However, common among people who have a superficial and/or selective understanding of heritable group differences is belief in conveniently inaccurate claims. One of these erroneous ideas that White nationalists in particular have latched on to is the belief in “ethnic genetic interests” – that is, that kin selection has led individuals to favor people of their own race/ethnic group over others. This of course is bunk. Natural selection doesn’t work that way, since individuals within an ethnic group aren’t closely related enough for this to work. This has been explained repeatedly, lately by Misdreavus:It is impossible for such a thing as a “race altruist gene” to evolve, because sacrificing yourself on behalf of strangers does nothing to increase the frequency of the gene under any set of circumstances. It doesn’t matter if the frequency of a such a gene “magically” originated with a frequency of 4 in 10 Chinese people. The Chinese who don’t have the gene, on average, would have a higher fitness, resulting in the frequency decreasing monotonically over time.He continues to argue there, which is worth a read for anyone seriously interested in the matter.
After anyone seriously interested in the matter gets done studying the confused rantings of JayMan's gay sidekick, I would recommend they at least take a few minutes to skim the work of the man who coined the term "ethnic genetic interests". Doing so would have saved JayMan and misdreavus at least some measure of embarrassment.
A copy of Frank Salter's On Genetic Interests: Family, Ethnicity, And Humanity In An Age Of Mass Migration is freely available online.
If you want to talk about ethnic genetic interests, read it.
The actual definition of ethnic genetic interests
Frank Salter defines "ethnic genetic interests" as:
The number of copies of a random individual'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, especially from different geographical races, ethnic genetic interest can be many orders of magnitude greater than familial genetic interests.
To deny that ethnic genetic interests exist is to deny that human population structure exists (or to hilariously misunderstand basic population genetics). Ethnic genetic interests exist regardless of whether or not one believes group selection has played any role in human evolution and regardless of whether or not people naturally favor others from their own group.
The issue of the degree to which group selection or kin selection favored the evolution of ethnocentric altruism in humans has no bearing on the reality of ethnic genetic interests. It's a separate issue, on which JayMan/misdreavus are also wrong (one can debate the issue, but not on the confused grounds JayMan and misdreavus have attempted to debate it).
Misdreavus's confusion about coefficients of relatedness
One area of confusion for misdreavus in the linked thread:
The coefficient of relatedness between a Swede and a non-related Swede is zero. The coefficient of relatedness between a Swede and a black African is also zero. You simply do not seem to understand this.I replied at the time:
This is exactly the misapprehension I just got done correcting for JayMan: http://racehist.blogspot.com/2015/02/kinship-coefficients-and-ethnic-genetic.html
The coefficient of relationship is simply twice the coefficient of inbreeding between the hypothetical children of two individuals. Inbreeding is defined relative to some population. It’s often convenient to disregard non-recent inbreeding in calculating coefficients of relationship, but this only makes sense with respect to a particular [approximately random-breeding] population, and, holding the base population constant, a Swede absolutely does not have the same coefficient of relationship to a sub-Saharan as to another Swede. Nor is there any difference in kind between the type of relatedness indicated through Fst and the type shared by close family members.
Even after having it explained to them again, Misdreavus (and apparently JayMan) still failed to understand this very basic concept.
Misdreavus's confusion about the viability of genes for altruism
This is another extremely basic issue: frequencies of genes harmful to an individual's fitness with respect to his group can increase globally if his group expands relative to other groups. If misdreavus had read and understood pretty much anything written about altruism and kin selection or group selection within the past half century, he would have grasped this.
If groups with high frequencies of ethnocentric people expand at the expense of groups with low ethnocentrism, genes for ethnocentric altruism can increase in frequency.
Even misdreavus's (anti-group-selectionist) idol understands this (or did at one point):
Imagine that in much of history, people lived in small groups that often fought with their neighbors. In that sort of situation, selection for group altruism is at least possible, since the group is full of close relatives, while the opponents are less closely related. Both sides are probably members of the same broad ethnic group or race, but that doesn’t matter : only the kinship coefficients matter.Related:
- Kinship coefficients and Ethnic Genetic Interests (JayMan embarrassing himself again)
- Remedial population genetics for Greg Cochran
- Hamilton on inclusive fitness and social behavior in humans
- More from Hamilton on kin recognition and intergroup hostility
- R. A. Fisher on group selection in humans
- Robert Axelrod on the evolution of ethnocentrism
- Gender and Politics Among Anthropologists in the Units of Selection Debate
But ''all traits are genetically inherited''...
ReplyDeleteThe definition claims "The size of ethnic genetic interest is relative to the kinship of genetic competitors." This makes sense to me: group selection mathematics emphasizes the importance of within-group variation to between-group variation, and how this is patterned with respect to competition.
ReplyDeleteYou then say "To deny that ethnic genetic interests exist is to deny that human population structure exists." In light of the above, though, one could also deny ethnic genetic interests - or, at least, say they are very weak - by asserting that the strength of selection/competition between ethnic groups is usually quite small - to small, when multiplied by the between-group variance, to play much role in genetic evolution of human behavior. This may not always be true, especially if we're talking about very small ethnic groups.
It seems your main point here is to say that (1) genetic structure exists at ethnic levels and (2) at least *some* amount of competition exists at these levels. Therefore ethnic genetic interests, sensu Salter, exist, by definition. Great. But, you also say that this is a separate debate from the question of whether people would have been selected to be innately ethnocentric. That is, the existence of ethnic genetic interests sensu Salter does not imply the existence of actual psychological and behavioral ethnic genetic interests in humans. It seems like the latter is what we really care about.
From an evolutionary perspective, Salter's definition seems to miss the mark a bit. The relatedness coefficient in Hamilton's rule is about the regression of allelic value of one's neighbor on oneself, relative to competitors. So it's about relative gene *frequencies*. The fact that Salter talks about "*number* of copies" and discounts kin suggests to me that he doesn't quite understand inclusive fitness - but maybe I should read him more closely. So, we can certainly ask whether the concept he defines exists, but it will have little to do with evolution. So, again, I'm not sure why I should care about this. The theory of inclusive fitness and group selection already identify the mathematical quantities that we should care about. What does Salter add?
Please, comment on this article:
ReplyDeletehttp://www.unz.com/pfrost/the-jews-of-west-africa/
RCB,
ReplyDeleteIn the introduction to the Transaction edition, Salter clarifies:
I now turn to two technical points. One commentator has identified a terminological error that needs to be corrected (David B. 2005c, point 1 ). Throughout On Genetic Interests I define an individual's genetic interests as consisting of his or her distinctive genes, some of which are also found in kin and fellow ethnics. Such genes exist, but kinship generally consists of shared frequencies of genes, even when the genes involved are not unique to any one individual. 'Distinctive genes' should be read as 'distinctive gene frequencies' or 'distinctive allele frequencies.'
