inclusive fitness

there’s some amount of confusion out there in the hbd-o-sphere (and beyond!) about inclusive fitness, which is understandable since the concept is not that straightforward — especially for those of us who are not scientists. i thought it’d try to dispel some of that misunderstanding by sharing my layman’s understanding of the concept — i think i grok the idea pretty well now (in a basic sorta way) — hope i don’t add to the confusion!

to start with, inclusive fitness is NOT some sort of biological law that organisms (including humans) will automatically be altruistic towards other individuals with whom they share a lot of genes (or vice versa if vice versa). if you hold that idea — and i get the impression that a lot of people do — get it out of your mind right now! you’ll feel better for it, trust me.

inclusive fitness is simply a concept or model which explains HOW certain social behaviors — especially altruism — might’ve evolved at all. period. full stop.

to understand inclusive fitness, we need to back up a sec first and think about fitness and what that is. very (very!) simply, fitness refers to an organism’s ability to survive and reproduce in a particular environment. traits — including behaviors — that enable an organism to survive and successfully reproduce will be selected for simply because that organism *is* able to survive and reproduce in its environment. this is natural selection. pretty simple, really, darwin’s dangerous idea.

when it comes to certain social behaviors in humans, it’s readily understandable why many of them were selected for. for example, mothers who devote a lot of time and energy to care for their infants — who obviously can’t take care of themselves and would die without any care — will be more fit than those mothers who don’t. the genes that predispose for those behaviors get selected for since children get half of their dna from their mothers, and the ones that are cared for are much more likely to survive.

what was — and to some extent still is — a big mystery is why other sorts of altruistic behaviors were ever selected for even though they hurt an organism’s fitness. how would self-sacrificing altruistic behaviors directed towards non-descendants ever be selected for? for instance, why on earth would somebody feel compelled to run into a burning building to save a neighbor (who wasn’t their child) at great risk to their survival and, therefore, to their fitness? we can see how “genes for altruistic behaviors towards offspring” could be passed down from mother (or parents) to kids, but how were genes for more general altruistic behaviors selected for?

here is where william hamilton‘s absolutely genius idea — inclusive fitness — comes in: perhaps certain social behaviors, which on the surface appear to reduce an organism’s fitness, and so shouldn’t get selected, might’ve been selected for if those behaviors were directed toward other close kin with whom individuals also share much dna in common.

everybody gets half of their dna from each of their two (for now, anyway) parents. but we also share dna with siblings and (blood-related) aunts and uncles and (wait for it…) cousins. given this inheritance pattern, probability says, for instance, that, in a randomly mating population, an individual should share 12.5% of their dna with a first cousin. so, if an individual with certain “genes for altruism” behaves altruistically toward their first cousins, odds are not bad that those first cousins might also have those same “genes for altruism.” here, then, we have a mechanism for how apparently self-sacrificing social behaviors can be selected for: since the altruistic individual 1) aids close kin with whom he shares much of his dna AND 2) probably in many instances shares the same “genes for altruism,” his being altruistic toward those kin 1) does not reduce his fitness AND 2) the “genes for altruism” get selected for, too. mystery solved. (see also: kin selection.)

one way i like to think of inclusive fitness — which, perhaps, isn’t entirely the right way to look at it, but i feel it helps my understanding — is that if you wanted to calculate an individual’s total fitness by adding up how many actual copies of his genes he passed on, you need to add together those found in his offspring and those in his close relatives’ offspring. in other words, you need to add together his own direct fitness plus his close relatives’ fitness to get his inclusive fitness (or his total fitness).

it seems likely that many of the altruistic (or spiteful, etc.) behaviors we’re talking about are pretty general in nature, i.e. not that specific behaviors like “be altruistic to your close kin” were selected for, but rather more like “be altruistic to the people around you, because they’re probably your close kin” were. it remains to be seen how much kin recognition plays a role in altruism in humans, but that’s a topic for another post anyway. for right now, i just wanted to make clear what inclusive fitness is — and isn’t. again, inclusive fitness is a concept which explains HOW altruistic behaviors MIGHT be selected for. it does NOT predict that individuals will DEFINITELY be more altruistic toward those with whom they share much dna.

the whole topic of inclusive fitness is, of course, much more complicated than all that, but i think this is a good basic intro to the concept. hope so, anyway! (^_^)

(note: comments do not require an email. citizens against altruism!)

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92 Comments

  1. No description of inclusive fitness would be complete without Hamilton’s rule (lifted from Wikipedia):

    rB > C

    where

    r = the genetic relatedness of the recipient to the actor, often defined as the probability that a gene picked randomly from each at the same locus is identical by descent.
    B = the additional reproductive benefit gained by the recipient of the altruistic act,
    C = the reproductive cost to the individual performing the act.

    When this is inequality is true, the altruistic trait is selected for.

    This formula, along with this table of the coefficient of relationship between human relatives (also lifted from Wikipedia):

    Degree of
    relationship Relationship Coefficient of
    relationship (r)

    Inbred strain 99%
    0 identical twins; clones 100%[4]
    1 parent-offspring[5] 50% (2^−1)
    2 full siblings 50% (2^−2+2^−2)
    2 3/4 siblings or sibling-cousins 37.5% (2^−2+2⋅2^−4)
    2 grandparent-grandchild 25% (2^−2)
    2 half siblings 25% (2^−2)
    3 aunt/uncle-nephew/niece 25% (2⋅2^−3)
    4 double first cousins 25% (2^−3+2^−3)
    3 great grandparent-great grandchild 12.5% (2^−3)
    4 first cousins 12.5% (2⋅2^−4)
    6 quadruple second cousins 12.5% (8⋅2^−6)
    6 triple second cousins 9.38% (6⋅2^−6)
    4 half-first cousins 6.25% (2^−4)
    5 first cousins once removed 6.25% (2⋅2^−5)
    6 double second cousins 6.25% (4⋅2^−6)
    6 second cousins 3.13% (2^−6+2^−6)
    8 third cousins 0.78% (2⋅2^−8)
    10 fourth cousins 0.20% (2⋅2^−10)[6]

    …demonstrates why “ethnic genetic interests” do not exist. Ethnocentrism cannot evolve via kin-selection because the coefficient of relationship falls off so much once you go past 2nd cousins.

    Also note the relationship difference between first cousins and double first cousins. You see the difference in terms of degree of relationship between outbred and inbred individuals. The higher coefficients of relationships within families in inbred societies make the payoffs from kin altruism that much higher, greatly affecting the selective pressures you encounter, especially once iterated for generations.

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  2. Nice, so if I get this right certain genes (alleles) in an individual might incline him to behave altruistically towards kin (or people he grew up close to) but in the absence of such genes that inclination will not exist. For a social animal like humans this means that it would not be surprising that such alleles exist, though they might be selected out in a society of largely unrelated individuals such as exist in the West. They would be less likely to selected out in clan-based societies for the simple reason that there would be a lot of relatives around to interact with. In fact, there is a reasonable chance that even more or new alleles favoring altruism might be selected for in clan-based societies given enough generations, whereas the tendency would be just the opposite in outbred societies.

    If I have that right there is still the quesition of how many generations does it take to make a big difference in the frequencies of such allelles. What kind of time scales are we talking about. I gather it is measured in centuries based on past posts.

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  3. @ Jayman – “…demonstrates why “ethnic genetic interests” do not exist. Ethnocentrism cannot evolve via kin-selection because the coefficient of relationship falls off so much once you go past 2nd cousins.”

    Aren’t you assuming that you start with a society of unrelated individuals? If a clan is prolific and endogenous, couldn’t it become an ethnic group with high coefficients or relatedness?

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    1. @Luke Lea:

      This would be your word on that. It tells you the average degree of relationship of individuals in each Euro ethny to his co-ethnics. I.e., not enough to make a difference. There’s no group (as far as I know) that’s related to each other as if they were second cousins or greater, which is what you’d need.

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  4. @jayman – “No description of inclusive fitness would be complete without Hamilton’s rule….”

    i was trying to keep the explanation simple and math free…for my fellow non-scientists. (~_^)

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  5. @luke – “Nice, so if I get this right certain genes (alleles) in an individual might incline him to behave altruistically towards kin (or people he grew up close to) but in the absence of such genes that inclination will not exist.”

    sure. all behaviors are heritable to some extent or another, so genes *must* be involved in altruism. there’s no way around that. no genes for altruism, no altruistic behaviors.

    @luke – “…though they might be selected out in a society of largely unrelated individuals such as exist in the West. They would be less likely to selected out in clan-based societies for the simple reason that there would be a lot of relatives around to interact with.”

    yes, and yes.

    @luke – “If I have that right there is still the quesition of how many generations does it take to make a big difference in the frequencies of such allelles. What kind of time scales are we talking about. I gather it is measured in centuries based on past posts.”

    not sure. presumably all sorts of other selection pressures on related behavioral traits would make the matter complicated! can’t see why something like forty generations wouldn’t be enough time for the frequencies of altruism-related alleles to increase (or decrease) significantly.

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  6. @jayman – “Also note the relationship difference between first cousins and double first cousins. You see the difference in terms of degree of relationship between outbred and inbred individuals.”

    yes. and the relatedness can actually be even greater in long-term inbreeding (cousin marrying) populations. the amount of genes shared between some first cousins in saudi arabia and pakistan can be twice as high as what you’d expect in a randomly mating population.

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  7. @ Jayman – “This would be your word on that. It tells you the average degree of relationship of individuals in each Euro ethny to his co-ethnics. I.e., not enough to make a difference.”

    Does that include Ashkenazis? How about Jewish relatedness through the millennia?

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  8. Another possible mechanism for this effect is that certain types of group cohesiveness enhance survivability, such as group defense against outsider attack. Behaviors that favor this type of cooperation and cohesiveness would persist (via genetic inheritance) because more of the group will survive to reproduce.

