Kinship coefficients and Ethnic Genetic Interests (JayMan embarrassing himself again)

Contra the White Nationalists, there’s no such thing as “ethnic genetic interests.”
JayMan supports the above JayMan assertion by linking to another JayMan comment, which sees JayMan copy-and-pasting from Wikipedia a table of inbreeding coefficients by degree of relationship, and asserting the table:
…demonstrates why “ethnic genetic interests” do not exist.
Let that sink in. "HBD" hobbyist JayMan sees coefficients of relationship near zero, and asserts (without being fully aware of what he's asserting) that this means I don't share any particular genetic relatedness with my third cousin relative to a tribesman from New Guinea or a triracial from Jamaica.

This is of course not what an inbreeding coefficient near zero indicates:

The solution to the problem posed in Figure 5.1 will be easy if we can calculate the inbreeding coefficient f H of individual H. The inbreeding coefficient of an individual is the probability that the two gene copies present at a locus in that individual are identical by descent, relative to an appropriate base population. Two genes are identical by descent if, and only if, they are descended from the same individual gene copy. Now of course we must stop somewhere as we trace back the ancestry of the two genes. Otherwise any two gene copies would be certain of being identical by descent, provided that life has a monophyletic origin. The function of the base population is to set the context of the problem. In the base population, all gene copies are assumed not to be identical by descent. [Joe Felsenstein. Theoretical Evolutionary Genetics.]

I'm not much more closely related to my third cousin than to a random member of the approximately-random-breeding population from which we both spring. I am genetically markedly more similar to my third cousin (and any other Northwestern European) than I am to someone like JayMan. This is the essence of "ethnic genetic interests".

From Henry Harpending's appendix to Frank Salter's book:

The coefficient of kinship between two diploid organisms describes their overall genetic similarity to each other relative to some base population. For example, kinship between parent and offspring of 1/4 describes gene sharing in excess of random sharing in a random mating population. In a subdivided population the statistic Fst describes gene sharing within subdivisions in the same way. Since Fst among human populations on a world scale is reliably 10 to 15%, kinship between two individuals of the same human population is equivalent to kinship between grandparent and grandchild or between half siblings. The widespread assertion that this is small and insignificant should be reexamined.

Note also that debates about group selection ultimately have no bearing on the reality of ethnic genetic interests, the existence of which is inarguable. When people from the population I belong to are replaced with members of genetically distant populations, this represents a loss of inclusive fitness for me, and one I see no reason to tolerate, irrespective of how strongly selection has operated at the level of groups in the past.

44 comments:

Anonymous said...

Great post.

I like JayMan but I think he has an emotional investment in opposing Ethnic Genetic Interests so he's not the most rational on this topic.

JayMan said...

Hey n/a,

No, actually the problem is I do understand the situation.

As I explained in the discussion linked in my comment, the fundamental problem comes back to the issue of selection. This is the foundation "inclusive fitness." An allele that appears which causes an individual to be altruistic to his distantly related co-ethnics won't become more common through kin selection for a simple reason: distantly related co-ethnics are unlikely to have it. The fitness payoff (i.e., the average frequency of this gene in relatives) is still given by the coefficients of relationship, as listed in the linked table. That's the only way such an allele could increase in frequency (drift notwithstanding – and of course, as we know, that has its limits). Now I suppose this could work if such alleles were already common, but how could they have gotten that way in the first place?

Hence, ethnic favoritism cannot evolve via kin selection. Harpending and Salter are wrong, and so are you. Now, ethnocentrism might be able to evolve through other methods, methods more akin to reciprocal altruism (which doesn't require relatedness to work), but that would still have nothing to do with "ethnic genetic interests", but be more a factor of good old in-group vs. out-group action.

As far as kin-selection goes, coefficient of relationship to distantly related co-ethnics matters more when there's inbreeding going on. When there's inbreeding, the payoff for helping relatives matters more. That's right, this puts us right back to HBD Chick's theory. That's as far as that matters.

n/a said...

JayMan,

No, you have no idea what you're talking about. You don't even know the definition of the thing you're asserting doesn't exist.

Frank Salter's definition of "genetic interests":

"The number of copies of an individual's
distinctive genes. These are most concen-
trated in the individual, then in first de-
gree relatives, thence in decreasing con-
centration to clan, tribe or ethny, geo-
graphic race, and species. In terms of
population genetics, genetic interest can
be quantified as aggregate kinship, as an
equivalent number of children or other
close kin."



His definition of "ethnic genetic interests":
"The number of copies of a random indi-
vidual's distinctive genes in his or her
ethny, not counting the copies in kin. The
size of ethnic genetic interest is relative to
the kinship of genetic competitors. When
competitors are closely related ethnies, the
interest can be relatively small. When
competitors are distantly related, espe-
cially from different geographical races,
ethnic genetic interest can be many orders
of magnitude greater than familial genetic
interests."