"discounts kin"
Salter's definition of "ethnic genetic interests" excludes close kin because he distinguishes between "ethnic" and "familial" genetic interests. Read the book.
"That is, the existence of ethnic genetic interests sensu Salter does not imply the existence of actual psychological and behavioral ethnic genetic interests in humans. It seems like the latter is what we really care about."
No, the first issue is more fundamental than the second.
"No, the first issue is more fundamental than the second."
ReplyDeleteHaving skimmed the first few chapters of Salter, I'm convinced that the book is mostly worthless if you're interested in the evolution of actual behavior. If you're interested in an arbitrary moral philosophy, though, then it's as good as any other.
Take this choice morsel:
"The genetic distance between English and Bantu is so great that, on the face of it, competition between them would take within-group altruism among random English (or among random Bantu) almost as adaptive as parent-child altruism, if the altruism were in the service of that competition. Thus it would appear to be more adaptive for an Englishman to risk life or property resisting the immigration of two Bantu immigrants to England than his taking the same risk to rescue one of is own children from drowning, unless the immigrants were bringing qualities of such economic value that they would permanently raise the Island's carrying capacity. "
Now what could he possibly mean by this? If he means what is usually meant by "adaptive" - that selection would favor an allele causing altruism toward other Englishmen, and malice toward Bantus - then the statement is incorrect. Take a more extreme example: say an allele arose in England that caused Englishmen to take assault rifles to Kenya and shoot up unsuspecting villages. Let's say such men could kill off a good 100 folks before being killed themselves. By Salter's definition, this would easily be beneficial to the Englishman's genetic interests. But clearly the allele will kill itself off whenever it arises: not selected for. (The drowning vs. immigration example he lists is also nonsense: an allele that expends a cost to resist even 100 Bantu immigrants gains nothing relative to those that don't.) Hell, even if the allele only produced this behavior 10% of the time (therefore not dying off immediately), its ultimate fate would still be extinction.
Maybe he has another definition of "adaptive" in mind. Maybe he even defines "adaptive" as "increasing ethnic genetic interests." That would be very convenient.
You're a smart guy, and you seem to think that this "ethnic genetic interests" concept is valuable. Help me understand why.
RCB,
ReplyDelete"Now what could he possibly mean by this? If he means what is usually meant by "adaptive" - that selection would favor an allele causing altruism toward other Englishmen, and malice toward Bantus - then the statement is incorrect."
Salter is correct.
"Take a more extreme example: say an allele arose in England that caused Englishmen to take assault rifles to Kenya and shoot up unsuspecting villages. Let's say such men could kill off a good 100 folks before being killed themselves. By Salter's definition, this would easily be beneficial to the Englishman's genetic interests."
No, killing people in a foreign land would not serve one's ethnic genetic interests (unless one is heading off a threat to one's own territory or facilitating the expansion of one's ethny into the foreign territory), since the dead Kenyans in this example would simply be replaced with other Kenyans, up to whatever the carrying capacity of Kenya is.
"But clearly the allele will kill itself off whenever it arises: not selected for."
The point you're missing here is that the English-Bantu FST is a statement about how likely it is that if I'm an Englishman and I have a given allele, another Englishman also has that allele (relative to a Bantu). The typical allele will not be a newly arisen one. Out of six billion bases and millions of SNPs, we inherit perhaps a few dozen de novo point mutations.
This is another of the things misdreavus had trouble wrapping his head around. The same sort of argument could be used to "prove" altruism between siblings or close cousins could never have arisen (every new mutation exists at first in only a single individual).
"(The drowning vs. immigration example he lists is also nonsense: an allele that expends a cost to resist even 100 Bantu immigrants gains nothing relative to those that don't.) Hell, even if the allele only produced this behavior 10% of the time (therefore not dying off immediately), its ultimate fate would still be extinction."
Again: see the point above about FST, and the second point of misdreavus confusion discussed in the point.
1. If we pick a random base in an Englishman's genome to be our Bantu immigration resistance allele, chances are the Englishman will be more similar to other Englishmen at that base than to Bantu (meaning this particular allele could be adaptive). Averaging over a large number of alleles (and traits of this sort will in reality be polygenic), it's a certainty that the Englishman will be more similar to other Englishmen than to Bantu (and thus, from the perspective of an individual Englishman, it would always be adaptive, depending on the sacrifice involved, to resist the replacement of Englishmen with Bantu).
2. Your frame of reference here needs to expand beyond England. If the English lived in total isolation, then yes sacrifice on behalf of the whole group would never be adaptive. But we live in a world with many competing groups. The English do not live in isolation, and the example we're discussing specifically involves migration and intergroup competition.
Alleles that are "maladaptive" within one's own group can be globally adaptive (in the context of intergroup competition), and groups that allow asymmetric immigration will be replaced. In this example, if instead of Bantu immigration resistance alleles (which you imagine will inevitably be driven to extinction by inter-English competition) the English carry only immigration non-resistance alleles, the immigration non-resistance alleles (along with all other English alleles) will tend toward extinction much faster in the face of unchecked Bantu immigration than would immigration resistance alleles attuned to Hamilton's rule in the face of thwarted Bantu mass immigration.
"The same sort of argument could be used to "prove" altruism between siblings or close cousins could never have arisen"
ReplyDeleteIt's true that self-sacrificial altruism will die off whenever it arises, even if it's among siblings. That's why Hamilton's Rule is only strictly true for weak selection. But it's not the same problem. As long as the altruistic allele can make it through the first few generations (so not self-sacrificial), then altruistic alleles for close kin will be selected for.
That's not true of the killer racist example. (Given your complaint, I'm going to now say that the racist Englishman allele kills immigrant Bantus in his own country. It still doesn't work.) Note that I said "suppose the allele only kills off foreigners every 10% of the time" (imperfect penetrance), to account for the weak selection. Now let's go further and imagine that 50% of the English population were magically mutated to have this allele (and Bantu have 0%). Strong F_ST! What happens? All copies of the allele quickly kill themselves off in a few generations. Selection eliminates the allele.
I recognize that English have higher genome-wide similarity to each other than Bantu. It's irrelevant to the evolution of selected loci.
I understand the frame of reference thing. Perhaps the English mutation causes one Englishman to kill off thousands of Bantu, and therefore, on the global scale, the allele actually increases in frequency for a bit (again, we have to assume that the allele is somehow at high frequency in England). That's cute, but wait around a few generations and see what happens. It goes it extinct in England. The problem you have is that you magically assume that a disfavored allele has somehow gotten to high frequency in one population but not another. In fact, it would never have gotten to high frequency. (It could drift there, but that's not an adaptive argument. Selection will still kill it off at equilibrium.)
