Talk:Kin selection

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Contents

[edit] Question

Question: What number of otherwise unrelated human beings should one sacrifice oneself for, according to this theory? The Anome

The question is not very meaningful as kin selection can't be applied to humans quite that easily. I changed 'person' into 'individual' to remove the unnecessary athropocentrism from the text. --- Timo Honkasalo 12:01 Feb 24, 2003 (UTC)

I guess I'm asking 'what is the average relatedness between two genes in human beings otherwise not known to be related?' -- for example, myself and another member of the human race chosen at random? The Anome 12:43 Feb 24, 2003 (UTC)

When calculating the value of r in the kin selection equation, it should be normalised so that the value when compared with oneself is 1, and 0 when compared to an 'outgroup'. It all depends on what you consider the outgroup to be. If you want to know, how large sacrifices a human would to for his relatives, compared to the human race as a whole, the r-value for a random human would 0. If you would want to know how large sacrifices a human would do for his fellow human beings in comparison to a dandelion, the thr r-value for a random human would be larger than 0, probably closer to 1. Usually possible inbreeding and assortative mating have to be also taken into account. Timo Honkasalo

How 'bout a redirect from "Hamilton's rule" (non-existent) to here? It was kind of difficult to find.195.84.83.115 11:01, 20 Apr 2005 (UTC)

[edit] Suspected vandalism

Some time ago an anon's sole contribution was tthe replacement of the "Tiger Salamander" piece with the following one.

for instance most young Plains Spadefoot Toads are detritivorous and congregate with kin but occasionally one will eat a shrimp and then become carnivorous, such individuals live more solitarily and at least when satiated prefer to eat non-kin than kin reducing the damage they might otherwise do to the survivorship of their relatives.

I strongly suspect vandalism here; and reverted this. In any case this anon's replacement destroyed the ogiginal intention to give an example when kin recognition is unnecessary. Therefore even if confirmed (I am not an expert in toads, only in vandals :-), it must be phrased differently. mikka (t) 19:15, 20 September 2005 (UTC)

[edit] Errors ID'd by Nature, to correct

The results of what exactly Nature suggested should be corrected is out... italicize each bullet point once you make the correction. -- user:zanimum

Reviewer: Chris Barnard, Professor of Biology, University of Nottingham, UK.

  • It’s not true to say kin selection was first suggested by Darwin. Darwin certainly recognized the problem posed by altruism, specifically in the form of sterile castes, but his solution owed more to group selection than kin selection. The modern concept of kin selection stems from informal comments by Haldane, later formally refined by Hamilton (but, to be pedantic, Hamilton did not use the term kin selection - he defined inclusive fitness; it was Maynard Smith who coined the term ‘kin selection’)
  • On the gene selection/individual selection issue, this starts off on a better footing than the previous definition, but slips a bit where it says ‘…siblings share 50% of an individual’s genes…etc’ – should say ‘has a 50% chance of sharing the individual’s gene for self-sacrifice etc’
  • In the third para from the bottom, I’d change the text from the second sentence on to: ‘This may be through recognizing some attribute of an individual that correlates with kinship, such as familiarity through having grown up together, or it may come about indirectly; for instance young tiger salamanders avoid cannibalizing other young salamanders and their eggs until they have left their natal vicinity thereby reducing the damage they would do to the survivorship of their kin.’ As it is the text perpetuates distinctions between forms of ‘kin recognition’ that have been shown to be misleading in the critical literature.

Reviewer Marcos Antezana

  • Darwin belongs in the article since he said that the sterile workers boost the "fitness" of the group of ants and he knew very well that this could only mean the reproductive output of the queen since only she can reproduce (duh). by the way, if the ant colony as a whole does not do well the queen's fitness suffers so this *is* group selection. Elliot Sober usng arguments along these lines demonstrated conclusively that kin selection is a form of group selection and that the latter can but need not involve kin selection, so trying to contrast the two is misguided.
  • the pfennig website i proposed as link should be used to improve the salamander example.
  • Haldane was right in his sly comments and he certainly was not thinking about invasion dynamics but rather of himself as owing some respect to his entire genetic endowment. invasion considerations and resistance to invasion could however be included later in the article once the main considerations are clarified.
    • Well kin selection is very different from group selection (ants are a special case). It may be that Darwin did discuss the subject, but clearly he isn't the most important researcher in the field and don't think he belongs in the first sentence. Also can you provide a reference which supports your assertion? Where exactly did he say that? As a side note, the article should probably mention group selection somewhere. Broken S 05:20, 23 December 2005 (UTC)

ce. group/kin selection: you should read sober's book or the whole issue of "biology and philosophy" that was dedicated to the problem after his book was published; ce. darwin: the quote belongs there for simple historical reasons. if you want to challenge the darwin quote in principle is up to you to come up with the goods. the quote was there before it was challenged with the specious "he must have meant group selection" argument. but i did the work for you because i am way too nice ;)