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  9. @luke – “Does that include Ashkenazis?”

    yes, that includes ashkenazis. *all* jews — not just ashkenazis — are as genetically similar to one another as though they are fourth or fifith cousins. and as jayman said above:

    “Ethnocentrism cannot evolve via kin-selection because the coefficient of relationship falls off so much once you go past 2nd cousins.”

    why? think of it this way. if you do haldane’s calculation — two brothers or eight cousins — the equivalent in second cousins would be 32 of them, for third cousins it would have to be 128, and for fourth, a whopping 512.

    who regularly goes around sacrificing their own fitness in a way that promotes the fitness of 512 of their fourth cousins? not enough individuals for it to make a difference.

    @luke – “How about Jewish relatedness through the millennia?”

    old testament jews appear to have married quite closely (uncle-niece and cousins — they’re the ones who probably introduced father’s brother’s daughter marriage into arabia), so they were most likely more related to one another back then. quite possibly like populations in the middle east today, but that’s just speculation on my part.

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  10. @tom – “Another possible mechanism for this effect is that certain types of group cohesiveness enhance survivability, such as group defense against outsider attack. Behaviors that favor this type of cooperation and cohesiveness would persist (via genetic inheritance) because more of the group will survive to reproduce.”

    there’s no good evidence for group selection in humans, and the population geneticists (like greg cochran) say that the math doesn’t work. also, to date, none of the pro-group selection arguments that i’ve seen have included the math for individual selection — which makes me think that they are probably overlooking it (individual selection).

    see also here:

    “Levels of Selection Are Artefacts of Different Fitness Temporal Measures”

    “I show by comparing the fitness of individuals with that of collectives of individuals in the same environment and over the same period of time – as required to decide if one or more levels of selection is acting in a population – that the selection of collectives is a by-product of selection at the individual level; thus, talking about two or more levels of selection represents merely a different perspective on one and the same process.”

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  11. Doesn’t explain why some people go out of their way to be altruistic to those least related to them, such as adopting orphans from Africa, or giving board and lodging to a dog!

    Darwinists are reduced to “evolution gone wrong” to explain these things.

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    1. @British Nationalist:

      That’s another force entirely: reciprocal altruism. I scratch your back if you scratch mine. The recipients don’t need to be related for such a relationship to develop; heck, they don’t even need to be the same species.

      Generosity is an outgrowth of this: you extend the first favor with the expectation the recipient will one day return it. The problem is that this only works in populations full of reciprocal altruists. Mix reciprocal altruists and kin altruists together, and hijinks ensue.

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  12. @ Jayman – “…demonstrates why “ethnic genetic interests” do not exist. Ethnocentrism cannot evolve via kin-selection because the coefficient of relationship falls off so much once you go past 2nd cousins.”

    Ethnic genetic interests clearly exist and could in theory be calculated precisely. Is it in the genetic interests of Africans if every African was replaced by a Chinese or vice versa? Clearly not.

    I agree that among large ethnies relatedness is too low for ethnocentrism to evolve directly but that’s where gene-culture co-evolution comes in. Whenever there is an interest that is visible to some bright spark – whether it’s St Augustine wanting less conflict or a tribal leader wanting greater defensive cohesion – but the interest is too diffuse for selection to work on its own then cultures develop to bridge the gap.

    And once a culture develops that rewards certain behaviors and punishes others then traits that support those preferred behaviors can evolve.

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  13. @Greying Wanderer:

    “Ethnic genetic interests clearly exist and could in theory be calculated precisely. Is it in the genetic interests of Africans if every African was replaced by a Chinese or vice versa? Clearly not”

    Actually, it doesn’t really matter. Past your 2nd cousins, the fitness impact to you (say mating, for example) is effectively the same if everyone else were Chinese or African. What doesn’t work doesn’t work.

    “And once a culture develops that rewards certain behaviors and punishes others then traits that support those preferred behaviors can evolve.”

    Sure, some other mechanism may be involved. But it’s not kin selection.

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  14. @hbd chick – “there’s no good evidence for group selection in humans”

    My comment is directed at trait evolution in individuals that result in behaviors that operate on group dynamics. For example, a gene mutation that expresses as a proclivity for cooperative behavior (and thereby leads to group cohesion and joint action) would have a reproductive advantage if the group survivability was enhanced. At root, it’s all individual selection, but some enhancements operate through group improvements. See pack animal behaviors as an example, and remember that we started our evolutionary journey as an animal species.

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  15. @BritishNationalist

    “Doesn’t explain why some people go out of their way to be altruistic to those least related to them, such as adopting orphans from Africa, or giving board and lodging to a dog!”

    Bear in mind the scientific and popular definitions of altruism are different.

    The scientific definition is doing things that improve your inclusive fitness so on balance it helps you (at a genetic level) more than it harms you. Altruism is actually selfish even if on the surface it looks selfless.

    The popular western definition is doing good deeds for its own sake.

    So genetically speaking your two examples aren’t examples of altruism. They do exist though so how come?

    .

    “Darwinists are reduced to “evolution gone wrong” to explain these things.”

    There’s nothing wrong with that – although I’d quibble with the phrasing. Evolution selects for a particular environment so it can’t go wrong for that environment but if the environment changes then traits that were beneficial can become harmful. Then it’s a question of whether the organism can evolve for the new environment quickly enough or not.

    .

    “explain these things”

    Part of the reason may be that genetic altruism is double edged. It’s not just being nice to kin it also leads to an incentive to be nasty to non-kin.

    For example if your village of 2nd cousins (3.13%) raids sheep from a village of 4th cousins (0.2%) then as long as it doesn’t harm them 16 times more than it helps you then your village is ahead genetically speaking.

    If so then creating larger co-operating groups might require reducing individual level genetic altruism (by coercion or otherwise).

    Oddly enough if those larger co-operating groups are then as a result of this process capable of building societies where their population dramatically increases then paradoxically the result of a reduction in individual genetic altruism can be genetically altruistic i.e. it could lead to much more of their dna than there was before.

    I think the model is the insurance policy. A large group of people reducing their individual fitness very slightly by say paying tax for an ambulance service actually increases their inclusive fitness through economies of scale.

    I’d say it is quite clear that from c. 1300 at least to c. 1914 ish this process dramatically increased inclusive fitness in NW Europe i.e. evolution didn’t go wrong.

    I’d say it was actually the huge productive capacity of those nations when fully harnessed for warfare during WWI that sparked the cultural unraveling.

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  16. “Darwinists are reduced to “evolution gone wrong” to explain these things.”

    Actually maybe in a sense it is/was evolution going wrong in that if an organism evolves to the point where it is capable of destroying itself then maybe it has gone wrong?

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  17. @Jayman

    “Actually, it doesn’t really matter.”

    It’s the same wherever you draw the boundary: self ~ 2 x brothers ~ 8 x 1st cousins ~ 30-ish x 2nd cousins ~ 500 x 4th cousins ~ whatever.

    self genetic interests
    sibling genetic interests
    1st cousin genetic interests
    etc
    human genetic interests
    carbon based lifeforms genetic interests

    “ethnic” is one boundary.

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  18. Families with psychopathic relatives show us that genetic familiar similarities are less important than phenotypical similarities. When you have families with higher behavior similarities it don’t mean that always will be like that.

    Then, probably, outbred people can have higher intra-familiar behavioral diversity than inbred people obviously because pattern mating.

    Centuries of inbred mating can accumulate and raise genetic similarities. Like, first cousin marriage, 12% of similarity. Heterozygosis reduce 88%??? This children marry other cousin, less heterozygosis… THIS HELP explain amish culture, when ”culture” and ”biology” are near to be 100% correlatives.

    Is like a line chart with ”culture line” and ”behavioral biology line”. In the begin of inbred cultures, the correlation quasi-always will be 0,5 between culture and behavioral biology. In outbred cultures, like modern culture, the correlation will can be 0,1 or less… In the end, the SYNCHRONICITY between culture and behavioral biology will be near to 100% of correlation.

    I think smart people, independent of pattern familiar historical, will tend to be behaviorally outbred or less outbred, look Malala (i love this name, hihihihihihihi)

    By logic, people to do friends with similar ideologies or neurological culture (similar intelligence, personal motivations, degree of sexuality, degree of kindness and/or psychopathy). To understand altruism, i think, we should start by friendship pattern and we already know that people make friends who are similar to him.

    A package behaviors were being selected and not behaviors were selected separately. This explains the extremist mind of human beings, they can not touch the surface of wisdom. Can not decide that the best answer will always be one that encompasses all others like magic cube. Civilizations are not complete accumulation of all the blessings and the elimination of all losses. Because when a population decides that ALL of your culture is wrong, it will not evolve, but only establish the old bio-cultural balance, so if consists of a regression.

    It is as if civilization was an old ford car, which evolves slowly in its improvement. It will still be a ford car.

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  19. @G.W.

    “’Actually, it doesn’t really matter.’

    It’s the same wherever you draw the boundary: self ~ 2 x brothers ~ 8 x 1st cousins ~ 30-ish x 2nd cousins ~ 500 x 4th cousins ~ whatever”

    Not really. With really small relationship coefficients, instances where the number of others will be large enough to matter will be rare, so there’d be little selective benefit.

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  20. I am not completely convinced by the ruling out of ethocentrism due to the low relatedness: I think that there is a hidden assumption in the Hamilton rule that the altruist trait is coded by a [set of] gene that have the same distribution as the rest of the genome, and that it can not be detected independently but only infered by probabllistic rules linked to relatedness…
    This is not necessarily true for ethnicity. Imagine that an allele that have a clear effect on phenotype (visually detectable, like EDAR variant) also increase altruism for ” people that looks like you”. Then it can be selected for, no?

    It look contrived, but it may be quite common, I think I remember studies showing that your (unrelated) friends have more alleles in common with you than random people from your place. If this is true, it is in line with what I try to tell here: Atruistic behavior for people that “look like you” is possible, because “looking like you” means also “more genetically similar”, regardless or relatedness….