Your immigration-resistance example is dumb. Resisting immigration is a common good: presumably everyone in English suffers the "cost" of having Bantus move in (whatever that is...). Those who pay the cost of resistance get nothing more than those who don't do anything. The allele is disfavored.
Let's try another example. Suppose there are two neighboring ethnic groups that are somewhat genetically diverged. But let's say that ethnic-level altruism is not selected for under these particular conditions: the combination of divergence + intergroup competition isn't quite strong enough to favor ethnic altruism at equilibrium. Whenever such an allele arises, it tends to die off in a few generations. Now suppose we take one group and mutate it to have completely different alleles at millions of loci across the genome. Many of the loci can be functional. But assume that the basic dynamics of migration and mating and competition are unchanged. Does this change the situation? If I were Salter, I would say probably yes: the ethnic groups are way different now, so selection could plausibly favor ethnic level altruism now. But the actual answer is no: since the dynamics of migration and mating have not changed, the force of selection on any novel ethnic-altruism allele is unchanged. It will tend to die off whenever it arises, just as before.
Seriously, man, it's time for you to drop Salter. The book is bullshit. There is actual population genetic theory of social behavior out there. Salter is not in that category.
Also your "carrying capacity" retort is nonsense. Imagine that the racist Englishman allele doesn't just kill 100 Kenyans, but permanently lowers their carrying capacity by that amount. The result? You guessed it: the allele quickly kills itself off.
ReplyDeleteWhether it's possible for ethnocentric behaviour to evolve and maintain stable allele frequencies is irrelevant to the existence of ethnic genetic interests. Suppose I have 1 million dollars in the bank. How it got there and what my spending habits are do not change the fact that I have 1 million dollars.
ReplyDeleteRCB,
ReplyDelete"Also your "carrying capacity" retort is nonsense. Imagine that the racist Englishman allele doesn't just kill 100 Kenyans, but permanently lowers their carrying capacity by that amount. The result? You guessed it: the allele quickly kills itself off."
It still doesn't matter if he "permanently lowers their carrying capacity", again, unless he is eliminating a threat to his own ethny or some of the territory or resources that would have been used by those killed go toward members of his own ethny instead.
If Kenyans are threatening the English or the English are colonizing Kenya, it may be adaptive for an Englishman to kill Kenyans. If one of those two conditions does not hold, it would not be.
"That's why Hamilton's Rule is only strictly true for weak selection. [. . .] Note that I said "suppose the allele only kills off foreigners every 10% of the time" (imperfect penetrance), to account for the weak selection."
This does not begin to approach the type of weak selection we would be looking at in any realistic model for a trait like this. No doubt thousands of loci influence traits like ethnocentrism, with each tending to account for less than 1% of variation in the trait.
"But it's not the same problem. As long as the altruistic allele can make it through the first few generations (so not self-sacrificial), then altruistic alleles for close kin will be selected for."
Yes, it's exactly the same problem. It's easy to imagine variants favoring ethnocentric altruism drifting to high frequency in the sorts of small, inbred groups that formed the basic units of organization among humans for most of their existence, and groups with high levels of these traits outcompeting those with low levels.
"Now let's go further and imagine that 50% of the English population were magically mutated to have this allele (and Bantu have 0%). Strong F_ST! What happens? All copies of the allele quickly kill themselves off in a few generations. Selection eliminates the allele."
You think 50% of the English would need to die to repulse Bantu immigration? Bantu immigration could be halted and reversed at much closer to a cost of 0 English lives, while unchecked Bantu immigration will drive the frequency of all English alleles toward zero.
And I specified immigration resistance alleles attuned to Hamilton's rule. This means, by definition, the person carrying these alleles would be sacrificing less than they would be gaining (in terms of preventing the replacement of genes similar to their own) in resisting the Bantu immigration. And, conversely, every Bantu that enters England is costing the English more than the Englishman would be sacrificing in preventing it. Your only recourse here is to deny that the English could ever realistically control their border at a less than excessive cost. But this is of course absurd, particularly in this example (given the disparity in technology and military ability between the English and Bantu, the geographic locations involved, etc.). And the fact that territoriality is a human universal would tend to argue against the notion that it could never be adaptive.
"but wait around a few generations and see what happens. It goes it extinct in England."
Not as quickly as it (or, rather, a non-functioning version of it) would go extinct if it didn't exist. And if it also helps the English expand into new territories at the expense of groups with lower frequencies (rather than the English simply staying in England and continually repulsing attacks), there's no reason the global frequency would need to decline at all.
"If Kenyans are threatening the English or the English are colonizing Kenya, it may be adaptive for an Englishman to kill Kenyans. If one of those two conditions does not hold, it would not be"
ReplyDeleteCertainly if an Englishman is at personal threat of being killed by invading foreigners, he should probably defend himself! But any costly effort to protect his ethnic group *as a whole* from invading foreigners is not going to be selected for, since all other Englishmen benefit from that action. (It's a "public good.") (Note that at no point did I mention the case of hostile invaders encroaching on England, by the way.)
"Yes, it's exactly the same problem."
No, it's not. Let me clarify. A novel, positively selection sib-altruism mutation faces a problem in the first generation: it's novel, so no kin share it! With luck, and if the cost is not too great, the allele will make it to the next generation, and then sibs *will* have the allele. If rb>c (taking usual assumptions), the allele will tend to spread from there. This is not the case for the suicidal racist Englishman allele. Even if, against all chances, the allele drifted to appreciable frequency at home, those who have the allele are going to be selectively disfavored (and quickly kill themselves off), because the benefits it provides are also enjoyed by the non-racist English alleles. This is true regardless of how many Kenyans such men manage to kill abroad or at home.
"You think 50% of the English would need to die to repulse Bantu immigration? "
No. I never said that, because it's irrelevant.
"This means, by definition, the person carrying these alleles would be sacrificing less than they would be gaining (in terms of preventing the replacement of genes similar to their own) in resisting the Bantu immigration."
This sentence basically sums up your lack of understanding about this whole issue. Selection doesn't favor individuals who "prevent the replacement of genes similar to their own." Recall the fact that genome-wide similarity is irrelevant to all of this. What would the *allele* be gaining? Selection will favor an allele if it can selectively bring more benefit to other copies of that allele than to alternative alleles (and at cost to itself). But helping your ethnic group helps all home alleles equally, meaning that selection will always drive the allele to extinction within the group.
"And the fact that territoriality is a human universal would tend to argue against the notion that it could never be adaptive."