It is a fact that I share genetic similarity with my co-ethnics relative to Africans. It is a fact that I suffer a loss of inclusive fitness when my co-ethnics are replaced with Africans. This is true regardless of whether or not group selection has driven adaptations -- related to ethnocentrism or anything else -- in humans. And it's true regardless of whether or not my group is recently inbred.

JayMan said...

You (and many others) can keep repeating it, but it still doesn't get around the root problem that the proposed mechanism for selecting for ethnic altruistic alleles doesn't work via kin selection. You can't get there from here.

n/a said...

I don't know how much simpler I can make this. "Ethnic genetic interests" and "ethnic altruism evolved via group selection" are two different things. Arguments about the latter have no bearing whatsoever on the existence former. Quantification of the former may have implications for theoretical discussions of the latter, but given your failure to grasp even basic concepts, you have nothing of value to contribute to that discussion.

gcochran said...

Jayman is right, of course.

Anonymous said...

Right about what? That this table demonstrates why ethnic genetic interests don't exist?

David said...

JayMan is wrong of course. The mathematics of ethnic genetic interests are very subtle & nothing in his comments indicates that he gets it.

Anonymous said...

Isn't "ethnic genetic interest" a political ideology? Seeking to maintain or increase the prevalence of the (poly)genes common in one's ethnicity may or may not make sense as a political goal, but I don't see how you can refute it logically or empirically because it's not a scientific theory.

Anonymous said...

JayMan's problem is more with reading comprehension than mathematics.

EGI is based on measures of kinship, not the evolutionary process by which that kinship developed. People can understand that without ever knowing how Fst's are calculated.

JayMan is a subtype of the tragic mulatto, who believes he is more intelligent than he is because the other blacks he sees in his daily life are so dumb.

n/a said...

Greg,

No, he's not. You either have no idea what the argument is about, or you also fail to grasp basic population genetics. Which is it?

n/a said...

Crude analogy of what we have here:

JayMan: You won't die if you jump off a tall building. / There's no such thing as jumping off tall buildings.
Me: Yes, you will. / Yes, there is.
JayMan: You (and many others) can keep repeating it, but it still doesn't get around the root problem that, it's impossible we evolved any aversion to jumping off tall buildings, because there were no tall buildings in the past.


Greg Cochran either thinks this is an argument about group selection, or, like JayMan, also fails to grasp that members of a given population who are treated as "unrelated" for purposes of calculating inbreeding coefficients are still related, relative to distant populations.

Anonymous said...

Is Cochran's wife Asian?

JayMan said...

@Anonymous:

"JayMan is a subtype of the tragic mulatto, who believes he is more intelligent than he is because the other blacks he sees in his daily life are so dumb."

For the record, the White people I have seen in my day-to-day life have been pretty f'ing dumb.

Anonymous said...

Even dumber than the blacks?

Since you're interested in IQ, let's see if we can explain EGI this way:

If the world was full of JayMan's, you would be neither intelligent nor dumb.

If the world was full of blacks, you'd be very intelligent.

If the world was full of HBDers, you would be of normal intelligence.

If the world was filled with n/a's, you'd be a cretin.

Now do you understand EGI?

Average Joe said...

For the record, the White people I have seen in my day-to-day life have been pretty f'ing dumb

How can you tell? Did you give them IQ tests or did you reach your conclusions about their intelligence because you did not like what they had to say?

RCB said...

My advice:

Learn mathematical population genetics. It will sort things out quickly.

RCB said...

My last comment was snooty. Let me try again:

As I understand them, Salter-type arguments state that, across the genome, races can look essentially like separate families. E.g. you can see high F_ST values. This is totally true. It's also totally irrelevant to natural selection on altruistic behavior - which is the issue at stake, I presume. That is, we are interested in using evolutionary theory to explain how people behave.

It's irrelevant because the vast majority of mating competition occurs within ethnicities and races (unless they're quite small). This has to be true - if there were complete global panmixia, then separate races/ethnicities would never arise in the first place. In kin selection models, r is defined relative to the mating pool in which an allele is competing. This means that r is essentially set at ~0 for the ethnic group. Hence no altruism.

There are other issues. E.g., genome-wide divergence is irrelevant to a new, altruism-causing mutation that is rare in all ethnic groups. All of this confusion probably arises because people hear about Hamilton's Rule but have probably never seen it derived mathematically. Crazy ideas follow.

There are particular cases (generally called "group selection") where these mechanisms can work, but few people think that they apply to humans.

RCB said...