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  21. @kai:

    “This is not necessarily true for ethnicity. Imagine that an allele that have a clear effect on phenotype (visually detectable, like EDAR variant) also increase altruism for ” people that looks like you”. Then it can be selected for, no?”

    The problem is that alleles for altruistic behavior itself are of interest. New alleles always originate in a single individual. Now how could said putative alleles have grown in frequency if the targets of altruism – hence selection for these alleles (distant relatives or even unrelated people) were highly unlikely to carry it? The fitness of the bearer goes down but the allele does not increase in frequency to compensate.

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  22. Also, another problem with standard interpretation of Haldane rule is that it is used in the very rare case of sacrifice, because it makes things simpler: the altruistic behavior is self-sacrifice (reducing fitness to 0) for saving people (increasing stiffness from 0). Should I die to avoid 2 of my brother to be executed? It is nice, simple and clearly altruistic, dying for saving 2+ brothers or 8+ cousins, but this does not happen a lot.
    Most of the time it is: should I help this guy (increasing his fitness, decreasing mine, but not necessarily in a symmetric way). If I am well off and my third cousin is in deep trouble, a small gift can help him a lot without much decreasing my fitness….So more compassion towards people more like you could well evolve due to inclusive fitness

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  23. @Jayman, yes for this to work the carrier of altruistic genes should be detectable. The genes for altruism itself must have an external marker (different from being altruistic, because then reciprocal altruism is enough an explanation – it get’s difficult to separate the 2 effects).

    But I find the higher genetic similarity between friends puzzling, could be that it’s easier to be friend the more similar you are, but even this needs an explanation…After all, friends can be well defined as people you trust and are more likely to help (i.e. the ones that benefits more from altruistic behaviors compared to others). A higher degree of genetic similarity in this context scream inclusive fitness selection…

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  24. It’s important to remember that Hamilton’s rule, as usually stated, is an approximation under weak selection. The reason is that r is the coefficient of relatedness under neutral dynamics, such that each allele is equally likely to be passed on to offspring. But under selection (e.g., kin selection), allele frequencies are changing, so r is usually not exactly right. The error is greater as relatedness grows weaker.

    A simple demonstration of this: suppose a new allele arises that causes an individual to sacrifice itself, at great benefit to its siblings. Regardless or rb>c, this allele will not spread: it kills itself whenever it arises.

    There are other complications. Hanging out with your kin increases the r of your social network, which might favor higher altruism. But this means you will also be competing with your relatives for mates, which discourages altruism. In some cases these effects exactly cancel each other out, such that dispersal has no effect.

    My experience is that it is easy to use Hamilton’s Rule and inclusive fitness to come up with wrong ideas, because they are complicated concepts. I still think just asking the question “Suppose a new allele arises that does (xyz). Will it spread?” usually gives better intuition. Better yet, use a pop gen model.

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  25. @tom – “My comment is directed at trait evolution in individuals that result in behaviors that operate on group dynamics. For example, a gene mutation that expresses as a proclivity for cooperative behavior (and thereby leads to group cohesion and joint action) would have a reproductive advantage if the group survivability was enhanced.”

    ok. but wouldn’t the other members of the group need to be close kin so that at least some of them (enough of them) would carry these same altruism genes? seems to me that we’re just back to kin selection then. -?-

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  26. @british nationalist – “Doesn’t explain why some people go out of their way to be altruistic to those least related to them, such as adopting orphans from Africa, or giving board and lodging to a dog! Darwinists are reduced to ‘evolution gone wrong’ to explain these things.”

    what jayman said above about reciprocal altruism.

    also, one shouldn’t look at the ever-expanding circle of altruism in certain human populations (primarily northwest europeans) as an example of “evolution gone wrong.” this is why i mentioned in the post that i think it’s likely that many of our social behavioral traits are probably general in nature. i think what makes sense is that, in human populations where high levels of reciprocal altruism appear to have been selected for (again, nw europeans are the prime example), that what’s been selected for is not a drive to be helpful to the unrelated neighbor who nevertheless looks rather similar to yourself, but just to be helpful to strangers in general. such non-specific urges could be quickly and easily applied to all sorts of “strangers” — even those of other species.

    have a look at staffan’s terrific post The Myth of the Expanding Circle or You Can’t Learn How to Be an English Vegetarian for more on this idea.

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  27. @grey – “Ethnic genetic interests clearly exist..”

    i think there’re a couple of things here:

    1) could ethnocentrism — i.e. a set of behaviors that would promote ethnic genetic interests — be selected for via kin selection? the answer is no (and i think we agree on that).

    2) how would we sit down and calculate ethnic genetic interests? it can’t be just by adding up the total number of genes (alleles) in common in an ethnic group versus those in another ethnic group. that, to me, sounds like it’s lewontin’s fallacy in reverse — the genetic similarity of a group isn’t just dependent upon how many alleles they share in common totally, but the frequencies of alleles at several loci at the same time.

    maybe this is exactly what frank salter did in his book On Genetic Interests. i dunno. haven’t read it! =o either way, from what i gather, salter’s argument is that looking out for one’s egi would be a logical, rational choice. but when are humans rational?! very infrequently. =/ can’t see how egi would’ve led to ethnocentrism, either.

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  28. @grey – “I agree that among large ethnies relatedness is too low for ethnocentrism to evolve directly but that’s where gene-culture co-evolution comes in.”

    ftr, my working hypothesis at the moment is that ethnocentrism pops up in some of my inbetweener groups — not as inbred as the arabs (who are all about kin), but not as outbred as nw europeans (who are all about EVERYbody including dogs and chimps, etc.).

    the swiss are a good example. maybe even the irish of today (esp. when they’re abroad in somewhere like boston!). the extreme nepotistic altruism of, say, arab groups is not there, because there’s been quite a lot of outbreeding. but the outbreeding (and manorialism) hasn’t been long enough for enough selection for extreme universalism to have set in. so, the altruism circle has been expanded quite a bit, but not to infinity.

    prolly need the modern world for ethnocentricsm to come to the fore, too. the people on one end of ireland need to know and hear about the people on the other end to bother including them in “their group.” (obviously also helps to have an external enemy, too.)

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  29. @grey – “Evolution selects for a particular environment so it can’t go wrong for that environment but if the environment changes then traits that were beneficial can become harmful.”

    yes! extreme universalism works great as long as (almost) everyone else in your society are also extreme universalists. problem is when you get people(s) who are not moving in. =/

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  30. @jayman – “New alleles always originate in a single individual. Now how could said putative alleles have grown in frequency if the targets of altruism – hence selection for these alleles (distant relatives or even unrelated people) were highly unlikely to carry it?”

    exactly!

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  31. BTW, the comment I made above (about why limited dispersal does not necessarily favor more altruism, despite increasing r) also explains why inbreeding does not necessarily favor higher altruism either.

    Intuitively: Under inbreeding, your sibs and cousins are more closely related to you than under random mating. But inbreeding causes them to be your main competitors for mates, too. These effects can cancel each other out – although it depends on the details. So “sticking close to home” increases inbreeding, but does not necessarily lead to greater altruism.

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  32. @rcb – “It’s important to remember that Hamilton’s rule, as usually stated, is an approximation under weak selection. The reason is that r is the coefficient of relatedness under neutral dynamics, such that each allele is equally likely to be passed on to offspring.”

    the reason that i’m interested in hamilton’s rule (as much as i understand it being a non-scientist!) is because it seems as though that, if the relatedness between two mating individuals is higher than what you’d expect in a completely randomly mating population (i.e. if you’ve got an inbreeding population), then each allele is not equally likely to be passed on to offspring. if mom and dad are first cousins (or double-first cousins), they’re much more likely to share duplicate copies of alleles, so those alleles are more likely to be passed on. no? then it seems as though, given favorable selection pressures, certain traits ought to be passed on more rapidly than otherwise.

    same, then, with the selection for altruistic behaviors via kin selection — increase the relatedness between close kin (siblings, aunts/uncles, cousins) and that ought to add some fuel to the fire, so to speak.

    this seems to fit the patterns we see in human populations: lots of nepotistic altruism in populations with frequent cousin marriage over the long-term (the arabized world, for instance), less of it in populations which have avoided cousin marriage for a very long time (northwestern european populations).

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  33. @rcb – “These effects can cancel each other out – although it depends on the details.”

    yes. prolly depends on the details. i imagine this could hold true for many other animals species, but one of the details re. humans is that in inbreeding populations, marriage is often arranged, so the competition for mates is…i dunno…not what it’s like in non-human species! (~_^)

    what you get in the arabized world, for instance, is that marriages are not only arranged, but typically males will by right hold the first option to marry his paternal parallel first cousin (his father’s brother’s daughter). the competition is really nearly eliminated here.

    there are all sorts of other funny mating arrangements out there in human societies that you’d think could affect the selection for altruistic behaviors.

    thanks for the thoughtful comments, btw! (^_^)

    Reply

  34. @ hbd – “ok. but wouldn’t the other members of the group need to be close kin”

    Yes, that’s the likely scenario. These behavioral traits likely developed hundreds of thousands of years ago, during the hunter-gatherer period. At that time, humans were living in small bands (all closely related) and a single advantageous mutation would have replicated and reinforced very quickly within the group before expanding outward. This outward expansion would only occur if the group was more successful vis-a-vis other bands that lacked the advantageous allele.

    There are lots of new studies coming out that indicate genetic-based behavioral traits are both extensive and determinative in all species, including homo sapiens.

    Reply

  35. Nice post hbd chick.

    I don’t grasp the field enough to enter the argument. I can do enough math to see that the equation is valid. However, it has not been proven that this is the only way that this behavior can arise and be sustained.

    Since we are doing equations; maybe somebody can help me with the following:

    NWE suicide bombers = 0*; non NWE suicide bombers = many

    *Any NWEs gone over to the other side as a suicide bomber?

    Jayman, will you go back to West Hunter and answer my family history assimilation question? I sincerely don’t understand how you can say these people were not assimilated.

