin selection is "multi-level" too. Using the term
"multi-level selection" fails to distinguish between kin selection and group
selection.
It is true that kin selection and group selection are equivalent -
in the sense that
<a href="http://kinselections.blogspot.com/2012/09/group-selection-and-kin-selection.html">modern versions of these theories make the same predictions</a>.
However that doesn't mean that proponents use the same methodologies. These
are still topics that are worth distinguishing between. The best way of doing
that is, I think to use the terms "kin selection" and "group selection".<P>
Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com0tag:blogger.com,1999:blog-6137515683316167603.post-67096142705614331652014-07-28T05:18:00.001-07:002014-08-19T02:37:11.196-07:00Group selection enthusiasm still rampant in the social sciences<img src="http://www.dailyworldfacts.com/wp-content/uploads/2012/04/civil-war-fact-civil-war-battle.jpg" align=right width=280>Most
evolutionary biologists got over group selection back in the 1970s.
However, there's one area of biology where it is particularly prominent:
the social social sciences. There's particular enthusiasm for <EM>cultural</EM>
group selection - as though cultural evolution plays by different rules
in this area.<P>
<a href="http://on-memetics.blogspot.com/2014/08/cultural-kin-selection.html">Cultural kin selection</a>
represents an alternative perspective which
makes little mention of selection acting on groups. Instead of humans
being part of innumerable overlapping groups, their genes and memes
are modeled as being <EM>related</EM> to the genes and memes of others.
This perspective has been much more enlightening in the organic realm,
and I think it will prove to be much more enlightening in the cultural
realm as well. The largely-fictional groups of group selection are
just a clumsy and awkward way of viewing the situation - most of the time.
The perspective has a long history of producing dud science.<P>
In most of evolutionary biology, 90% of scientists are using kin selection and 10%
are using group selection. In the social sciences, it's the other way around.
I think that this fairly clearly indicates the existence of a problem.<P>
Why is group selection still so popular among social scientists? For a long
time many of them held out hope that it would prove to be a new theory
of social behaviour. These hopes now appear to have been crushed by
<a href="http://kinselections.blogspot.com/2012/09/group-selection-and-kin-selection.html">the
failure of the theory to make different predictions</a> from the
long-established orthodoxy of kin selection.<P>
Social scientists have a long history of not understanding how evolutionary
biology applied to their subject areas. As a result there's a large
<a href="http://on-memetics.blogspot.com/2013/04/cultural-evolutions-scientific-lag.html">scientific
lag</a> afflicting the study of cultural evolution. Group selection enthusiasm
seems to be clearly one aspect of that: the numerous social scientists involved are stuck back
in the 1960s somewhere, where the virtues of the kin selection perspective were not yet
widely appreciated.<P>
Lastly, group selection in the social sciences seems to have found
<a href="http://kinselections.blogspot.com/2013/02/kin-selections-vs-templeton-foundation.html">extensive
funding</a> from the John Templeton Foundation. In a surreal twist to the final
battle between science and religion, group selection enthusiasm has been funded to
the tune of millions of dollars. That kind of marketing budget can buy a considerable
quantity of confusion.<P>
Tim Tylerhttp://www.blogger.com/profile/06623536372084468307noreply@blogger.com1