"This difficulty, though appearing insuperable, is lessened, or, as I believe, disappears, when it is remembered that selection may be applied to the family, as well as to the individual, and may thus gain the desired end. Thus, a well-flavoured vegetable is cooked, and the individual is destroyed; but the horticulturist sows seeds of the same stock, and confidently expects to get nearly the same variety; breeders of cattle wish the flesh and fat to be well marbled together; the animal has been slaughtered, but the breeder goes with confidence to the same family." from the The Origin of Species by Charles Darwin. Chapter 7 - Instinct

replace family with queen and you are set. the passage is even better than i thought since it covers also less clean cut situations than the queen/workers case.

  • Thanks for the quote, but I was asking for a reference citing Darwin as an early proponent of kin selection (not a direct reference to Darwin). Still your quote does convince me (I still think he shouldn't be in the first paragraph, though, because exactly what he is talking about is arguable). Also please sign your posts to talk pages by using ~~~~. I have made the Haldane reference an actual quote and cited it. Broken S 04:47, 24 December 2005 (UTC)
  • darwin is talking about the paradox of evolving sterile workers in social ants (since it cannot be via individual selection). the cows killed "for study purposes" are the sterile workers, the herd to go back to is the queen ant. this was very fine by darwin indeed. about the haldane quote: i have seen it quoted differently, i.e., as "*more* than two identical twins, more than etc". knowing a few haldane stories i believe he is more likely to have said "more than". Marcosantezana 05:17, 24 December 2005 (UTC)
    • Agreed, I have seen it quoted many ways also. Both wikiquote and the source I found (a book of quotations) gave it without the "more than". Some sources even say 3 brothers. I think this version is the most authentic, but feel free to change it if you can find a more authoritative source. I've given up on the Darwin issue. It's fine either way, but I still want a parenthetical reference if we are going to defy Nature. Broken S 05:24, 24 December 2005 (UTC)

one could perhaps cite the crucial passages as an introduction. only animus can motivate one to deny darwin his credit for this. the text speaks for itself. no need for exegets that hate the spotted owl. Marcosantezana 05:45, 24 December 2005 (UTC)

    • That's much better Broken S 04:25, 26 December 2005 (UTC)
      • I just removed Darwin's quote again, see my edit summary. BrokenSegue 04:37, 10 February 2006 (UTC)

[edit] Sectionalizing

This article is in desperate need of some sections. No good divisions come to mind. The article as it stands is a mish-mash. How about "History" and "Theory" sections. I'm not sure if that's the best though. Broken S 04:25, 26 December 2005 (UTC)

i agree but it will get better. the problem is that the history explains why people came to care at all about the concept so in a way it's necessary. we will fix it. it's work in progress. Marcosantezana 23:37, 26 December 2005 (UTC)

The introduction should probably have something about how the old group selectionist arguments are overcome by kin selection and how this brings it in line with the overall 'selfish gene' point of view. Shyamal 05:02, 5 January 2006 (UTC)

[edit] Improvements

The 1st sentance needs to be a definiton of kin selection. Also is there a date for hamiltons "first formal quantitative treatment of kin selection "? -Ravedave 21:28, 27 December 2005 (UTC)

  • Well, I added the 1st sentance, I don't know how correct it is. I think it needs to work in that it is a theory... If anyone could help out it would be appreciated. -Ravedave 18:42, 28 December 2005 (UTC)

I think a section or at least reference to haplodiploidy would be worthwhile given its scientific importance in the study of K/S. Secondly, some clarification on the apparent paradox of group selection vs individual benefits and how K/S theory was able to explain it would be useful. williruntus.

[edit] Removed content - worth keeping?