    Reply

  36. @Jayman

    re: …demonstrates why “ethnic genetic interests” do not exist. Ethnocentrism cannot evolve via kin-selection because the coefficient of relationship falls off so much once you go past 2nd cousins.

    Could you clarify what you mean? It’s difficult to tells because you employ the term “ethnic genetic interest” in an idiomatic way. I commented elsewhere:

    “Jayman’s comment — that “Coefficient of relationships…demonstrates why ‘ethnic genetic interests’ do not exist” — is either semantically of conceptually confused. “Ethnic genetic interest” is Frank Salter’s term which means something like the “distinctive genes carried by an ethnic group.” As used by Salter, the term is equivalent to neither “inclusive fitness” nor “ethnic/kin altruism”*. Ethnic genetic interest, so defined, could not not exist, except in a scenario were ethnic groups did not pick out biological natural divisions (races). What Jayman means, I think is that, (a) as a contingent fact of history (e.g., due to groups with large genetic differences not living in close proximity) “ethnic altruism was not selected for”; it is also possible that he means that (b) “ethnic altruism is not relative fitness (in the biological sense) increasing and therefore that it could not have been selected for. One can have (a) without (b). Mathematically, (b) is untenable. The problem with (a) is that ethnic altruism could have been derived indirectly though inclusive fitness enhancing selection which promoted general genetic similarity favoritism, such as Rushton’s GST. Those who think otherwise need to explain the phenomenon and heritability of ethnic favoritism. Personally I would take Salter and Harpending more seriously then I would Jayman.

    Cite:Salter, F., & Harpending, H. (2013). JP Rushton’s theory of ethnic nepotism.Personality and Individual Differences, 55(3), 256-260.

    There are a number of pathways by which populations could have acquired a congenital disposition for [racial favoritism]. The most plausible is that this is a serendipitous, from the perspective of ethnic genetic interest, over-generalization of a general genetic similarly favoritism mechanism, resultant from kin selection. The genetic similarity between two random White Europeans relative to Black West Africans, for example, is equivalent to that between halfsibs (Harpending, 2002). While not inbred, Europeans are still, owing to geographic isolation, linebreed — hence a race. You would only need a mental module which could recognize magnitudes of genetic similarity of this degree (which we have), and which would incline one to favor individuals this relatively related — and not care if individuals were actually immediate kin. A mechanism that limits favoritism only to immediate kin, that can distinguish between relative genetic relatedness and absolute, would seem to me to be less plausible — it would also contravene a common sense interpretation on Hamilton’s rule. And why would this have evolved? It would seemingly have only under the dual conditions that generalized favoritism (conditioned on degree of relatedness) was maladaptive — yet mathematically it is not — and that you had roving bands of very differently related people around, making this an issue. But you have to argue that the latter case did not hold to argue against a non-serendipitous form of evolved ethnic altruism”

    I’m not sure about your argument, since there are several related issues that you might be discussing: (a) could ethnic altruism historically have evolved via ethnic selection? (b) could apparent ethnic altruism represent an over-generalization of a mechanism evolved via kin selection (broadly construed) e.g., Rushton’s general similarity theory? (c) would ethnic altruism, in our multiracial world, be adaptive (or inclusive fitness increasing)? etc Salter’s primary position concerned (c) –I think that it’s pretty sound. Whether this situation could have led to selection for ethnic altruism, given the historic frequencies of encounters between substantially dissimilar groups is another matter.

    *e.g., In: On Genetic Interests: Family, Ethnicity, And Humanity in an Age of Mass Immigration”

    “In chapters 2 and 3 we saw that ethnies carry large stores of their members’ distinctive genes, especially when contrasted with ethnies of different racial background. It is therefore plausible to assume that it would be adaptive to direct a great deal of altruism towards one’s ethny. After all, if it is adaptive for a parent to make sacrifices for a family containing a total genetic interest of a few children, it is easy to conclude, that efforts to preserve a population carrying the equivalent of thousands of millions of children must at least be adaptive. The adaptiveness of ethnic patriotism would seem to follow directly from a rule formulated by W.D. Hamilton for deciding whether altruism is adaptive… In fact this common sense proposition is controversial, when it is discussed at all. Dawkins dismisses the notion that racial similarity denotes any significant degree of kinship. Commentators responded with equal brusqueness to Rushton’s argument that competition between ethnies could amount to group selection of similar genes. The likelihood is hardly discussed, rare exception coming out against the idea that ethnic altruism is or even could be adaptive.

    The crucial issue for individual actors is whether ethic altruism is adaptive; can they sacrifice their individual reproduction for the sake of their ethnics without losing genetic interest, that is, without contributing to the elimination of their distinctive genes from the gene pool.

    Inclusive fitness is widely misunderstood to mean genetic interests, though the two concepts are distinct. It is often stated that inclusive fitness is the number of copies of ego’s distinctive genes existing in offspring and collateral kin. The definition actually describes familial genetic interest. Inclusive fitness is the effect of an individual’s behavior on the reproduction of his distinctive genes in himself and others (usually kin and fellow ethnics).”

    Reply

  37. @tom – “These behavioral traits likely developed hundreds of thousands of years ago, during the hunter-gatherer period.”

    sure. but i think that they’ve also been tweaked and retweaked in more recent times, especially since the advent of agriculture. recent and (rather) rapid human evolution — which is ongoing — yada, yada, yada.

    Reply

  38. @cracker – “However, it has not been proven that this is the only way that this behavior can arise and be sustained.”

    no, not the *only* way, prolly. but one of the ways and a significant way. and very relevant, i think, given that many human populations have experienced long-term inbreeding (eg. cousin marriage), whereas others have not; and as i said in an above comment, if kin selection *is* a thing, then one would think that altering the degrees of relatedness in a population might alter how the selection for certain altruistic behaviors happens (how quickly, for instance, or maybe even in which directions).

    @cracker – “NWE suicide bombers = 0*; non NWE suicide bombers = many”

    nwe = the product of long-term outbreeding, therefore there are low levels of nepotistic altruism in the population.

    non-nwe (like arabs) = the product of long-term inbreeding, therefore there are very high levels of nepotistic altruism in the population.

    maybe.

    Reply

  39. @chuck – you quoted salter & harpending as saying:

    “The most plausible is that this is a serendipitous, from the perspective of ethnic genetic interest, over-generalization of a general genetic similarly favoritism mechanism, resultant from kin selection.”

    i think this is the most likely route as to how ethnocentrism pops up in some populations. it is a sort-of misapplication of drives and feelings that were selected for via kin selection and meant to be applied to kin.

    my impression is (and this could be completely wrong — somebody needs to check it) that, if we just look at europeans for a sec, the most ethnocentric today are groups like the scots-irish, the scots, the irish, the swiss (sort-of), the poles (maybe). it’s NOT the english, the dutch, the french, the germans, the scandinavians.

    that pattern corresponds to the outbreeding/inbreeding, medieval manorialism/non-manorialism, core europe/peripheral european pattern that i’ve been discussing ad nauseam here on the blog. my working hypothesis is that the medium level of nepotistic altruism of these not-so-outbred groups (like the scots-irish or the swiss) manifests itself as ethnocentrism in our modern world (where communications between the members of the population are excellent). groups with really low levels of nepotistic altruism — the english, the northeastern french, the dutch — they exhibit very low levels of ethnocentrism. they are, rather, uber-universalists. (no ethno-stuff for them!)

    that’s my current working hypothesis. could always be wrong.

    so, ethnocentrism can’t evolve via kin selection; ethnocentrism is a “misapplication” of medium-levels of nepotistic altruism; and ethnic genetic interests *might* be a thing, but humans would have to fulfill them using logic and reasoning, and that’s difficult since most humans are *not* logical.

    Reply

  40. @kai:

    “But I find the higher genetic similarity between friends puzzling”

    One of the few studies I know on the matter found results that we’re statistically significant, so that could just be BS. The other matter is “population stratification” (finding Italians are best friends with other Italians out of Whites in New York City, for example), so that might not mean what we think it means.

    Reply

  41. @RCB:

    “It’s important to remember that Hamilton’s rule, as usually stated, is an approximation under weak selection. The reason is that r is the coefficient of relatedness under neutral dynamics, such that each allele is equally likely to be passed on to offspring. But under selection (e.g., kin selection), allele frequencies are changing, so r is usually not exactly right. The error is greater as relatedness grows weaker.”

    Sure. Not enough to throw off the general pattern, though. All told, the situation largely “reduces” to Hamilton’s rule.

    “A simple demonstration of this: suppose a new allele arises that causes an individual to sacrifice itself, at great benefit to its siblings. Regardless or rb>c, this allele will not spread: it kills itself whenever it arises.”

    Probably. Also probably gives way to alleles that do so only when one must. Self-preservation still reigns.

    “There are other complications. Hanging out with your kin increases the r of your social network, which might favor higher altruism. But this means you will also be competing with your relatives for mates, which discourages altruism. In some cases these effects exactly cancel each other out, such that dispersal has no effect.”

    If that was the case, the Islamic world as we know would not exist. Reality check: it does.

    Reply

  42. @HBD Chick:

    “that what’s been selected for is not a drive to be helpful to the unrelated neighbor who nevertheless looks rather similar to yourself, but just to be helpful to strangers in general. such non-specific urges could be quickly and easily applied to all sorts of
    ‘strangers’ — even those of other species”

    Or at the very least, not discriminate, so to speak, thanks to high levels of reciprocal altruism genes (which includes genes for generosity). When, at that point, the categories are effectively arbitrary, and “expanding circle” is to be expected. Note though, that’s true for exactly one broad group of people around the world.

    Reply

  43. @HBD Chick:

    “ftr, my working hypothesis at the moment is that ethnocentrism pops up in some of my inbetweener groups…the swiss are a good example. maybe even the irish of today (esp. when they’re abroad in somewhere like boston!). the extreme nepotistic altruism of, say, arab groups is not there, because there’s been quite a lot of outbreeding. but the outbreeding (and manorialism) hasn’t been long enough for enough selection for extreme universalism to have set in. so, the altruism circle has been expanded quite a bit, but not to infinity.”