Don't be stupid. Lots of animals are territorial. You don't need to be ethnically-minded to be territorial. Individual selection can explain why animals don't like others encroaching on their territory. (I don't doubt that ethnocentrism exists, by the way. My point is that Salter's incoherent theory does nothing to explain this phenomenon.)
"Not as quickly as it (or, rather, a non-functioning version of it) would go extinct if it didn't exist."
What you're talking about here, I think, is that invading enemies would have an easier time killing off people who were not ethnically interested. I don't doubt that. But that doesn't mean that selection will favor the ethnic altruism allele. See "public good" explanation.
RCB,
ReplyDeleteIt makes no difference if the Bantu immigration is "hostile" or not. If England is subject to unfettered Bantu immigration, allele frequencies in England will converge on those of the Bantu.
If the diploid population is infinite, selection is absent and the immigrant gene frequency is fixed, then the gene frequency on the island converges to the immigrant frequency
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1213928/
"But any costly effort to protect his ethnic group *as a whole* from invading foreigners is not going to be selected for, since all other Englishmen benefit from that action."
You're having trouble grasping the fact that in the example we're discussing the genetically distant immigrants represent a greater threat to the Englishman's fitness than inter-English free-riding.
This is true by definition, in the problem we're discussing. So your options are (1) deny the problem could ever exist in reality (that groups could ever meaningfully compete or that immigrants displace natives); or, again, (2) deny that a country like England could ever control it's border at a cost lower than the benefit.
"the allele will make it to the next generation, and then sibs *will* have the allele."
No, there is no certainty siblings "*will* have the allele." It's still a statistical question.
"No. I never said that, because it's irrelevant."
No, it's not irrelevant. It's all that matters for this discussion. I don't care about your "suicidal Englishman" example. If the suicidal Englishman prevents the displacement of enough Englishmen by Bantu, then his act is adaptive. If not, it's not. But the question is whether it's possible to act adaptively in resisting immigration / ethnic displacement.
"What would the *allele* be gaining?"
Again, FST is both a statement about genome-wide similarity and about the chance that any given locus will be shared identical by descent relative to the outgroup. The allele will be gaining better odds that copies of itself survive.
"But helping your ethnic group helps all home alleles equally, meaning that selection will always drive the allele to extinction within the group."
Again, not to the extent that intergroup competition is a meaningful force and the allele is attuned to Hamilton's rule.
"What you're talking about here, I think, is that invading enemies would have an easier time killing off people who were not ethnically interested."
As I explained above, it doesn't matter if immigration is "hostile" or not. Asymmetric immigration leads to population replacement either way.
To the extent immigration resistance alleles are attuned to Hamilton's rule (so that whatever cost is expended is more than made up for in benefit), by definition they will be more adaptive than the alternative of allowing unfettered, non-reciprocal immigration by genetically distant groups.
More from Salter:
ReplyDeleteIn a recent study of biochemical polymorphism in farm populations of the house mouse, Petras 48 found no evidence of other than random mating within local groups, but on the basis of gene-frequency differences between groups he estimated FST = 0.18. Substituting in the formula explained in Appendix B (page 221) this gives b = 0.305, which shows that within groups mice should treat the average individual encountered as a relative closer than a grandchild (or half sib) but more distant than an offspring (or full sib), referring to an outbred population. [. . .]
APPENDIX B
Viscous and subdivided population models
An outline of this analysis can be introduced by quoting an identity, which Wright proves:
(1 - FIT) = (1 - FST)(1 - FIS)
In this identity, FIT is the correlation of uniting gametes relative to the array of gametes of the whole population, FIS is the correlation of uniting gametes relative to the array of gametes of their own subdivision, and FST is the correlation of gametes drawn randomly from a subdivision relative to the array of gametes from the whole population.
For evaluation of b from equation (1) (page 213), FIT must be used in the calculation of both rAB and FA. If instead FIS is used, an inclusive coefficient is obtained relevant only to selection within the subdivision. It could easily happen that an altruistic trait that was 'too generous' to increase in frequency within its subdivision (except by drift), increases, nevertheless, in the population as a whole due to the more rapid expansion of those subdivisions which contain altruists in higher frequencies.
Oh wait that's Hamilton.
You're quoting Hamilton like I don't understand multilevel selection. I do. I'm a coauthor on a paper that favorably reviews group selection - in humans! Salter isn't in the same league as Hamilton. Not even close. He shows no evidence of understanding this. His book amounts to little more than moral philosophy with some genetic statistics sprinkled in.
ReplyDeleteI understand that certain models can maintain enough genetic variation among groups to counteract the group-variation-killing forces of within-group selection, between-group selection, and migration to maintain significant group selection at equilibrium (they are strict conditions - usually very small groups with lots of fissioning so as to constantly restock between-group variation - could work for HG bands). Does Salter show any appreciation of this? I don't see it. (I also don't buy that any of the genetic group selection models that do work are applicable to large human ethnic groups (like "The English), but I'm less convinced there.)
"It makes no difference if the Bantu immigration is "hostile" or not."
It's funny, because I actually started all of this with examples of non-hostile immigration. You then brought up the example of "If Kenyans are threatening the English...", so I switched to that. Your argument is hard to pin down.
"If the suicidal Englishman prevents the displacement of enough Englishmen by Bantu, then his act is adaptive."
No. It isn't. All English alleles benefit equally by his act. Therefore the alleles that pay the cost to fight immigrants will be at a fitness disadvantage, and go extinct in England. When an allele is selectively disfavored in *every* group it's in, you need pretty special dynamics to keep it alive! Again, the usual mechanism (Wilson's models) is constant turnover of small groups, with founder effects. This creates strong drift, which in turn maintains the between-group variation that facilitates group selection. This isn't what large scale human ethnic groups look like - certainly not at the scale of nations. Where is the discussion of this in Salter? (That's partly a sincere question - maybe it's there and I missed it.)
"You're having trouble grasping the fact that in the example we're discussing the genetically distant immigrants represent a greater threat to the Englishman's fitness than inter-English free-riding."
Let's make this simple. Explain to me how an English allele that incurs reproductive costs to fight off Ethiopian immigrants (of any number) to his country (hostile or not), thereby producing a public good for all Englishmen, will not be selectively disfavored in England. Bonus points if you explain how such behavior is more adaptive than saving one's own kids from drowning (to take from Salters book). If you can do this with a dynamically complete pop gen model, and I can't show that it's inconsistent or obviously invoking additional assumptions that Salter never implied, I'll be particularly impressed and humbled (but probably too proud to admit it). Note: saying "it satisfies Hamilton's rule" isn't enough. Show me *how*.
"To the extent immigration resistance alleles are attuned to Hamilton's rule..."