I removed this content by Speh, it might be worthwhile to keep once edited, but it definitly needs a source. Any comments? -Ravedave 21:01, 4 January 2006 (UTC)

--- When the theory of Kin Selection become well established the excitement around it led to its being over used in many cases. People have made the mistake of jumping to the conclusion of kin selection when another explanation is really behind the observation. For example, in elephants babysitting is observed by some lineages, in which older sibs look after their young sibs while the parents are away. Initially the conclusion of kin selection was jumped to, but this is not the case. In fact babysitting in elephants has evolved under system where the parents force the older sibs to babysit their younger sibs. This can be shown mathematically to be a much easier system to evolve, as all of the benefit is confered onto the babysitting gene. If the babysitting gene manifests itself in the offspring, then all of the cost is on that gene - any offspring who has the gene must look after its younger sibs, who may not have that same gene. If the babysitting gene is in the younger sib only, the gene receives no evolutionary benefit. Under parental pressure, all of the benefit is on the babysitting gene, and the benefits of being babysat simply have to outweigh the costs of babysitting.

---

Yes, I think it would benefit from a source, but I don't have time this weekend. My dominant niggle with it is that the relatedness of full sibs to each other is the same as the relatedness with their parents. At least African elephants do not have biparental care, and I do not know to what extent females mate with the same males in successive years. This would make the explanation given by Speh to be more likely to hold, because (half) siblings will be less related to each other than to their mother, but I am worried by the fact that the plural is being used for parents, when only the mother actually cares!
Finally, it must be questioned whether the sample size of the phenomenon being described (Kin Selection become well established the excitement around it led to its being over used in many cases) is actually greater than 1, so I think one would have to be able to cite several papers where this is the case. It is also generally true that new, popular explanatory principles tend to be overused, so I'm not sure that it merits inclusion in this particular article. There's probably even a name for the phenomenon, so it really perhaps belongs somewhere else? - 1/2! - Samsara 17:12, 14 January 2006 (UTC)
To give an example where a new theory is being overused: fluctuating asymmetry. As I said, I don't have time to look it up, but from memory, there are a handful of very implausible papers out there. - Samsara 17:18, 14 January 2006 (UTC)

[edit] The Darwin Quote

I don't want to edit war over this (I follow WP:1RR),but care to explain how that quote is relevant? The Nature review said that the Darwin reference was improper and I'm starting to agree. That sounds like group selection to me. He never once mentions family connections. I'll agree that it's iffy whether or not he's talking about kin selection (he might be- he does say that the seeds would be from the same "stock" which could imply kinship or could just imply same species, he is vague), but the amount of text devoted to darwin is excessive and also POV (ex. "Darwin commented insightfully"). I might agree with keeping it if it served as a particularly good example, but the vegetable example seems more like artificial selection and therefore isn't the most instructive example. No? (P.S. please include more substantive edit summaries, yours wasn't helpful) BrokenSegue 20:21, 10 February 2006 (UTC)


"when it is remembered that selection may be applied to the family, .... " is that the word "family" ? the amount of text is excessive? why? darwin is given credit by everybody who is not a sycophant of hamilton, and this passage is the one everybody including haldane and hamilton has drawn from. by the way, the citation is there after you asked for it. are you bipolar, amnesic, both? who told you that "nature" is holy? read maynard smith's nature article on group vs kin selection and then consider the fitness causation criterion for the level of selection. this will take care of your "is this group selection" point. "same stock": do you think the guy was stupid? we knew he needed heritable differences within the especies, ergo... finally is artifical selection excluded by the first sentence in the "kin selection" article? that is an irrelevant distinction. 05:36, 11 February 2006 (UTC)

Please please. Keep this civil. Did you not see my comment "after some reflection". I have changed my mind. I am neither bipolar nor amnesic. The reference is indeed an improvement. "Darwin is given credit by everybody who is not a sycophant of hamilton" can you find a text that says that? In, "The Ant and the Peacock: Altruism and Sexual Selection from Darwin to Today" by Helena Cronin, the author wrote, "At times, Darwin reads like a group selectionist and at times like an individual selectionist with a kin selection solution...Sadly, it seems, there can be no definitive answer as to what Darwin really had in mind." It's as I said. It isn't clear what he thought. I'm not arguing my point of view is entirely superior, but that it is disputed. In "Adaptation" Michael R Rose wrote, "Darwin's solution was to invoke superordinate family selection." Oh, and in "The Nature of Selection: Evolutionary Theory in Philosophical Focus" Elliott Sober said nearly exactly what I was thinking, "Darwin suggested a group selectionist explanation. But if the tribe were composed of genticaly related individuals..." As to the Maynard Smith article in Nature, did you mean the one titled, "The origin of altruism" [seems to be a book review]? Or another (he has written a lot of articles in Nature)? So, at the vey least the text should be modified to suggest a conlfict or some doubt (I'd also prefer if it would be shrunk considerably, as Darwin is unimportant in the creation and application of kin selection.). BrokenSegue 12:56, 11 February 2006 (UTC)