    I’m still working on it myself. If kin selection is out, then that leaves selection for some form reciprocal altruism – discriminatory reciprocal altruism?

    In other words, I’m not sure it’s simply a quantitative matter between inbreeding and outbreeding.

    Reply

  44. @TomA:

    “Yes, that’s the likely scenario. These behavioral traits likely developed hundreds of thousands of years ago, during the hunter-gatherer period. At that time, humans were living in small bands (all closely related) and a single advantageous mutation would have replicated and reinforced very quickly within the group before expanding outward. This outward expansion would only occur if the group was more successful vis-a-vis other bands that lacked the advantageous allele.”

    That’s a group-selectiony argument. Don’t use it.

    Reply

  45. @Cracker1:

    “Jayman, will you go back to West Hunter and answer my family history assimilation question? I sincerely don’t understand how you can say these people were not assimilated.”

    Short answer: assimilation (for things that don’t depend patently on content, like language, dress, cuisine, etc) is genetic in nature. Think about that and it may clear things up for you.

    Reply

  46. @Chuck:

    “The problem with (a) is that ethnic altruism could have been derived indirectly though inclusive fitness enhancing selection which promoted general genetic similarity favoritism, such as Rushton’s GST. Those who think otherwise need to explain the phenomenon and heritability of ethnic favoritism. Personally I would take Salter and Harpending more seriously then I would Jayman.”

    How about Greg Cochran?

    “There are a number of pathways by which populations could have acquired a congenital disposition for [racial favoritism]. The most plausible is that this is a serendipitous, from the perspective of ethnic genetic interest, over-generalization of a general genetic similarly favoritism mechanism, resultant from kin selection. The genetic similarity between two random White Europeans relative to Black West Africans, for example, is equivalent to that between halfsibs (Harpending, 2002”

    It don’t fly, Chuck. The reason is simple: if an altruistic act isn’t going confer a fitness benefit when outsiders are absent (thanks to Hamilton’s rule), it isn’t going to suddenly confer more fitness when outsiders are present: the degree of relationship to your co-ethnics is the same in both scenarios, and so is the fitness payoff to you.

    Reply

  47. @ HBD chick

    “so, ethnocentrism can’t evolve via kin selection; ethnocentrism is a “misapplication” of medium-levels of nepotistic altruism; and ethnic genetic interests *might* be a thing, but humans would have to fulfill them using logic and reasoning, and that’s difficult since most humans are *not* logical.”

    First, one of my points was that both you and Jayman are using the term “ethnic genetic interests” incorrectly. It’s Salter’s coinage and by it he meant neither “ethnic altruism” nor “inclusive fitness”. Your idiosyncratic usages are adding confusion to the discussion, IMO. That said, the above mentioned pathway is what I would imagine. Dawkins has proposed this and so to Cosmides and Tooby (1989), who provide the standard argument against direct selection for ethnic altruism:

    “Finally, it is important to bear in mind that our complex innate psychological mechanisms evolved during the Pleistocene, and were created by histories of selection (see, e.g., Daly & Wilson 1988): modern phenomena such as friction between people of different “races”, war between nation-states, and so on, cannot be adaptations to modern circumstances, but rather reflect the operation of Pleistocene adaptations misfiring under modern circumstances. In fact, non-relatives from one’s own “race” are only slightly more genetically similar than non-relatives from a different “race” (Lewontin 1982), and this modest difference could not have led to any behavioral adaptations, because in the Pleistocene, humans would not commonly have encountered people from different “races”. Instead, competition could only have been between neighboring groups; typically, intergroup conflict would have reflected cooperation with nearer kin against more distant kin. Although in such small-group conflicts, the relatedness of many of the participants in the same coalition must have been very low, the impact of an individual’s decisions on coalition formation, coalition fissioning or exclusion, and coalitional aggression, summed over the members of the two groups, would often have aggregated into substantial inclusive fitness effects. ”

    This “misapplied” kin selection hypothesis should be confirmable in the manner which you suggested. We would see if, on the international level, inbreeding correlated with increased ethnocentrism/racism. That it would, would be a direct prediction of a “misapplied” model.

    @ Jay Man

    “It don’t fly, Chuck. The reason is simple: if an altruistic act isn’t going confer a fitness benefit when outsiders are absent (thanks to Hamilton’s rule), it isn’t going to suddenly confer more fitness when outsiders are present”

    Literally read, this sentence is untrue. First, terms (since there is semantic disagreement):

    genetic interest ~ genetic representatively in a gene pool
    individual fitness ~ ability to maintain genetic representatively via direct reproduction
    inclusive fitness — ability to maintain genetic representatively via direct and indirect reproduction
    altruism ~ behaviors which do not directly increase representatively via direct reproduction

    When dealing with relational fitness, altruistic behavior P, which decreases Zhou the Han’s genetic interest in relation to his neighbor Jianbo’s, can increase Zhou and Jianbo’s genetic interest in relation to Zhou, Jianbo, and //Hui !Gaeb the Khoisan. The introduction of outsiders into our consideration puts the genetic relatedness in different relief. We are now considering a broader gene pool. With respect to the global gene pool, Zhou can increase his more Zhou-like genes by increasing Jianbo’s, just as prior Zhou could increase his more Zhou-like genes by increasing those of his brother. Conceptually, we might expand the gene pool further; we could consider, for example, alternate universe Zhous, Jianbos, and //Hui !Gaebs. From this perspective, our universe Zhou could potentially increase the meta-universe Zhou’s genetic interest by promoting our universe //Hui !Gaeb’s at the expense of his and our universe Jianbo’s (if the cost was offset in the other universes). The above is a crazy scenario which attempts to make clear that inclusive fitness and genetic interest are both relational and abstract concepts, they make sense only in context to a specific gene pool (i.e., set of individuals under consideration) and they do not entail selective pressure (which would require competition between our groups).

    So, in fact, an altruistic act which decreases fitness in relation to one gene pool can increase fitness in relation to another — which is the whole point of inclusive fitness. The questions are: does it and, as a result, was altruism selected for. (I grant that I am uncharitably reading your sentence.)

    “It don’t fly, Chuck”

    I don’t know what you mean.

    First a clarification: As Chick noted, ethnocentrism (ethnic altruism) can represent a ““misapplication” of medium-levels of nepotistic altruism”. One question would be whether this “misapplication” is serendipitous or maladaptive. This would primarily be a mathematical issue. With respect to some gene pool the behavior either does or does not increase the actors’ genetic interest. Salter dispositively showed that ethnic altruism would increase individual genetic interest with respect to the global gene pool.

    Now, what could you mean? You might mean that (1) ethnic altruism was not directly selected for. There are a lot of arguments that could be made in this regards, such as Cosmides and Tooby’s. These seem coherent. (If you mean that it couldn’t in principle, say if broad ethnic and racial groups where historically in competition, this is nonsense.)
    Alternatively, you might mean that (2) a general mechanism, which allows for the overgeneralization of kin altruism, could not have been or was not selected for. I don’t see how you possibly could argue for this based on inclusive fitness theory.

    Reply

  48. @JayMan –

    “If that was the case, the Islamic world as we know would not exist. Reality check: it does.”

    I’m confused. Can you explain to me how the existence of Islam negates the the theory of altruism in viscous populations?

    The point is that “sticking close to home” (viscosity) will increase both inbreeding and average relatedness to one’s social network. But this will not necessarily favor greater altruism within that network. You have to adjust r relative to the mating population you are competing with, and sticking close to home means you are competing more with close relatives. This is a decades-old result in population genetics, not my own musings.
    E.g. http://link.springer.com/article/10.1007/BF01237667#page-1

    Reply

  49. Jayman

    Again, if you mean that as a contingent fact of history (e.g., due to groups with large genetic differences not living in close proximity and so competing) ethnic altruism was not selected for, I can’t dispute the claim. If you mean something else, perhaps you could clarify.

    Reply

  50. @jayman – “I’m still working on it myself. If kin selection is out, then that leaves selection for some form reciprocal altruism – discriminatory reciprocal altruism? In other words, I’m not sure it’s simply a quantitative matter between inbreeding and outbreeding.”

    yup! i agree.

    i think elimating the close mating, tho, can at some point set the stage for a possible increase in the selection for greater and greater amounts of reciprocal altruism. i wondered about this scenario before. could make sense.

    the thing is, i don’t think that the populations with extreme amounts of reciprocal altruism traits (i.e. core europeans) are very ethnocentric. i think they’ve moved beyond that circle in the ever-expanding circle of the ingroup.

    could always be wrong! (^_^)

    Reply

  51. @chuck – “We would see if, on the international level, inbreeding correlated with increased ethnocentrism/racism.”

    can’t be too much inbreeding! can’t be arab-level inbreeding. they have a very difficult time doing ethno-anything.

    Reply

  52. @chuck – “First, one of my points was that both you and Jayman are using the term ‘ethnic genetic interests’ incorrectly. It’s Salter’s coinage and by it he meant neither ‘ethnic altruism’ nor ‘inclusive fitness’. Your idiosyncratic usages are adding confusion to the discussion, IMO.”

    ok. maybe i’m misunderstanding.

    you say above that egi means: “…something like the ‘distinctive genes carried by an ethnic group.'”

    well…yeah. but that’s just restating the obvious, then. something we all know, i.e. that different populations (including ethnic groups) have distinctive sets of genes.

    i thought there was something more to salter’s egi. -?-

    Reply

  53. @rcb – “Can you explain to me how the existence of Islam negates the the theory of altruism in viscous populations?”

    not islam, but the islamic world. a better way of describing it is perhaps the arabized world.

    i think jayman’s referring to what i also mentioned above, i.e. the *very* high rates of inbreeding in the arabized world where they also seem to have *very* high rates of nepotistic altruism (see sailer’s Cousin Marriage Conundrum which appeared in The Best American Science and Nature Writing 2004, ed. by steven pinker).