I agree, of course, that if an allele's action satisfies Hamilton's rule, then it will be selected for. No disagreement there! What I have argued, and you have failed to refute, is that the examples I suggest (and most of Salters) could not possibly satisfy Hamilton's rule unless you make some heroic assumptions that Salter wouldn't understand. In other words: to say that something is in your "ethnic genetic interests" is, I *assert*, not equivalent to saying that something is in your inclusive fitness interests. At best, "ethnic genetic interests" is a vague genetic concept that encourages people to make stupid conclusions about social evolution. Let's just use real evolutionary theory, please.
Resisting immigration may not (at the moment) be adaptive within the English context. But in a global context, it most certainly is (if successful). What is maladaptive in one context or one point in time may be adaptive in a different context or become adaptive later on.
ReplyDelete"You're quoting Hamilton like I don't understand multilevel selection. I do."
ReplyDeleteYet you earlier appeared to be disputing the relevance of FST to the question of how one "should" behave in intergroup interactions (on the basis of imagining a new mutation and supposing it would necessarily be culled by selection before becoming established in the population).
Hamilton also speculates at length about the effects of group selection in humans, and doesn't hesitate to attribute tribal behavior in humans to selection on intergroup competition.
"(I also don't buy that any of the genetic group selection models that do work are applicable to large human ethnic groups (like "The English), but I'm less convinced there.)"
The question of what would be adaptive will always come down to cost, benefit, and relatedness, regardless of scale.
"You then brought up the example of "If Kenyans are threatening the English...", so I switched to that."
The point I was trying to get across is that what matters (in determining whether or not it is potentially adaptive for an Englishman to kill Kenyans) is that the English and Kenyans are in competition. It doesn't matter if the competition takes the form of warfare or demographic displacement. If the Kenyans represent no military, demographic, or resource-competition threat (and the English are not colonizing Kenya), it would never be adaptive for an Englishman to go to Kenya and kill Kenyans.
"No. It isn't. All English alleles benefit equally by his act. Therefore the alleles that pay the cost to fight immigrants will be at a fitness disadvantage, and go extinct in England."
Again, if the English simply sit in England, turning back wave after wave of immigrants, forever, the immigration resistance alleles would eventually die out. But this would happen much more slowly than the English people would die out if they fail to turn back mass immigration of genetically distant people (which they could have done and still can do at fairly minimal cost). Slightly reduced fitness is preferable to massively reduced fitness.
"When an allele is selectively disfavored in *every* group it's in, you need pretty special dynamics to keep it alive!"
If the allele is not simply an immigration resistance allele, but one that increases ethnocentric altruism in general, it's likely to aid intergroup competition in general (including offense/expansion, not just defense/preservation). In which case, as I've repeatedly noted and as you've said you're fully aware, it could well be favored in the total population.
In reality, the English did not simply sit in England. They colonized much of the world and expanded demographically into new territories.
"Let's make this simple. Explain to me how an English allele that incurs reproductive costs to fight off Ethiopian immigrants (of any number) to his country (hostile or not), thereby producing a public good for all Englishmen, will not be selectively disfavored in England."
ReplyDeleteIt will be disfavored relative to other English alleles in the scenario where the English resist immigration. It will not be disfavored relative to its share in the global population in the alternate scenario where unlimited Bantu immigration is allowed. A smaller share of an existing pie is larger than an equal share of a no-longer-existent pie.
If we clone England, and genetically engineer England 1 with our immigration resistance alleles and England 2 with immigration non-resistance alleles and turn loose flotillas of sub-Saharan Africans on both, the immigration resistance alleles in England 1 will be around long after the immigration non-resistance alleles in England 2.
"What I have argued, and you have failed to refute, is that the examples I suggest (and most of Salters) could not possibly satisfy Hamilton's rule"
If killing Bantu immigrants in England prevents them from displacing enough English (or causes enough other immigrants to leave or turn back), it will be adaptive. Period. (Of course, in reality, an event like this would simply be used by elites as an excuse to further dispossess the English. Actual methods for resisting immigration and repatriating immigrants in the modern world typically are or would be considerably less violent and present considerably less cost to those enforcing them. Given that Europe is likely currently financially subsidizing its colonization by the third world, "costs" of preventing immigration right now may be negative.)
The only way this would not be true is if you believe the immigrants would not be displacing natives, which is essentially the same as saying England can accept unlimited immigration (unlimited population growth, increased competition for resources and mates, etc.) with no effect on the English. Even if we pretend unlimited immigration would not suppress English birth rates below their current levels, the immigrants are coming from countries with higher birth rates, can be expected to maintain higher birth rates in England for some period of time, and have behind them a stream of prospective immigrants that is effectively limitless compared to the ~40 million English in England. Birth rates of the native British are below replacement. The population of Africa is projected to quadruple to 4 billion by the end of the century. If that population growth is constrained to Africa, and Africans maintain themselves with their own resources, the effects on England will be minimal and selection can be expected to eventually pull native British birth rates back up to at least replacement level (while the population of Africa would eventually correct itself to a sustainable level). If the English allow free immigration, the English will be replaced.
"Yet you earlier appeared to be disputing the relevance of FST to the question of how one "should" behave in intergroup interactions"
ReplyDeleteI know the relevance of FST. I know you can rearrange the group-selection version of the Price equation to be framed in terms of FST. I also know that this FST refers to the *selected locus*, not genome-wide FST, as Salter apparently believes. Now you might say "populations with high genome-wide FST are more likely to be diverged at functional loci." In general, yes, but not for alleles that are selectively disfavored in all groups: selection keeps their frequency at ~0 everywhere, so there is no variation for selection to act on (again, unless rapid turnover of small groups constantly replenishes FST at this locus). Ethnic altruism alleles fall into this category, since, by definition, they provide a public good to the ethnic group at a cost.
"Slightly reduced fitness is preferable to massively reduced fitness."
An allele that produces a public good at a cost dies off faster than one that receives the public good at no cost. That's called "selectively disfavored." Whenever there is a polymorphic population, then, the immigration-resisting allele will die first. This is true whether or not the population is being bombarded by immigrants. Get it into your skull.
"In which case, as I've repeatedly noted and as you've said you're fully aware, it could well be favored in the total population"
You have to assume that the allele is at reasonably high frequency in England (but not elsewhere) for this to be true (otherwise the allele is just sacrificing itself to help the other allele - like when it first arises. It's the 0 FST thing again.). Given that it's selectively disfavored within England, you have to wonder how it got to high frequency in the first place, especially in a population of 60 million. A miracle of drift, perhaps! But let's say it happens to be true. Then, yes, if these folks can happen to kill off enough foreigners, the global frequency of the allele can rise for a time. Hurrah! But, like I said before: wait around a bit. Selection will kill the allele off in England: then no one will have it. Remember that the FST Price equation formulation is a one-generation description of evolution, not a model. If you artificially fix FST to be high one generation, you can get the global result you mention for a few generations, until things return to equilibrium. But if you want this sort of large scale group selection to work long-term, you need to somehow produce a system in which the allele sticks around despite being selectively disfavored within every group. I've mentioned a few of those; Salter doesn't know shit about it.