"When people from the population I belong to are replaced with members of genetically distant populations, this represents a loss of inclusive fitness for me"

No, it doesn't. Check out "How not to measure inclusive fitness," a classic, 30-year-old paper by Alan Grafen, to see why not. Or just read Hamilton's original definition.

Anonymous said...

E.g. you can see high F_ST values. This is totally true. It's also totally irrelevant to natural selection on altruistic behavior - which is the issue at stake, I presume

And you presume incorrectly.

Again, no mathematics is required. Just some basic reading comprehension, which is badly lacking among Dr. JayMan, genius Cochran, and assorted fanboys.

RCB said...

So what is the interest, then? Anyone can make up some quantity and call it "ethnic genetic interest." That doesn't mean it will predict evolutionary dynamics or actual human behavior. The quantities I've seen described here and elsewhere have this problem.

n/a said...

"As I understand them, Salter-type arguments state that, across the genome, races can look essentially like separate families. E.g. you can see high F_ST values. This is totally true. It's also totally irrelevant to natural selection on altruistic behavior - which is the issue at stake, I presume."

No, that was not the primary issue at stake in this thread. And, no, it's not "totally irrelevant" to selection on altruisitc behavior.


"In kin selection models, r is defined relative to the mating pool in which an allele is competing. This means that r is essentially set at ~0 for the ethnic group. Hence no altruism."

You are confused. Relatedness both within and between groups matters in discussions of altruism.

Also, you seem to be imagining an ancestral environment in which groups had no opportunity to compete with genetically differentiated groups, but this is almost certainly wrong. Human prehistory is likely to have featured highly inbred bands or villages in direct competition with one another. Yanomama villages show FSTs on the same order as continental races.


"Or just read Hamilton's original definition."

Explain what about Hamilton's definition you think means I'm wrong. Hamilton expressly discusses multiple levels of selection in later papers (see "Innate social aptitudes of man").

RCB said...

See the paper "Genetic relatedness in viscous populations" by David Queller (a real evolutionary theorist). Or, this sentence in the abstract: "Hamilton's rule still gives the correct condition for the evolution of altruism if relatedness is measured with respect to the local competitive neighbourhood." If your main competitors are those of your own ethnic group (must be true - that's how ethnic groups are made, by well-separated mating pools), then r~0 for this group.

I am aware that FST between small neighboring groups can be high. That's because they are small. We are talking about ethnic-level altruism here (I thought!). Stranger-danger isn't ethnocentrism.

Finally: if foreigners move into your town, no, your inclusive fitness does not drop. Inclusive fitness removes all neighbor-modulated fitness effects. In other words, any aspect of your own fitness that is not due to your own behavior is not counted. This includes things like foreigners moving into town. See the definition in Hamilton 1964 - the original paper. Also read "How not to measure inclusive fitness" by Grafen. This is old stuff: population geneticists have known it for decades.

Bottom-line: If there were extremely strong competition between large, genetically-distinct ethnic groups for long stretches of human history (somehow stronger than within-group competition!), then yes, ethnic-level sentiments would have evolved. AKA group selection. It's hard for this to work because selection within groups (and gene flow) quickly erodes FST for large groups *at altruistic loci under selection*. (Neutral loci throughout the genome may maintain high FST - but, to repeat, these are irrelevant).

n/a said...

"Finally: if foreigners move into your town, no, your inclusive fitness does not drop."

Rephrased: by aiding members of genetically distant populations in replacing members of my own population, I would without question lower my inclusive fitness. By aiding members of my population in avoiding replacement by members genetically distant populations, I might increase my inclusive fitness (depending on the numbers, relatedness, and sacrifice involved).


"I am aware that FST between small neighboring groups can be high. That's because they are small. We are talking about ethnic-level altruism here (I thought!). Stranger-danger isn't ethnocentrism."

Think general mechanisms for recognizing and discriminating in favor of kin, not a specific adaptation for "discriminating in favor of people of European origin".


"See the paper "Genetic relatedness in viscous populations" by David Queller (a real evolutionary theorist)."

This paper challenges nothing I've said:

Pamilo (1984, 1989) has devised hierarchical r-statistics, analogous to F-statistics, that measure
relatedness at different levels in subdivided populations. The results of this paper give some
guidance in selecting among these possible measures of r and also expand the set of choices to a
continuum. Relatedness should usually be measured with respect to the local population, where
most of the competition resides. However, if the investigator has reason to believe (based on
intuition or data) that an individual displaces a deme-mate x of the time and displaces an
individual in another deme 1-x of the time, these values can be used as weights to obtain
appropriate values of p for relatedness estimates. More generally, the message that needs to be
remembered is that relatedness is not just a statement about the genetic similarity of two
individuals, it is also a statement about who their competitors are.

RCB said...