    @rcb – “This is a decades-old result in population genetics, not my own musings.”

    maybe some scientist could sit down and check this out for inbreeding human populations. look maybe for nepotistic altruism versus reciprocal altruism, etc. (^_^)

    Reply

  54. “i thought there was something more to salter’s egi. -?-”

    the something more is the mathematical argument that one can — and the philosophical argument that one should — increase individual genetic interest through ethnic altruism — whether or not ethnic altruism was directly selected for in the past.

    egi would be igi stored in the ethnic gene pool

    Reply

  55. the something more is the mathematical argument –BY WAY OF INCLUSIVE FITNESS THEORY (ift) — that one can …

    so he applies ift to show why people should, given garden variety concern for igi, be racist — i think that that’s a pretty atypical application

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  56. @chuck – “the something more is the mathematical argument that one can — and the philosophical argument that one should — increase individual genetic interest through ethnic altruism — whether or not ethnic altruism was directly selected for in the past.”

    right. that’s what i thought. that’s why i said:

    “ethnic genetic interests *might* be a thing, but humans would have to fulfill them using logic and reasoning, and that’s difficult since most humans are *not* logical.”

    don’t see how that’s me using the term incorrectly. -?-

    Reply

  57. @Chuck:

    “This “misapplied” kin selection hypothesis should be confirmable in the manner which you suggested. We would see if, on the international level, inbreeding correlated with increased ethnocentrism/racism. That it would, would be a direct prediction of a “misapplied” model.”

    No, misapplied kin selection can’t really explain ethnocentrism. In the post linked above, Cochran explains how states should have bred such right clean out of people. (The possible exceptions being people living today without a long history of living in states, which isn’t the majority, by numbers.)

    Further, HBD Chick’s theory demonstrates that there is great variation along the clannish-universalist dimension, indicating that there was plenty of time for evolution to fine tune people in the proper directions on this matter.

    “When dealing with relational fitness, altruistic behavior P, which decreases Zhou the Han’s genetic interest in relation to his neighbor Jianbo’s, can increase Zhou and Jianbo’s genetic interest in relation to Zhou, Jianbo, and //Hui !Gaeb the Khoisan. The introduction of outsiders into our consideration puts the genetic relatedness in different relief.”

    Here’s the problem with that notion: let’s think of an allele that induces its bearer to be altruistic towards kin (as per degree of relationship). In a homogenous society, it works as we’d expect through Hamilton’s rule, increasing in frequency if it accurately targets kin (since they are also likely to possess this allele). The fitness payoff is as per the chances of said allele being in a relative. Now, the important part is this: this is the only way such an allele can increase in frequency! (Drift notwithstanding.) If you introduce an outside group, the frequency distribution of said alleles doesn’t change. The exist in him and his close relatives as per the coefficient of relationsh ip. Hence all this stuff about genetic background is irrelevant. All that matters is the coefficient of relationship, as it is normally calculated.

    What you and Salter, etc are saying could work if the genes were already common. But that leaves us with a problem: how could they have gotten that way in the first place?

    We can go back to misfiring kin selection, but I think the turmoil in most clannish societies should make us think twice about the idea that kin selection actually misfires to that extent.

    “Salter dispositively showed that ethnic altruism would increase individual genetic interest with respect to the global gene pool.”

    For ethnocentrism to have evolved, it must have done so through something akin to reciprocal altruism: selection acting on each actor individually, and NOT through some form of inclusive fitness, which just doesn’t work.

    Reply

  58. @HBD Chick:

    ““ethnic genetic interests *might* be a thing, but humans would have to fulfill them using logic and reasoning, and that’s difficult since most humans are *not* logical.”

    don’t see how that’s me using the term incorrectly. -?-“

    Where does logic (or at least, its utilization by humans) come from? ;)

    Reply

  59. @ Jayman

    Do you disagree with this statement:

    “Salter dispositively showed that ethnic altruism would increase individual genetic interest with respect to the global gene pool.”

    If you do then you don’t understand the argument.

    JM:”Here’s the problem with that notion: let’s think of an allele that induces its bearer to be altruistic towards kin (as per degree of relationship). In a homogenous society, it works as we’d expect through Hamilton’s rule, increasing in frequency if it accurately targets kin (since they are also likely to possess this allele). The fitness payoff is as per the chances of said allele being in a relative. Now, the important part is this: this is the only way such an allele can increase in frequency!”

    There is a difference between whether a behavior is inclusive fitness increasing, with respect to gene pool x, and whether the behavior was selected for given its fitness benefits with respect to gene pool y.

    JM:”Hence all this stuff about genetic background is irrelevant. All that matters is the coefficient of relationship, as it is normally calculated.”

    By this logic, the genetic relatedness between Joe the European and fourth cousin Sam the European is practically no different from that between Joe and Zhou the Han. Uh, no. This was the point of Harpending (2002); relative to Zhou, Joe and Same are half-sibs.

    JM:”What you and Salter, etc are saying could work if the genes were already common. But that leaves us with a problem: how could they have gotten that way in the first place?…For ethnocentrism to have evolved, it must have done so through something akin to reciprocal altruism: selection acting on each actor individually, and NOT through some form of inclusive fitness, which just doesn’t work.”

    Statements like these suggest to me that you are confusing issues: whether ethnic altruism was selected for versus whether, in global context, it would be fitness increasing (that is, whether it would increase one’s genetic representativity). On the other hand. you might just be wrong.

    JM: “We can go back to misfiring kin selection, but I think the turmoil in most clannish societies should make us think twice about the idea that kin selection actually misfires to that extent.”

    It would not be misfiring but rather “misfiring”. It the latter case it could be perfectly tuned to Hamilton’s rule. And it could be perfectly fitness maximizing in global context. The cross cultural interactions just wouldn’t have driven selection for this generalizable form of altruism.

    I think you are misreading Cochran; but if he thinks that a “misapplied” kin altruism model is improbable, he is wrong.

    Reply

  60. @HBD chick

    HBD chick: “right. that’s what i thought. that’s why i said:

    “ethnic genetic interests *might* be a thing, but humans would have to fulfill them using logic and reasoning, and that’s difficult since most humans are *not* logical.”

    don’t see how that’s me using the term incorrectly. -?-”

    The “might” along with the “fulfill them” suggested that you were not talking about the “the aggregate count of … genes” stored in an ethnic group.

    Salter: “Inclusive fitness is widely misunderstood to mean genetic interests, though the two concepts are distinct. It is often stated that inclusive fitness is the number of copies of ego’s distinctive genes existing in offspring and collateral kin. The definition actually describes familial genetic interest. Inclusive fitness is the effect of an individual’s behavior on the reproduction of his distinctive genes in himself and others (usually kin and fellow ethnics)….Fitness is the effect of investing in copies of one’s genes, It is not the aggregate count of those genes — the genetic interest.” (On Genetic Interests: Family, Ethnicity, And Humanity in an Age of Mass Immigration)

    Reply

  61. @chuck – “The ‘might’ along with the ‘fulfill them’ suggested that you were not talking about the ‘the aggregate count of … genes’ stored in an ethnic group.”

    so increasing “individual genetic interest through ethnic altruism” (your words) is NOT “ethnic genetic interests,” then? there is no goal here (that might be fulfilled) — no aim to promote the reproduction of one’s own genes (via logical and mathematical reasoning)? ethnic genetic interests is simply (to paraphrase salter) the number of copies of ego’s distinctive genes existing in his ethnic group.

    what does he call the goal (or possibility) to increase “individual genetic interest through ethnic altruism” then?

    salter’s definitions of fitness and inclusive fitness are rather unusual, which is where the confusion is coming in, i think. fitness IS normally calculated as the aggregate count of genes (or the individuals that carry those genes), and inclusive fitness as “the sum of [an organism’s] own reproductive output (its direct fitness) and the reproductive output of relatives that share its genes (indirect fitness).” [pg. 42]

    Reply

  62. @rcb – “Hopefully y’all have access to this. Grafen’s paper ‘How not to measure inclusive fitness.'”

    unfortunately, no. =( but thanks for the reference! =)

    Reply

  63. @Chuck:

    “@ Jayman

    Do you disagree with this statement:
    ‘Salter dispositively showed that ethnic altruism would increase individual genetic interest with respect to the global gene pool.'”

    Yes, I disgree.

    “If you do then you don’t understand the argument.”

    The thing is, I do understand the argument, and that is why I am at odds with it.

    “There is a difference between whether a behavior is inclusive fitness increasing, with respect to gene pool x, and whether the behavior was selected for given its fitness benefits with respect to gene pool y.”

    No, fitness means one thing, and one thing only: whether the number of these alleles increasing in the gene pool. This can only happen via selection acting on individuals or via inclusive fitness.

    “JM:’Hence all this stuff about genetic background is irrelevant. All that matters is the coefficient of relationship, as it is normally calculated.’

    By this logic, the genetic relatedness between Joe the European and fourth cousin Sam the European is practically no different from that between Joe and Zhou the Han.”

    Exactly! That’s the thing, and why I said this:

    At least with respect to your inclusive fitness, once you go outside your family (past 2nd cousins), it doesn’t matter the other is a cousin or from a different race. Any inclusive fitness impact to you is effectively the same. This is, by the way, how universalism evolved in an outbreeding group.

    “This was the point of Harpending (2002); relative to Zhou, Joe and Same are half-sibs.”

    And my point, and Cochran’s, is that this is incorrect, at least not for our purposes.

    “Statements like these suggest to me that you are confusing issues: whether ethnic altruism was selected for versus whether, in global context, it would be fitness increasing”

    If it exists (and is too common to be the result of mutation pressure), it must have been selected for or is the result of the misfiring of something that was selected for something else. That’s our only possibilities, right?

    “I think you are misreading Cochran; but if he thinks that a “misapplied” kin altruism model is improbable, he is wrong.”