i apologize for my impatience. you should use your own insight to judge what darwin wrote. sober may be a good philosopher but who told you he must have the last word when reading a biologist? furthermore when sober wrote that he had not yet developed his fitness causation criterion. you should read the book. the nature review of maynard smith shows you on the other hand how biologists have trouble with epistemology. sober's fitness causation criteron is not "disputed" and has not been challeged credibly by anybody. cronin's text is flawed by her ignorance of sober's criterion. finally remember that one of the main advantages of wikipedia is that people with big names cannot simply impose their will. the stuff must make sense or should be rejected. the crucial thing here is that you absorb why sober says that kin selection is group selection. actually after you do that, you should edit the group selection article accordingly, to stress the causation criterion and do so a truly big service to everybody in this bickering group and, more importantly, to the wider wiki reading community. Marcosantezana 17:12, 11 February 2006 (UTC)

Perhaps I didn't make myself clear enough. I'm not saying that it is nonsense to think that Darwin first originated the idea, but I'm saying that it is disputed. Sober might very well be mistaken, but it is one source. I remind you that all facts (and interpretations) must be non-original (WP:NOR). Even if you and I disagreed with the big names, unless you have published some noteworthy work (I haven't), then we can't include our views. I'd say that your position is not backed up by much literature (are you suggesting that Kin selection is equivalent to group selection?). In the next few days I'm going to type up a compromise version and reference it. BrokenSegue 04:41, 12 February 2006 (UTC)

sober concluded that kin selection is group selection in his book (after writing the inconclusive stuff early on that you unearthed). his identification of the causation of fitness differences as the sole criterion to decide on units of selection/level of selection issues is the major advance of the last 50 years in the analytical philosophy and epistemology of natural selection. you shoud read the book before further discussing these complex issues. citing old literature or works written by people who have not done their home work won't cut it. only then you should write your "final version" (and you will be also able to clean up other articles). enjoy  :) Marcosantezana 05:05, 12 February 2006 (UTC)

[edit] Needs a large scale reversion of Marcosantezana's edits

Marcosantezana (talk · contribs · deleted contribs · logs · block user · block log) has caused considerable neutrality difficulties on other evolution related pages, hence Wikipedia:Requests for arbitration/Marcosantezana. And this page seems to be hevely influenced by him. However, the pre-Marcos version contains errors according to natures' reviewer. So a simple minded wholesale revert & reintroduction of abreviated version of new information is impossible; meaning this page needs the attention of an evolutionary biologist. JeffBurdges

This page is in dire need of a biologist. I have been trying to keep tabs on the page as much as I can, but it is over my head. -Ravedave 05:32, 4 April 2006 (UTC)
In context with the struggle that we have with him at the Natural selection page, it will not happen directly. KimvdLinde 13:59, 4 April 2006 (UTC)

[edit] Unequiocal evidence for the operation of Hamilton's Rule in humans

It is unethical to manipulate genetic relatedness in humans. Furthermore, without exception, all studies of Hamilton's rule in humans to this date suffer one common drawback: the evidence is entirely observational, and it is therefore in no case possible to exclude categorically all confounding variables (interaction time, reciprocity etc.), so as to make an unequivocal test of the claim that human altruistic behaviour is modulated by the genetic relatedness, when other factors are held constant.

In the light of this, what would constitue unequiocal evidence for the operation of Hamilton's Rule in humans ? Varga Mila 08:47, 20 May 2006 (UTC)

[edit] On Tiger salamanders and social insects

I have deleted the discussion of tiger salamanders and social insects from the 'mechanisms' section.

1) The salamanders were presented as an example of altruism in the absence of kin recognition; however, the altruism is facultatively expressed only in the presence of kin, so this is very confusing.

2) The discussion of the social insects had no bearing on the *mechanism* of kin selection, as far as I could see.

    • No, social insects are an excellent illustration of some of the mechanisms through which kin selection acts. Perhaps you mean that social insects don't tell us any more about the theory behind this; while this may well be true, it is not true that examples are often far more valuable than reams of theory? For this reason I will be adding some information on kin selecion in the social insects.

[edit] Racism

I have deleted the 'see also' link to Racism, only for it to be reinstated. I am going to delete it again, now. I can see no justification for the link between kin selection and racism. This is not some quibble about the naturalistic fallacy (which states that what 'is' is the same as what 'ought' to be; clearly nonsense). Rather, I don't see how racism as a behaviour is one that is in any way linked to kin selection. If there is a proper evolutionary literature on the origins of racism, and if the argument invokes kin selection, a short discussion of this would make an interesting contribution to this article on kin selection. And once that had been added, it might be worth linking to the racism article.