Yes, I suppose that if you reframe the example so as to imply something about your own behavior, then it could affect inclusive fitness. Of course, the invading foreigners would have to hurt you somehow, or prevent you from helping non-foreigners (assuming non-foreigners don't also carry the xenophobia gene in question!).

I doubt that humans ever evolved a kinship recognition system based on phenotypic matching to close relatives, but I could be wrong. If it doesn't exist, then it can't be extended to larger ethnic groups. I don't know this empirical literature well, though. (Of course, most of us do know, with high accuracy, who our close relatives are.)

My point in invoking Queller's paper: if the vast majority of mating competition occurs within ethnic groups (probably almost always the case in human history), then ethnic-group-wide altruism isn't going to evolve. Doesn't matter if the ethnic groups have an FST of .99 across the genome on average.

n/a said...

"Of course, the invading foreigners would have to hurt you somehow, or prevent you from helping non-foreigners (assuming non-foreigners don't also carry the xenophobia gene in question!)."

Being in my territory, competing for resources and mates with me or people like me, "hurts" me.

"My point in invoking Queller's paper: if the vast majority of mating competition occurs within ethnic groups (probably almost always the case in human history),"

And, again, your model of human history/prehistory is wrong: http://racehist.blogspot.com/2015/03/hamilton-on-inclusive-fitness-and.html

RCB said...

"Being in my territory, competing for resources and mates with me or people like me, "hurts" me."

I misspoke: if we're using inclusive fitness accounting, then their behavior is not counted. Them moving in would have to prevent me helping those who do share my genes, relative to those who don't. If we are using a direct fitness accounting, the problem is that both foreigners and non-foreigners will compete with you for mates. (And no, a foreigner mating with you does not "hurt" you.) The REAL problem with this little model is that we're not even putting it in a context of an evolving locus. If the trait under evolution is an allele that fights foreigners who move into town, then the allele that doesn't fight is going to have higher fitness (it reaps whatever benefits the group gets from having a xenophobic allele around, without paying the cost of fighting), and will displace the xenophobia allele. Making a mathematical model would quickly clarify this.

Citing a non-empirical paper doesn't disprove that most human competition occurs within large ethnic groups, rather than between. I'm totally open to the idea that competition between small bands (usually not strongly distinguished by identifiable phenotypic differences) could have shaped some human behavior. We see a lot of it in ethnographies. And it happens in chimpanzees too. But selection for altruism in large, genetically distinct ethnic groups? No. Hamilton wouldn't have believed it. Vast majority of competition would be within these groups.

Make a model. You will see how hard it is for these dynamics to work out. I'm tempted to do this and post it online. Seems like there is some demand for it.

n/a said...

Hamilton himself in "The genetical evolution of social behaviour. II":

4. Discrimination in Social Situations
Special case (b) of the previous paper has shown explicitly that a certain
social action cannot in itself be described as harmful or beneficial to inclusive
fitness; this depends on the relationship of the affected individuals. The selective advantage of genes which make behaviour conditional in the right sense
on the discrimination of factors which correlate with the relationship of the
individual concerned is therefore obvious. It may be for instance, that in
respect of a certain social action performed towards neighbours indiscriminately, an individual is only just breaking even in terms of inclusive fitness. If
he could learn to recognize those of his neighbours who really were close
relatives and could devote his beneficial actions to them alone an advantage
to inclusive fitness would at once appear. Thus a mutation causing such
discriminatory behaviour itself benefits inclusive fitness and would be
selected
. In fact, the individual may not need to perform any discrimination
so sophisticated as we suggest here; a difference in the generosity of his
behaviour according to whether the situations evoking it were encountered
near to, or far from, his own home might occasion an advantage of a similar
kind. [. . .]

Nevertheless, if any correlate of relationship is very persistent, long-
continued weak selection could lead to the evolution of a discrimination
based on it even in the range of distant relationships
. One possible factor of
this kind in species with viscous populations, and one whose persistence
depends only on the viscosity and therefore may well be considerably older
than the species in question, is familiarity of appearance. For in a viscous
population the organisms of a particular neighbourhood, being relatives,
must tend to look alike
and an individual which used the restrained symbolic
forms of aggressive behaviour only towards familiar-looking rivals would be
effecting a discrimination advantageous to inclusive fitness.

RCB said...

You should know that the viscosity argument mentioned in that Hamilton paper often doesn't work. This was not realized until some years after those original 1964 papers. The reason, again, is that sticking close to home causes you to compete with your relatives for mates. r is measured relative to your average mate competitor, which means that for a highly endogamous ethnic group, r~0 on average - regardless of how genetically diverged any other groups are. See the Queller paper I mentioned - it goes over this in the first few paragraphs.