    You want to ask him? ;)

    Anyways, I’d actually suggest that, because I think we’ve both said enough on the matter, so I think I’ll leave off here unless something new comes up. :)

    Reply

  64. Fairly simple models can help here. Even verbal ones (although these can lead to nonsense if you’re careless).

    Suppose an allele arises that causes someone to be racist toward a large outgroup. Or perhaps to be altruistic toward her own group. Or, in general, to prefer her own group at the expense of others. Will selection favor this allele? Most likely not at the ethnic level. The reason is that, when the allele is rare, race/ethnicity is not predictive of its frequency – the frequency is virtually 0, everywhere. Helping your coethnics does very little to increase the frequency of this allele. If the gene is altruistic (i.e. costly), then the main effect is that the allele wastes energy helping people who don’t carry it. So selection will tend to disfavor large-group racial altruism, *even if races are well distinguished across the genome (at other loci).*

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  65. @rcb – “The reason is that, when the allele is rare, race/ethnicity is not predictive of its frequency – the frequency is virtually 0, everywhere. Helping your coethnics does very little to increase the frequency of this allele. If the gene is altruistic (i.e. costly), then the main effect is that the allele wastes energy helping people who don’t carry it.”

    yes. that’s my understanding of it. (whew! (~_^) )

    thanks!

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  66. @Jayman

    “With really small relationship coefficients, instances where the number of others will be large enough to matter will be rare, so there’d be little selective benefit.”

    Yes, I agree there is too small a benefit for selection to work however the benefit is there mathematically so if a bright spark recognizes that and manages to create an ethno-centric culture that operates on that basis then even if none of the people following that culture are doing it for kin-selection type reasons and are purely conforming to their native culture then they get the benefit. The only person operating on a kin-selection basis in that case might be the original bright spark.

    The other question is could that ethno-centric culture then itself select for ethno-centric traits? dunno. If they did then that might explain the flood of volunteers in 1914 but at the same time would explain the long-term flaw.

    I personally don’t think this is it but I think EGI clearly explains why almost every culture either has or had an ethno-centric culture as long as you define the ethno bit flexibly enough (i.e. from extended family to clan to whatever circle you like which for some of my relatives includes pretty much all carbon life forms).

    #

    “Now how could said putative alleles have grown in frequency if the targets of altruism – hence selection for these alleles (distant relatives or even unrelated people) were highly unlikely to carry it? The fitness of the bearer goes down but the allele does not increase in frequency to compensate.”

    What if it’s attractive?

    It’s in the interest of a man to marry a woman who is altruistic towards the kids. There’s no sacrifice of fitness in this case. The woman increases her fitness by being soppier around babies (up to a point) so her soppy genes spread.

    There are two people involved.

    Reply

  67. @ka

    “Also, another problem with standard interpretation of Haldane rule is that it is used in the very rare case of sacrifice, because it makes things simpler”

    Yes, a *small* fitness cost to potentially help *large* numbers of loosely related people makes perfect sense and why almost every society that produces a large enough surplus almost immediately creates a national health system. The big exception to this being the US where even before 1965 the population wasn’t homogenous enough.

    #

    “yes for this to work the carrier of altruistic genes should be detectable. The genes for altruism itself must have an external marker (different from being altruistic, because then reciprocal altruism is enough an explanation – it get’s difficult to separate the 2 effects).”

    I went for my wife on the basis of how she was with friend’s kids but as you say hard to tell if that was one or the other.

    Reply

  68. @grey (and others) – dunno why some comments are getting caught in comment moderation. =/ it’s been happening the last few days. i hope wordpress fixes whatever the problem is soon. sorry!

    Reply

  69. “Actually, it doesn’t really matter. Past your 2nd cousins, the fitness impact to you (say mating, for example) is effectively the same if everyone else were Chinese or African. What doesn’t work doesn’t work.”

    I think that the ethnic genetic interests idea is sort of sloppy and unnecessary, but I do think that demographic changes do matter on individual and inclusive fitness grounds, and I think that people generally perceive them on those terms.

    For a Chinese person, the social environment changing from Chinese people to African people will result in a new social environment with different selective pressures, and this will have an impact on the Chinese person’s individual fitness and the fitness of their immediate relatives. So the individual or inclusive fitness impact is not the same, but depends on the social environment, which in term will depend on demographic makeup. The average Chinese person who suddenly finds himself in, say, an African environment in which fitness depends on dancing ability or basketball skill will obviously find his individual and inclusive fitness lower relative to, say, a Chinese environment in which fitness depends on academic ability.

    Reply

  70. (some posts disappeared – hope i din’t spam unintentionally)

    @hbdchick

    “so, ethnocentrism can’t evolve via kin selection; ethnocentrism is a “misapplication” of medium-levels of nepotistic altruism; and ethnic genetic interests *might* be a thing, but humans would have to fulfill them using logic and reasoning, and that’s difficult since most humans are *not* logical.”

    I’d quibble on that because I’d say ethno-centrism does evolve via kin selection for low levels of ethny. At the levels it can evolve at ethno-centrism = altruism. It might seem semantic but to me the behavior is the same between whichever two concentric relatedness circles you pick as the border e.g. Pakistani grooming gangs are “racist” at the level of extended families, others are “racist” at the level of tribe, others at the level of nation, others at actual race etc.

    That aside I’d agree that at the level usually defined by “ethno” ethno-centrism isn’t going to evolve at that level on its own because the relatedness isn’t high enough for the selection mechanism.

    So the options are

    1) the physical mechanisms that evolved at lower population levels which made ethno-centrism / altruism work at those more related levels somehow overflow onto people who aren’t related enough – so it exists as an overflow

    2) a bright spark recognizes that ego ~ 512 x 4th cousins and therefore creates cultural adaptations that promote ethno-centrism. these adaptations might have a purely cultural effect, they might simply magnify the effect of (1) or they might culturally select for ethno-centrism. if it’s either of the first two then it would break down if the supporting culture broke down. the humans involved wouldn’t need to be logical – only the bright spark who figured it out would be logically acting out of inclusive fitness.

    3) something else

    Personally I think EGI clearly exists (at every level of “E” from self upwards) but ethno-centrism at the scale of contemporary populations probably requires culture (but maybe didn’t or not as much when the population was smaller).

    but anyway.

    #

    What I think is more interesting is the idea that out breeding didn’t necessarily select for anything but simply reduced selection for something else instead.

    If you take skin color moving from the tropics to the mid latitudes did they get lighter because lighter skin was being selected for or simply because darker skin *stopped* being selected for.

    If black and white are the extremes and you take away selection pressure in either direction then random mutations and time should settle at medium brown.

    Similarly if altruism (defined here as being nice to kin and nasty to non-kin) is strongly selected for in clannish populations then as soon as you take that pressure off people should start to drift to the middle of the spectrum as contrary random mutations stop being selected against.

    The opposite end of the altruism spectrum would be being nasty to kin and nice to non-kin with the mid point being treating everyone the same: not as nice to kin and as nasty to non-kin as the more clannish altruists but somewhere in the middle.

    Reply

  71. Regarding group selection, it seems that Edward O. Wilson, one of the primary proponents of group selection, adopts Richard Dawkins’ “extended phenotype” concept to explain group selection: While he continues to use the term “group selection”, it seems like what he’s really describing is individual selection, with the group being the extended phenotype of the individuals.

    From “The Social Conquest of Earth”, page 55, chapter “The Creative Forces”:

    Selection has remained at the individual level, queen to queen. Yet selection in the insect societies continues at the group level, with colony pitted against colony. This seeming paradox is easily resolved. As far as natural selection in most forms of social behavior is concerned, the colony is operationally only the queen and her phenotypic extension in the form of robot-like assistants.

    From “The Leafcutter Ants: Civilization by Instinct” by Wilson and Hölldobler page 9:

    “[T]he entire colony represents an extended phenotype of the queen and her mates on which evolutionary selection operates”

    From http://seedmagazine.com/content/article/the_hive_mind:

    “There are traits that are expressed in the workers that are not expressed in the queen,” says Hölldobler. “The colony’s traits are the phenotype of the queen, but it is not really the queen’s phenotype. It is the extended phenotype. The colony, even the nest structure, is part of this extended phenotype.”

    Moreover, Wilson and Hölldobler explicitly credit Dawkins by name for the concept of the extended phenotype that they explicitly make reference to in their work.

    Reply

  72. HBD chick

    “so increasing “individual genetic interest through ethnic altruism” (your words) is NOT “ethnic genetic interests,” then? there is no goal here (that might be fulfilled) — no aim to promote the reproduction of one’s own genes (via logical and mathematical reasoning)? ethnic genetic interests is simply (to paraphrase salter) the number of copies of ego’s distinctive genes existing in his ethnic group…what does he call the goal (or possibility) to increase “individual genetic interest through ethnic altruism” then?

    When meaning “a good, evolutionarily speaking” he often says “ultimate interest” and when meaning “number of copies of our distinctive genes”, he usually says “individual/family/ethnic interest”. The two usages are related, and he flows back and forth, since the latter is identified with the former. So he will say, for example:

    “Taken together with the limits imposed by carrying capacity, these kinship coefficients mean that, other factors being equal, immigration is more harmful to the receiving population’s genetic interests the more genetically distant the immigrants”
    …or
    “But this argument has not as yet been supported by quantification of ethnic genetic interests—the number of copies of an individual’s distinctive genes carried in his or her ethny. Without such estimation it is impossible to determine who has it right, those who see ethnies as extended families, or those such as Lewontin (1972) who deny that humans have genetic interests in their ethnies.”
    …or
    “Thus genetic interests are the number of copies of our distinctive genes carried by reproducing individuals. Individual genetic interest is the number of copies carried by offspring. Familial genetic interest is carried by close kin, and ethnic genetic interest by one’s ethnic group. Genetic interests are often confused with ‘inclusive fitness.’ The latter concept was coined by Hamilton (1964) to describe his theory of altruism. It refers to the effect that an individual has on the reproduction of his distinctive genes, not to a static gene count. ”

    Above one could substitute “genetic interests” with either “evolutionary good” or “number of distinctive genes”. So “no” and “yes” depending on how you want to read this. I was just trying to clarify that:
    (a) egi is not “ethnic favoritism” and it is not “inclusive fitness” per se
    (b) there is no “might” about “ethnic genetic interests”, unless by *might* you meant something like “meaning dependent” and not “could plausibly be”. [Imagine: “races *might* exist”..].