"A smaller share of an existing pie is larger than an equal share of a no-longer-existent pie."
Again, wait around a bit. The share of each pie is decreasing everywhere, always.
"If we clone England, and genetically engineer England 1 with our immigration resistance alleles and England 2 with immigration non-resistance alleles and turn loose flotillas of sub-Saharan Africans on both, the immigration resistance alleles in England 1 will be around long after the immigration non-resistance alleles in England 2."
Perhaps this is how your selectively disfavored alleles got to high frequency! Divine genetic intervention! Yes, of course a group that has more immigration-resistance alleles will better resist immigration. But in no scenario does a novel, costly immigration-resistance allele displace the other. That would have to happen for any of this to work.
"If killing Bantu immigrants in England prevents them from displacing enough English (or causes enough other immigrants to leave or turn back), it will be adaptive. Period."
ReplyDeleteIf a Bantu is try to displace me, it is adaptive for me to kill him. If an Englishman is trying to displace me, it is adaptive for me to kill him. If a Bantu is trying to displace a distantly related countryman, it is not adaptive for me to kill him (because alleles that don't do this still get the benefit of having other ethnic altruists kill the Bantu for them, at no cost). That's what we're talking about.
"If the English allow free immigration, the English will be replaced."
Maybe so. But the ethnic altruist alleles in England will die off faster, because they are selectively disfavored. This will happen whether or not Bantus try to take over England, actually.
I know breaking up is hard to do, but it's time for you to get serious and quit Salter.
RCB,
ReplyDeleteThere's no point in continuing to go over this. Among other things, you are denying the possibility of meaningful group selection in humans. Hamilton disagrees. Fisher disagrees. I disagree.
I may work on some simulations to illustrate a few points, but it's unlikely I'll get to that in the near future.
Like I said, I'm an author on a paper that suggests meaningful group selection in humans, so you're wrong about me. I've disagreed with Salter's particularly crazy ideas, and I think that selection does not currently favor altruism at the level of large ethnic groups, because the math just doesn't work out. For smaller human groups? In the past? Maybe.
ReplyDeleteI'm aware of Hamilton's speculations (in part, due to your sharing, which I appreciate). (Not aware of what Fisher has to say on this point - I thought he was a vehement opponent of group selection, but you probably know better.) At some point, though, you need to stop hiding behind big names and start grappling with the body of population genetic theory that has been produced since 1970. And the arguments I've made here. When you do this, you will realize that Salter is full of shit. (Whether genetic group selection in humans is important is less clear, but none of the arguments you've given actually work at the level of England vs. Bantu.)
For now, I eagerly await the day that you produce a model in which an allele that pays a cost to resist foreign immigrants selectively replaces an alternative allele that provides parental care (Salter's example!).
For now, I eagerly await the day that you produce a model in which an allele that pays a cost to resist foreign immigrants selectively replaces an alternative allele that provides parental care (Salter's example!).
ReplyDeleteIt's perhaps unfortunate that Salter used the term 'adaptive', because it's not adaptive in the narrow sense population geneticists mean. What's needed is an adjective that means 'advantageous from a neo-Darwinian perspective'. That's how he is using adaptive.
And n/a's point about having a smaller share of an existing pie rather than none is important. Again, the behaviour may not be adaptive in the sense you mean it (where the advantage is considered only with regards to fellow Englishmen), but it is advantageous from a neo-Darwinian perspective.
Madison Grant's actions may not have been 'adaptive', but that's a limitation of the definition.
This type of criticism of Salter's work has been going on since it came out. How ethnic genetic interests (essentially, population differentiation) developed, and whether alleles that seek to preserve or expand it are or can be evolutionarily stable is not relevant to the existence of these genetic interest. People attacking EGI based on whether it is adaptive to pursue either miss the point or are ethnically compromised and have an interest in creating distraction (e.g. gnxp subcons and freaks).
Finally, what is adaptive human behaviour is not stable. After 1933, certain behaviour and traits quickly became maladaptive.
Anonymous:
ReplyDeleteLet me state this without "adaptive" semantics: If an allele were to arise right now which caused an Englishman to resist foreign immigrants (e.g. by killing them) at some reproductive cost, the allele would tend to go extinct. Especially if this traded off with not caring for your kids. I don't care how many immigrants are repelled by the behavior. It will die off in England, and hence everywhere in the world.
Now, I don't know any definition of "adaptive" that applies to alleles that always tend toward immediate extinct relative to others. And I would call myself a neo-Darwinian! In fact, I have never heard an evolutionary theorist use it this way. So I think you are making things up.
You would do well to remember *my* "share-of-the-pie" point: when within-group selection is causing an allele's share of the pie to decrease *in every population*, you're gonna have a hard time maintaining it at high frequency! I've mentioned conditions that can make it work (constant replenishment of group variation via fissioning of small groups, with strong between-group competition). Salter is strangely quiet on this matter.
You basically admit at the end that ethnic genetic interests cannot be used to make conclusions about evolutionary outcomes. This is, indeed, my main point. So, what *can* ethnic genetic interests be used for? I see nothing except an arbitrary moral philosophy: Salter has defined a quantity, and has asserted that it is good to preserve it. Great, but don't pretend you're doing evolutionary theory.
"People attacking EGI based on whether it is adaptive to pursue either miss the point or are ethnically compromised and have an interest in creating distraction"
You're wrong: I'm neither one of those things. I'm just a guy who understands evolutionary theory, and who happens to think that "adaptive" means "will tend to spread and replace alternatives in a population." You know, what it actually means.
RCB,
ReplyDelete"For now, I eagerly await the day that you produce a model in which an allele that pays a cost to resist foreign immigrants selectively replaces an alternative allele that provides parental care (Salter's example!)."
That you imagine this is impossible shows you don't really grasp inclusive fitness theory in general.
Altruism, Spite, and Greenbeards [2010]
Stuart A. West* and Andy Gardner*
Hamilton’s theory of inclusive fitness showed how natural selection could lead to behaviors that decrease the relative fitness of the actor and also either benefit (altruism) or harm (spite) other individuals. However, several fundamental issues in the evolution of altruism and spite have remained contentious. Here, we show how recent work has resolved three key debates, helping clarify how Hamilton’s theoretical overview links to real-world examples, in organisms ranging from bacteria to humans: Is the evolution of extreme altruism, represented by the sterile workers of social insects,driven by genetics or ecology? Does spite really exist in nature? And, can altruism be favored between individuals who are not close kin but share a “greenbeard” gene for altruism? [. . .]