I don't know of any evidence that humans (or any other primate?) use phenotypic matching to identify close relatives with any accuracy. Seems to me that there are much better cues to identifying close relatives: e.g., who you grew up with as a kid, or kinship terms. It's not clear that these mechanisms would extend to between-group variation: a stranger is a stranger, no matter what he looks like. After all, most humans do know who their close relatives are, and probably have for a long time. A phenotypic matching mechanism would probably be too error prone to contribute much beyond these other mechanisms (remember that half of the additive genetic variance occurs within full sib groups, if mating is nearly random within ethnic groups! that's a lot of room for error). I would happily be wrong about this, though, if there is good evidence.

Now, if ethnocentrism is in fact the result of misfiring kinship recognition systems, then this would be a maladaptive outcome. I.e., it would almost certainly not be selectively favored via inclusive fitness, for reasons we've discussed. So it still wouldn't make sense to call it a matter of "genetic interests." But the result would be the same.

I should mention: there are alternative explanations to ethnocentrism and racism. One is cultural group selection, a new-ish theory that posits that cultural transmission of behavior much more easily maintains group differences in behavior, and so could facilitate group competition on larger scales than is feasible via genetics. This would presumably lead to cultural-group favoritism. The jury is out on this, but I mention it is a plausible alternative. More generally: the fact that humans are often ethnocentric doesn't HAVE to be explained by genetic differentiation between groups.

n/a said...

"You should know that the viscosity argument mentioned in that Hamilton paper often doesn't work."

And, again, nothing Queller says conflicts with Hamilton's insights or with anything I've said:

At first, this result may seem surprising from the standpoint of inclusive fitness theory. Taylor (1992a), following a general approach outlined by Grafen (1985), has shown that it is perfectly consistent. The trick is to use Hamilton's rule as intended, but to remember to explicitly include all individuals whose fitness is affected by a behaviour. If you help a neighbour to produce more offspring, then you are ultimately hurting someone else, whose offspring are displaced by the increased competition.

n/a said...

From Hamilton's introduction to a 1971 paper in Narrow Roads of Gene Land:

From the positive side of my miscellany, what seems worth noting? Perhaps the most novel and useful formula is that in the first appendix, which sketches the handling of relatedness in cases where there is inbreeding (a topic soon to be expanded in the paper of Chapter 8). Care is needed with the formula, however: the inbreeding coefficients must be based at an appropriate population scale. 3 One of the most surprising conceptual points in the paper is also concerned with altruism under inbreeding and may seem even contradictory to altruism being correlated with relatedness. This is in my verbal caution that low dispersal by itself (population 'viscosity') as a way of reaching high relatedness has snags. The point is that, to be effective, altruism must put offspring into competition with non-altruists, not bunch them in a wasteful competition with their own kind. Recently this point has been treated explicitly and quantitatively with some quite surprising conclusions.3-5

If relatives cannot avoid the bunching then the group itself must expand at the expense of others. For this to work most effectively expansive groups need to replace other groups; in essence, members need and are expected to evolve a degree of xenophobia. 6 This line leads on to thoughts that for the re-slanted spiritual descendants of the prim Victorians of my second introduction remain quite paralysing and I have to admit that at the time the thoughts were painful enough to me. It is hard even to feel and harder still to write in a way that runs counter to a current world view, especially a moral one, and it is all the harder when the way is re-shaping a plane of perfection to which all civilized cultures are thought to be striving. A scientist or philosopher with a programme of such heresy has to be tough if he or she is to communicate it and, while doing so and for long after, must endure the tortures of Orestes. 1 ,7 For me it was the discussion of the darker of the 'innate aptitudes' that I believed must exist in all human populations and most individual humans, and of the selection our forbears must have undergone through competition between populations, creating their warlike inclinations (including as a sideline, their relish in cruelty) that caused me most pain to write. The feelings remained acute when I had to write similarly for my next symposium contribution (Chapter 9): yet I am glad I gave the discussions I did. Theory concerning the nature of the 'beast within' was why I had been invited and I continue to believe that only from a basis of honest description can there be hope of taming what we have and may not like.

n/a said...

"I don't know of any evidence that humans (or any other primate?) use phenotypic matching to identify close relatives with any accuracy."