    As for the egi / inclusive fitness distinction, Salter draws it out on page 120-123 of “Ethnic genetic interest”: “Inclusive fitness has some counter-intuitive properties. An individual can have large ethnic genetic interest but, unless he or she is investing in that interest, little or no inclusive fitness…”

    On the individual and family level genetic interest and inclusive interest will more or less correspond, and the former will be a measure of the latter, so sometimes no conceptual distinction is drawn. This is similar to how in biology (spatial) “populations” correspond with races in the wild, so races are often identified with populations; there is a logical distinction that becomes important in certain instances, though.

    I think Salter’s use is consistent with Hamilton’s:

    “Inclusive fitness may be imagined as the personal fitness which an individual actually expresses in its production of adult offspring as it becomes after it has been first stripped and then augmented in a certain way.(1964)”

    But it doesn’t matter since “ethnic genetic interest” is Salter’s, so, when employing the term, one should grant the distinction which it’s based on or use a different expression or put “” around it.

    Reply

  73. Lastly I think the health system argument works both ways. On the one hand I think it illustrates ethno-centrism but on the other it shows it’s quite weak at high population levels as it only involves a very low fitness cost – an individual’s tax contribution – and a very large number of potential beneficiaries. Even in a very homogenous country what percentage of income would be the tipping point where people balked?

    The other interesting (to me) point about this is from an EGI point of view progressive taxation is effectively a way of trying to equalize the fitness cost.

    Reply

  74. I think ethnocentrism can be explained in terms of individual and inclusive fitness. Since an individual’s fitness will depend on social environment, and social environment depends on demographic makeup, an individual can adopt ethnocentric attitudes to promote his individual and inclusive fitness. Adopting an ethnocentric attitude that, say, prevents immigration and thus restricts the labor supply and keeps wages higher can be good for an individual’s economic prospects and thus his fitness. I think it’s this sort of concern about individual and immediate family fitness that primarily explains ethnocentrism, rather than thinking about 5th cousins or how related non-family members are and that sort of thing.

    I think another way that ethnocentrism can be explained by individual and inclusive fitness is the fact that getting other people to behave ethnocentrically can be very good for a particular individual’s fitness. Obviously being king of a group can be very good for the king’s fitness. Getting other people to swear allegiance to you, behave ethnocentrically, help you conquer territory, and crown you king of the nation can be a very good strategy with enormous fitness payoffs for a man who can pull it off. It’s not necessarily the best strategy for the infantryman on the front lines however.

    Reply

  75. @ Jayman

    JM: “The thing is, I do understand the argument, and that is why I am at odds with it.”

    You must be using the terms differently.

    Ask Greg: “Do you agree with Salter that ethnic altruism would increase an individual’s number of copies of distinctive genes in the global gene pool — for example, that if Zhou the Han helped many “unrelated” Han reproduce, at his personal reproductive expense, there would, at the end of the day, be, from a global perspective, relatively more Zhou-like genes than if he did not. To be clear, I am not asking if you think that ethnic altruism could have been or was selected, given various historical considerations, but whether, as Salter contends, this is a way to increase the number of one’s distinctive genes in the species’ gene pool.”

    But I’m glad that we got to the heart of the matter.

    Reply

  76. @grey – “some posts disappeared – hope i din’t spam unintentionally”

    shoot. i thought i got them all out of the spam box. sorry! i hope i didn’t delete any of your comments unintentionally! =/

    the comments system has been weird all week. dunno what’s going on with it. some very plain comments with no links at all have gotten tossed in spam. annoying!

    Reply

  77. Regarding the health system, the contemporary moral and philosophical justification for welfare state liberalism isn’t based on theories of or appeals to altruism, but on the work of the American philosopher John Rawls, who bases welfare state liberalism on “justice”, which he defines as what a rational individual actor would regard as fair and choose in an “original position” under a “veil of ignorance”.

    https://en.wikipedia.org/wiki/Original_position

    https://en.wikipedia.org/wiki/Veil_of_ignorance

    The idea is that if one were in an “original position” in which one would develop a society to be in, but were behind a “veil of ignorance” and thus had no idea of what position, wealth, level of talent and abilities, etc., one were to have, then one would being rational focus on ensuring that the society was fair and accomodating regardless of how one would end up in the society. One could be born crippled, or poor, or with low intelligence, and thus it would be rational for a person in an “original position” to develop the society to make sure that the society is fair and provides a decent standard for such people.

    Reply

  78. Can anyone locate:

    Selection of selfish and altruistic behavior in some extreme models. In: Man and beast: Comparative social behavior. J. F. Eisenberg and W. S. Dillon. Washington, D.C., Smithsonian Institute Press: 59 – 91.

    I would like to read the full of Hamilton’s discussion referenced by Salter (2008):

    “If there is free mixing within subdivisions an encounter concerns a randomly selected pair from the subdivision. The correlation of gametes from such a pair is zero with respect to their subdivision. Thus an altruistic trait expressed in random encounters is certainly counterselected within the subdivision. The correlation of gametes with respect to the population is Fst, which is always greater than zero, depending on the degree to which the gene frequencies of the isolates have differentiated. Thus if there is a gain to inclusive fitness on the basis of the coefficient 2Fst/(1+Fit) the genes for the trait are positively selected in the population as a whole” (Hamilton, 1971)

    Salter (2008), relatedly cites Harpending:

    “Total kinship is then derived in a familiar way. Two individuals within the same subdivision are related as]’0 =- 1/(2N – 1) with respect to the gene frequency pi l of the subdivision, and Epi,=f0 + (1 – f,)pil,. With respect to the total population, take the expectation of pi, here; EEpi,0 =fo + (1 -fj)[pi(l – Fs) + FstI, and equation (1) gives Jo = Fst + (1 – Fst)[- 1/(2N – 1)]. Thus two different individuals are negatively related with respect to the local population, but they may be positively related with respect to the total population or the species. This will mean that helping behavior within the subdivision will be selected against locally, because kinship is negative locally, but it may be positively selected within the species because kinship between donor and recipient is positive with reference to the global base population.” (Harpending, 1979)

    This is what I was referring to when I noted that inclusive fitness is relative to the gene pool under consideration e.g., local versus global.

    Intuitively obvious this seemed.

    Reply

  79. @hbdchick

    no it was fine – our posts crossed in the um post, my comment was out of date when i pressed enter.

    Reply

  80. Whatever dog you pick in this fight, it certainly has an interesting history. For example, according to my 2009 version of David Buss’ standard text on evolutionary psychology, referring to E. O. Wilson’s “Sociobiology,” “The bulk of its theoretical tools – such as inclusive fitness theory, parental investment theory, parent-offspring conflict theory, and reciprocal altruism theory – had already been developed by others. What it did do is synthesize under one umbrella a tremendous diversity of scientific endeavors and give the emerging field a visible name.” Apparently Buss isn’t very good at reading the tea leaves, because even in those days Wilson was carrying on something of a dalliance with group selection. He always has. Of course, in his two latest books, he leaves nothing to the imagination, outing himself as such a full-fledged group selectionist that he makes David Sloan Wilson look timid. Relying on the work of Harvard mathematician Martin Nowak and others, he is now claiming that inclusive fitness and kin selection are basically defunct, and mathematically insupportable.

    Going further back, group selection played an interesting role in the “adjustment” of the whole history of the Blank Slate orthodoxy. There is no question that the man most influential and effective in demolishing that “scientific” aberration was Robert Ardrey. The fact was certainly no secret to the Blank Slaters themselves, as can be seen, for example, in that invaluable little piece of historical source material, “Man and Aggression,” edited by arch-Blank Slater Ashley Montagu. However, Ardrey was a “mere playwright,” and, as we all know, the likes of Shakespeare, Ibsen and Shaw can’t possibly have known anything about human nature. It would have been too shy-making for the academics and other “men of science” to admit that their “science” had been corrected by the likes of Ardrey. They had to find a way of dropping him down the memory hole. Enter Richard Dawkins, who claimed in “The Selfish Gene” that Ardrey and others had been “totally and utterly wrong” about (you guessed it) group selection. Pinker then seized on Dawkins’ phrase to claim that Ardrey had been “totally and utterly wrong,” period in his “history” of the Blank Slate. This particular bowdlerization of history has been quite effective, and Ardrey has been more or less an unperson ever since.

    As one who has done computational physics during most of my career, I find the mathematical models that purport to “prove” group selection, inclusive fitness, etc., quite crude. They’re not what we would call “full physics models.” In other words, they don’t don’t include anywhere near all the relevant variables, nor can they, because we don’t have a time machine that can take us back to the times during which the relevant behavioral characteristics were selected for. As a result, I don’t think we can say with absolute certainty whether and/or to what extent group selection happened or not. In other words, the “group selection team” and the “inclusive fitness selection” team are just two more instances of sports fandom.

    Reply

  81. @Brian

    Regarding the health system, the contemporary moral and philosophical justification for welfare state liberalism isn’t based on theories of or appeals to altruism,

    I’d imagine the rationalization for creating a national health system could be different in every homogenous enough nation with surplus enough to pay for one.

    .

    @Helian

    I find the mathematical models that purport to “prove” group selection, inclusive fitness, etc., quite crude.

    The blank slatists stunted western science for nearly 100 years. I expect the models would be a lot more robust by now if that hadn’t happened.

    Reply

  82. Lack of cognitive diversity can also cause endogamy. The first biological predictors to endogamy. Cognitive diversity of europeans explain their predisposition to individualism.

    Reply

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