Inclusive-fitness theory explains altruism but also has a darker side. Spiteful traits, which are harmful to both actor and recipient, may be favored (4). Considering the classic two-party version of Hamilton’s rule, if c is positive (which is costly to the actor), and b is negative (which is costly to the recipient),then rb–c>0 can be satisfied if relatedness (r) is negative. Negative relatedness may seem a bizarre concept, but it simply means that the recipient is less related to the actor than is an average member of the population (SOM text).
Another way to think about spite is to distinguish between the primary recipient of the harming behavior (the individual physically attacked) and those secondarily influenced as a byproduct of this (those experiencing reduced competition from the harmed individual) (16,17). From this perspective, spite can be favored if the actor is more related to the secondary recipients (who benefit) than to the primary recipients (who are harmed). Spite can therefore be thought of as altruism toward the secondary recipients: Harming an individual can be favored if this provides a benefit to closer relatives (Fig. 2C). These two different encapsulations of spite are different ways of looking at the same thing, using either a two- or three-party Hamilton’s rule (SOM text) (16, 17). If the actor is more closely related to the second- ary recipients than the primary re- cipients in a three-party rule, then this leads to a negative relatedness in the two-party rule.
[. . .] recent theory has shown that a positive relatedness to secondary recipients can be obtained more easily than previously thought, suggesting that at least from a theoretical perspective spite is plausible (19) if there is (i) large variance in relatedness between competitors; (ii) kin discrimination, with harming behaviors aimed at individuals to whom the actor is relatively unrelated (making the actor relatively more related to the secondary recipients); or (iii) strong local competition so that harming the primary recipient provides appreciable benefits to secondary recipients. When these conditions are met, spiteful behaviors can be favored so as to reduce competition for relatives. Consider the extreme example of when two genetically identical (clonal) individuals are
competing with a nonrelative to whom they are unrelated. One of the clone-mates could be selected to harm the nonrelative, at a cost to itself, if this harming reduces the nonrelative’s ability to compete for resources and hence benefits the other clone-mate.
http://faculty.chas.uni.edu/~spradlin/SandE/Readings/altruism%20spite.pdf
"start grappling with the body of population genetic theory that has been produced since 1970."
ReplyDeleteDavid Queller [1994]:
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.
http://www.researchgate.net/profile/David_Queller/publication/226956654_Genetic_relatedness_in_viscous_populations/links/0f3175367c85930dbf000000.pdf
On Indirect Genetic Effects in Structured Populations [2001]:
If individuals are distrib- uted into social groups or physical neighborhoods, then population subdivision exists, and interactions may be more likely to occur within demes than among them (Wade 1996). Such population structure is a biological reality for many, perhaps most, organisms (Loveless and Hammrick 1984; Avise et al. 1987; Whitlock 1992; Hanski and Gilpin 1997; Kelly 1997; Wade and Goodnight 1998) and allows for selection to act at multiple levels. Other approaches to the study of social evolution have shown that population structure can be important (e.g., the ev- olution of cooperation in viscous populations [Taylor 1992; Queller 1994a, 1994b; van Baalen and Rand 1998]). Wade (1982, 1985) showed the relationship between ge- netic subdivision, F ST , and relatedness, r, demonstrating that population structure provides another way to satisfy Hamilton’s rule.
http://faculty.virginia.edu/brodie/files/publications/strpopiges.pdf
Alan Grafen [1990]:
ReplyDeleteThe Evolution of Kinship Recognition Systems
The substance of the second principle quoted from Hamilton (1964) in the introduction was that kinship recognition systems should evolve in such a way that organisms treat each other according to their ancestral relatedness. Hamilton himself gave no special justification for asserting that ancestral relatedness would be the important quantity, but treated it as a corollary of his general theory of inclusive fitness. I believe this position to be wholly sound, and so some justification must be given for providing a longer and more explicit argument here. The difficulty arises when kin recognition occurs by a process of matching, as defined above.
The difficulty is that the process of matching creates genetic similarity between an individual and those it 'recognizes' as kin, over and above any genetic similarity that arises from common anestry. The question is then whether Hamilton's rule should apply with the ancestral relatedness as Hamilton claimed originally, or whether it should apply with an inflated relateness which takes into account this extra genetic similarity brough about by the process of matching. [. . .]
The above arguments show that selection on the whole system of kin recognition by matching will evolve close to the way suggested by Hamilton's principles. The using loci will evolve so that other individuals are treated according to their ancestral relatedness, and the matching locus will evolve towards a high degree of polymorphism so that the picking out of relatives from a mixed group of relatives and non-relatives will become very reliable.
[. . .] The first consequence to be drawn is that even when kin recognition occurs by matching, the resulting behaviour will be in accord with Hamilton's rule using ancestral relatedness. The enhanced relatedness at the matching locus does not affect the behaviour unless the using loci are very tightly linked to the matching locus (a green beard gene). There is not reason to expect this to occur in general.
http://users.ox.ac.uk/~grafen/cv/kinrec.pdf
Also, while I do believe you're arguing in good faith and that you believe you're thinking about these questions objectively, I think if you engage in a bit of self-examination you'll find you probably had little resistance when reading about Hamiltons's "spite" in the abstract. Yet you were completely incredulous at Salter's example in humans. So I don't think your refusal to seriously consider Salter is entirely unrelated to political preconceptions.
ReplyDelete"That you imagine this is impossible shows you don't really grasp inclusive fitness theory in general."
ReplyDeleteRemember, I don't think selection for group-beneficial behavior on the basis of population differentiation is impossible. I understand group selection. But Salter's examples in the context of English and Bantu (and other nation states) are ridiculous. It doesn't work.
Your collection of random quotes does nothing to answer the very simple challenge I gave you: explain how a novel allele that pays a reproductive cost to resist foreign Bantu immigrants to England would not tend to die off within a few generations (Salter's example). Do it without invoking heroic assumptions about small group size and frequent fissioning, which Salter never mentioned. You can quote people about spite and negative relatedness all you want (which I do understand, btw), but you still have not shown how this can work in that context. If you don't do this in the next response, I'll assume it's because you can't.