First few results from Google Scholar:

Visual kin recognition and family resemblance in chimpanzees (Pan troglodytes).
JR Vokey, D Rendall, JM Tangen, LA Parr… - Journal of …, 2004 - psycnet.apa.org

Social perception of facial resemblance in humans
LM DeBruine, BC Jones, AC Little, DI Perrett - Archives of sexual behavior, 2008 - Springer

Facial resemblance enhances trust
LM DeBruine - Proceedings of the Royal Society of …, 2002 - rspb.royalsocietypublishing.org

Father–offspring resemblance predicts paternal investment in humans
A Alvergne, C Faurie, M Raymond - Animal Behaviour, 2009 - Elsevier

Where are kin recognition signals in the human face?
MF Dal Martello, LT Maloney - Journal of Vision, 2006 - journalofvision.org

Human ability to recognize kin visually within primates
A Alvergne, E Huchard, D Caillaud… - International journal of …, 2009 - Springer

Reactions to children's faces: Males are more affected by resemblance than females are, and so are their brains
SM Platek, DM Raines, GG Gallup, FB Mohamed… - … and Human behavior, 2004 - Elsevier

Kin recognition and the perceived facial similarity of children
LT Maloney, MF Dal Martello - Journal of Vision, 2006 - journalofvision.org

Recognition of neonates by facial-visual characteristics
RH Porter, JM Cernoch, RD Balogh - Pediatrics, 1984 - Am Acad Pediatrics

Cross-cultural perceptions of facial resemblance between kin
A Alvergne, R Oda, C Faurie, A Matsumoto-Oda… - Journal of …, 2009 - journalofvision.org


And re: weaselish "with any accuracy", absolute precision is not required. Any signal of relatedness is potentially useful.


"Seems to me that there are much better cues to identifying close relatives:"

I fully expect there are other cues as well. So did Hamilton. Again, any signal of relatedness is potentially useful, and different signals may be useful in different situations. Specific adaptations for bonding mothers to babies do not rule out other adaptations for kin recognition in other situations.


"Now, if ethnocentrism is in fact the result of misfiring kinship recognition systems"

Accurate, not "misfiring".


"I should mention: there are alternative explanations to ethnocentrism and racism."

Which you seem inclined to prefer for ideological reasons.

RCB said...

"And, again, nothing Queller says conflicts with Hamilton's insights or with anything I've said"

Well, sort of: I'm saying that Hamilton DID mistakenly put too much stake in viscosity as a selective force for altruism (in 1964). He later figured out the problem, though. In other words, he did not originally use Hamilton's rule "as intended", ironically enough. He says as much in that paragraph you cite ("surprising conclusions").

More generally, the stuff you cite should make clear to the readers of this blog that large FST's across groups alone cannot be used to infer selection for ethnic-level altruism. Viscosity increases FST, but may not cause any increase in altruism. Need to understand the range of reproductive competition. This is my main point; it sounds like we agree here.

BTW, I wrote down some models investigating the evolution of endogamy in a particular population scenario (basically, when two ethnic groups "come together" into the same area). Thinking of making PDFs and distributing, to show that relatedness alone will not favor endogamy (i.e., it's not in your "genetic interests" to mate with your own kind, unless you invoke other mechanisms, e.g. lower fitness among the foreigners). Sharing would allow others to check the math and suggest changes, if folks are interested.

RCB said...

First: I have no ideological preferences here, and I don't see any evidence that would make you believe that. Only theoretical interests. I've seen others to fling around political attacks here; that's not my style.

As for those citations: thanks, I'll check them out. I'm skeptical (people want these things to be true, because Hamilton's rule is so compelling; strong possibility for publication bias), but I admit ignorance.

The amount of accuracy in kinship recognition does matter, because it attenuates r. If you can only identify your brother 1/5 of the time, then r drops from 1/2 to 1/10 (or smaller, anyhow). Acting altruisically toward "possible kin" can be worse than doing nothing at all.

And, I say misfiring under the assumption that selection will almost never favor altruism at the level of the large ethnic groups (argued before why this is usually the case). If that's true, then any altruism based on extending kin recognition mechanisms to this level will be maladaptive - hence, misfire.

n/a said...

"Viscosity increases FST, but may not cause any increase in altruism. Need to understand the range of reproductive competition. This is my main point; it sounds like we agree here."

Yes. And this would be relevant to the discussion if humans were subject to the sorts of constraints (like limited dispersal) under which this actually becomes an issue.

Hamilton: Social selection cannot occur in completely regular 'mating systems'. It may be noted here, however, that the most 'system-like' version of an 'isolation-by-distance' model, which is supposed to preclude long-range migration and elastic expansion from vigorous areas, is rather hostile to altruism.

RCB said...

Please explain, then, exactly what model of mating and dispersal you have in mind. This would help the discussion immensely. The models I've referred to usually assume many patches with island-like dispersal (which does allow long-distance migration), in which more cooperative populations produce more dispersers than others ("hard" selection - a form of expansion by successful groups). Do you mean to imply that this is an irrelevant model for human evolutionary history?

Your last comment sounds to me like you're suggesting that there is some case in which genome-wide FST is somehow relevant to selection for altruism at a particular locus (my main dispute throughout). If this is your argument, I emphasize my extreme skepticism, but will of course hear you out. If this is not what you're arguing, then I think we're done here.

n/a said...