I added the link. The connecton between these two concepts seems quite obvious to me, allthough I'm terribly biased (but who isn't?). I admit that my understanding is rather limited, but Kin selection,Group selection,Inclusive fitness,Racism,Ethnic nepotism and Altruism seems to be related. I will dig up some sources instead of ranting.

Here's something interesting to work with:

"The National Front was saying something like this, "kin selection provides the basis for favoring your own race as distinct from other races, as a kind of generalization of favoring your own close family as opposed to other individuals." Kin selection doesn't do that! Kin selection favors nepotism towards your own immediate close family. It does not favor a generalization of nepotism towards millions of other people who happen to be the same color as you. Even if it did, and this is a stronger point, I would oppose any suggestion from any group such as the National Front, that whatever occurs in natural selection is therefore morally good or desirable. We come back to this point over and over again. I'm definitely not one who thinks that "is" is the same as "ought." (Richard Dawkins)[1]

Apparently, some political groups have made the connection. (Note the very emotional response by Dawkins.)

Sceptic: How do you evaluate the work of Irena"us Eibl-Eibesfeldt, J.P. Rushton, and Pierre van den Berghe, all of whom have argued that kin selection theory does help explain nationalism and patriotism?
 Dawkins: One could invoke a kind "misfiring" of kin selection if you wanted to in such cases.[...]I could imagine that racist feeling could be a misfiring, not of kin selection but of reproductive isolation mechanisms.[2]

Apparently, some research (controversial, of course) has been made.

I also recall that Dawkins briefly mentioned racism as misfired kin selection in The Selfish Gene.
We'll see where this ends. --CAD6DEE2E8DAD95A (hello!) 17:50, 25 July 2006 (UTC)

[edit] Defenitions

"...In evolutionary biology, kin selection refers to changes in gene frequency across generations that are driven at least in part by interactions between related individuals, and this forms much of the conceptual basis of the theory of social evolution. " Who has formally defined it as such, published in which peer reviewed journal? TongueSpeaker 11:42, 6 May 2007 (UTC)

The concept was formalized by JBS Haldane (1955) and W. D. Hamilton (1963), the actual term "kin selection" was (I think) first applied to the concept by John Maynard Smith (1964) where he says "These processes I will call kin selection and group selection respectively. Kin selection has been discussed by Haldane and by Hamilton. ... By kin selection I mean the evolution of characteristics which favour the survival of close relatives of the affected individual, by processes which do not require any discontinuities in the population breeding structure."
  • Haldane, JBS. 1955. Population Genetics. New Biology 18:34-51
  • Hamilton, WD 1963. The evolution of altruistic behavior. Am. Nat. 97:354-356.
  • Maynard Smith, J. Group Selection and Kin Selection, Nature Vol.201, pp.1145-1147, 1964.
Pete.Hurd 19:58, 20 July 2007 (UTC)


I'm curious: there's no discussion on this page about concerns about the application of kin selection asa scientific construct... Also, does anyone know of an experiment validating kin selection? In the absence of such, can it even qualify as a scientific theory (as opposed to a hypothetical mechanism)?

“behaviors that help a genetic relative are favored by natural selection.”

Sure, but behaviours that help a non-genetic relative are also favoured by natural selection, as in symbiosis. What is favoured is what is beneficial, which may be kin selection, but it may also be something wholly different.

I don't know, I think this article needs some counterpoint for completeness. ##### —Preceding unsigned comment added by 67.187.96.11 (talk) 16:09, 1 April 2008 (UTC)

[edit] Hamilton's rule

I removed this from an HTML in the article:

Technically, the correct definition for relatedness (r) in Hamilton's rule describes it as a regression measure. Regressions, unlike probabilities, can be negative, and so it is possible for individuals to be negatively related, which simply means that two individuals can be less genetically alike than two random ones on average. This has been invoked to explain the evolution of spiteful behaviours. (This passage directly contradicts the definition given in Hamilton's original formulation, which I have here in front of me, and which matches that given in the text above - it is not possible to have a NEGATIVE probability!; this is apparently someone else's definition of "r", and if it is to be included, then it needs to be cited with proper attribution, and contrasted with Hamilton's definition, otherwise it is terribly confusing).

--SallyForth123 21:20, 30 July 2007 (UTC)