"The models I've referred to usually assume many patches with island-like dispersal (which does allow long-distance migration)"

As far as I know, this is wrong. If you bother to read and understand the papers you cite, I think you'll find the models in which increased competition and increased relatedness directly cancel all limit long-range migration and/or local population expansion. Obviously, neither of these limits holds for humans, and Hamilton was aware of the issue prior to his 1971 and 1975 papers.

"in which genome-wide FST is somehow relevant to selection for altruism at a particular locus"

You apparently fail to appreciate that "genome-wide FST" is also a statement about the expected similarity at "a particular locus".

RCB said...

"If you bother to read and understand the papers you cite"

This is the second time you've presumed something about me that was false.

Taylor (1992) "Altruism in viscous populations - an inclusive fitness model" is probably the most cited paper. With regard to long-range dispersal, note: "(4) Dispersal. With probability t, each female disperses to a distant patch, distant enough that she will find no relatives there."

I've actually rederived some of these results myself, so I'm pretty sure I do understand them. As viscosity goes up, FST at the altruistic locus goes up, but altruism does not.

The cancellation noted in Taylor's paper doesn't always happen, by the way. Sometimes viscosity can favor higher altruism among local groups, as I understand. But the point remains that FST alone (even at the locus of interest) cannot be used as an argument for increased altruism. Got to know other things.

n/a said...

Okay, I'd read Taylor's other 1992 paper, but not that one. I believe now that you're attempting to argue in good faith. Unfortunately, the models you're bringing up are still irrelevant for what you're trying to use them for (challenging the possibility that human ethnocentrism could have evolved via group selection): they do not allow for any sort of kin recognition (altruism is dispensed purely based on proximity).

What happens when ancestry indicators are available? A relevant model:

We show with an evolutionary model how contingent altruism
can be sustained even when arbitrary heritable indicators of
relatedness, called ‘‘tags’’, coevolve with the strategies gov-
erning behavior. Discrimination based on tags is not assumed,
but rather evolves endogenously in a viscous population (i.e.,
local reproduction and local interaction) and is selected for
even when phenotypic matching is very coarse-grained.
We
also show how to extend Hamilton’s rule to establish the
conditions under which kin recognition can support discrim-
inating altruism even when coevolution causes the reliability
of indicators of relatedness to vary with each individual’s
evolving social environment.
[. . .]
The resulting agent-based model is based on a model
previously developed to study ethnocentrism in humans
(Ax-
elrod and Hammond 2003). The present model is not meant
to be a literal representation of biological processes. Instead,
our model is designed to illuminate the consequences of the
fact that kin discrimination typically entails coevolution of
three things: the strategies governing behavior, the reliability
of the tags on which the behavior may be conditioned, and
the population structure that determines who interacts with
whom
.
[. . .]
The algebraic method above is the first published analysis
of selection for kin recognition with simultaneous variation
at the indicator and altruistic loci. This method helps us un-
derstand the conditions under which kin recognition can sup-
port discriminating altruism even when the reliability of in-
dicators of kinship depends on the individual’s social envi-
ronment.

n/a said...

Continued:

The value of being able to distinguish tags can be under-
stood in terms of inclusive fitness theory that takes into ac-
count the degree of relatedness between two agents (Hamilton
1964; Lacy and Sherman 1983; Riolo et al. 2001). While
proximity alone can be an indication of relatedness, being
able to distinguish among heritable tags, as in the armpit
effect (Dawkins 1982; Hauber and Sherman 2000; Hauber et
al. 2000; Mateo and Johnson 2000; Isles et al. 2001), allows
a still better indication of relatedness, for example among
sessile cnidarians (Grosberg and Quinn 1989; Grafen 1990).
The discriminatory abilities required for the armpit effect are
likely to be widespread. The self-recognition required for
multicellularity provides them from intimate contact, and the
need to distinguish conspecifics for mating provides them
more generally for animals
. In both cases, a hardwired com-
parison known as the green beard effect (Hamilton 1964;
Dawkins 1976; Haig 1996; Grafen 1998; Keller and Ross
1998) would seriously slow evolution and make speciation
almost impossible.

Viscosity is ubiquitous because few populations complete-
ly mix from one generation to the next. Hamilton (1964)
believed that simple viscosity was a widespread sufficient
cause of fairly weak altruism, and various models have found
that viscosity can indeed foster cooperation (Getty 1987; Pol-
lock 1989; Nowak and May 1992; Nakamaru et al. 1997).
However, this general claim is now considered doubtful. The
balance between increased relatedness and increased com-
petition between neighbors may tilt toward or away from
cooperation (Taylor 1992; Wilson et al. 1992; West et al.
2002). Taylor and Irwin (2000) have suggested that with
overlapping generations, and with altruism dispensed as ben-
efits to fecundity, there is a tendency for population viscosity
to support altruism. The 15.6% cooperation found in our
model with one tag is on the one hand more than zero, sup-
porting Taylor and Irwin, but on the other hand is rather
limited. Adding observable tags shows that proximity can
sustain cooperation based on contingent altruism, even if the
very correlation of tags and relatedness evolves
. By putting
both the matching and the altruism under explicit genetic
control, the model shows how altruism conditional on heri-
table tags can evolve despite substantial costs of cooperation.
Thus, the present model, which combines viscosity, the arm-
pit effect, and endogenous use of discrimination in a genet-
ically explicit way, creates a very general expectation of
widespread, and not necessarily weak, conditional altruism
in nature.


Altruism via kin-selection strategies that rely on arbitrary tags with which they coevolve
R Axelrod, RA Hammond, A Grafen
http://deepblue.lib.umich.edu/bitstream/handle/2027.42/72180/j.0014-3820.2004.tb00465.x.pdf?sequence=1&isAllowed=y

RCB said...

You're right that those Taylor models don't allow kin recognition. I don't know much about kin recognition models, myself. I only know there has historically been some debate over the importance of kin recognition in general, and how it can work theoretically (some models support it, some don't). Found two more recent reviews, should be helpful:
http://www.sciencedirect.com/science/article/pii/S0960982207017083
http://link.springer.com/chapter/10.1007/978-3-642-02624-9_3#page-1
I can't claim any expertise here.

The models we've discussed are still relevant because they define the scope of altruism that is adaptive. That is: if my arguments regarding the scope of competition (i.e., mostly within ethnic-group) are right, and if altruism based on kin recognition did evolve in humans, then it mostly evolved to help people distinguish close relatives (or bandmates) from others. If, say, a large, foreign, mostly endogamous population is suddenly transported into/near your neighborhood, then this evolved kin-recognition might misfire, making you think that your greatest competitors for mates (your coethnics) are actually close relatives. This would probably be maladaptive; presumably, in the long run, selection would tone down the mechanism such that you wouldn't get caught up on ethnic differences. In the meantime, of course, you'd have many generations of ethnocentrism!

M said...

Wow. I love this stuff, even though I barely understand it.

I sense in my dimness that RCB and crew are technically right, but side with n/a, anyhow. I didn't want to be replaced. And it definitely "hurts" me!

n/a said...

M,

No, I'm right, and your intuition is correct.

The articles RCB brings up have little to do with the questions at hand.

The Taylor papers, for example, seem to be intended to model insects, in one case, and plants in the other, in the absence of any sort of kin recognition. These are primarily of interest to people, e.g., studying the evolution of eusociality in insects (and specifically, whether or not limited dispersal in and of itself favors cooperation). Even without kin recognition, these models don't rule out the evolution of cooperation. Which is good, considering eusocial insects do exist.

If you want to look at the evolution of ethnocentrism in humans, you create a relevant model. When Robert Axelrod did, he found:

The main result of the simulation is that the ethnocentric strategy becomes common even though, unlike previous models,3 favoritism toward similar others is not built into the model. In the final 100 periods of ten 2,000-period runs, 76 percent of the agents have the ethnocentric strategy, compared to 25 percent if selection had been neutral (Table 1, row a). This result shows that in-group favoritism based on simple tags and local interactions can overcome egoism and dominate a population even in the absence of reciprocity and reputation and even when "cheaters" need to be suppressed.

This too is good, since, in the real world ethnocentrism obviously exists.

But even if we pretend ethnocentrism does not exist in humans, or that it's impossible for ethnocentrism to have evolved via group selection / kin selection (and, again, it's not -- RCB is fixated on an irrelevant technical discussion and Cochran simply "knows" group selection can't be an important force in humans), this still doesn't change the fact that ethnic genetic interests exist. Cochran and JayMan failed to read "On Ethnic Genetic Interests" (freely available at
https://archive.org/details/OnGeneticInterestsFamilyEthnicityAndHumanityInAnAgeOfMassMigration2006ByFrankKempSalter ) or even know the definition of what they were asserting didn't exist.

Also see:
http://racehist.blogspot.com/2015/03/hamilton-on-inclusive-fitness-and.html
http://racehist.blogspot.com/2015/03/more-from-hamilton-on-kin-recognition.html
http://racehist.blogspot.com/2015/03/r-fisher-on-group-selection-in-humans.html

Anonymous said...

"but I think he has an emotional investment in opposing Ethnic Genetic Interests so he's not the most rational on this topic."

He's a cuckservative