I recall that Queller's "relatedness in viscous populations" is aimed primarily at explaining why "sticking close to home," which increases deme FST, does *not* in general yield altruism among the demes, as it only serves to increase competition *within* the demes. See the quotes: "Hamilton argued that limited dispersal would lead to what he called population viscosity, elevating local relatedness sufficiently to allow altruism towards neighbours in general. Recently it has been discovered that this second mechanism does not necessarily work (Taylor, 1992a,b; Wilson et al., 1992)." And "But it has only recently been shown, by computer simulation (Wilson et al. 1992) and analytical models (Taylor, 1992a,b), that the relatedness-enhancing and competition- enhancing effects of viscosity population exactly cancel under a variety of viscous population structures." This makes sense: when you stick close to home, your competitors are all co-ethnics. So, I'm not sure what your point is in quoting this paper. (Again, I recognize that there are conditions under which group selection can be maintained in the long term, so don't respond like I'm arguing against that fact.)
Given that your strategy usually involves picking quotes that sound good, I've got another challenge for you. Send emails to Grafen, Queller, West, and Gardner. You can find their addresses online, no doubt. CC me in the email - you have my email address. Ask them what they think about Salter's work. Ask them specifically about the example where he says that killing foreign Bantu invaders is more adaptive than caring for your own kids. Ask them if they think "ethnic genetic interests", he defines it, is a useful concept. If you get positive reviews, I'll leave you alone, and you can brag about the victory on your blog.
As to my political preconceptions - I don't think I've given any indication as to what they are. Disagreeing with you doesn't make me a Marxist. The fact is I've spent a lot of time getting to understand the intricacies of Hamilton's rule and group selection, so when I see people completely botching them (which is easy to do), I speak up.
Why don't you e-mail Harpending and ask what he thinks of Salter's work?
ReplyDeleteexplain how a novel allele that pays a reproductive cost to resist foreign Bantu immigrants to England would not tend to die off within a few generations
ReplyDeleteAnd while we're at it, let's determine how many angels can dance on the head of a pin!
Do you ever consider how retarded the model is that you're requesting.
You and Cochran and assorted types think there's a 'Breivik gene'.
Far more likely that the 'Breivik phenotype' is an outlier for some other complex behaviour, which could well be adaptive.
"Why don't you e-mail Harpending and ask what he thinks of Salter's work?"
ReplyDeleteI could; I have chatted with him a bit on other pop gen topics. In general, of course, appeal to authority is a bad way to make arguments. I only suggested those other names because n/a's defense rests on a collection of quotes from them. I doubt they would actually support Salters ideas - if they did, I would take it up with them.
"Do you ever consider how retarded the model is that you're requesting."
It's not my idea. It's Salters: "Thus it would appear to be more adaptive for an Englishman to risk life or property resisting the immigration of two Bantu immigrants to England than his taking the same risk to rescue one of his own children from drowning." I agree it's stupid. Comically so.
"Far more likely that the 'Breivik phenotype' is an outlier for some other complex behaviour, which could well be adaptive."
Sure. But then ethnocentric altruism would be a maladaptive byproduct of selection for other behaviors. Not adaptive, as Salter claims.
Not adaptive, as Salter claims.
ReplyDeleteIt's not necessarily adaptive. But again, it doesn't matter for the existence of EGI. That's the point you're missing. Read the example about the bank account.
This statement was reasonable at some level:
I'm convinced that the book is mostly worthless if you're interested in the evolution of actual behavior.
But the evolution of behaviour is not a main topic of the book. I don't even remember it being discussed, but maybe it was somewhere.
And when you say things like:
If you're interested in an arbitrary moral philosophy, though, then it's as good as any other.
Seriously, man, it's time for you to drop Salter. The book is bullshit.
, then it's clear that you're not an objective reader.
Based on your comments, the only thing you've pointed out that is disputable is whether pursuing EGI can be adaptive in the sense meant by population geneticists.
I'm willing to concede that it may not be adaptive in that sense (perhaps I should not have said it isn't before, because it may be adaptive under certain conditions). But that's only because I think arguing about it distracts from the existence of EGI, which is not disputable.
Let's consider Salter's statement from a neo-Darwinian perspective. I don't know what you understand by neo-Darwinian, but I mean that something is good which increases the frequency of alleles that I share. If my ethny is wiped out, that is bad from a neo-Darwinian perspective. Maybe you consider that an event without neo-Darwinian repercussions.
Anonymous -
ReplyDeleteThe whole purpose of my argument here has been to show that Salter's book does not provide correct predictions about natural selection (probably not all of his predictions are wrong, but it's easy to find some that are), that the behaviors he describes (endorses?) are not adaptive in any sense meant by people who do actual evolutionary theory.
If you read my earlier comments, you'll see that, indeed, I grant the existence of EGI, as defined by Salter. Anyone can define a quantity, measure it, and write a book about how acting in favor of that quantity is "adaptive" (using a strange definition of the latter term). Again, my argument is that the book is mostly useless for someone who wants to understand how selection *actually* acts on behavior. If you agree with this, then we have no quarrel. n/a evidently does *not* agree with this, hence our long debate.
"neo-Darwinian, but I mean that something is good which increases the frequency of alleles that I share"
Notice your use of moralistic language here: something is "good" or "bad." Again, this is fine: you can believe whatever moral philosophy you want. But this isn't the definition of "adaptive" among evolutionary theorists, neo-Darwinian or otherwise. As such, it provides no value in predicting what will actually evolve, and therefore what humans are selected to do. This is what I'm interested in (if you're not, then fine - again, we would have no quarrel).
And yes, you can objectively arrive at the conclusion that something is bullshit.
RCB,
ReplyDelete"a novel allele that pays a reproductive cost"
Salter says it would be adaptive for the Englishman. Not for a "novel allele" in the Englishman. An Englishman's genome does not consist of a novel allele. It contains millions of variants of various ages, many of these shared with other Englishmen relative to Bantu.
And, yet again, the sorts of traits we're talking about will be massively polygenic, and any given allele will generally have little effect on the trait value, meaning novel alleles will not be instantly culled by selection and can easily drift to high frequencies in small groups. Ethnocentric groups outcompete non-ethnocentric groups. Bands grow into tribes. Tribes grow into nations.
When you imagine a novel allele and assert that if it is not favored within a population subdivision it must inevitably be selected against, you are denying the possibility of group selection whether you realize it or not.
"I recall that Queller's"
We're not talking about the broader paper. The example we're discussing specifically involves long-distance migration and competition between genetically distant groups. The quote means what it says, and is obviously correct.
"Send emails to Grafen, Queller, West, and Gardner."
ReplyDeleteGrafen, for one, coauthored a paper with Hammond and Axelrod:
To explore the evolution of tag-based altruism, our model embodies three mechanisms. To allow altruism, but not direct reciprocity, fitness is determined by neighbors interacting in a one-move prisoner’s dilemma. To allow behavior that is conditioned on indicators of relatedness, strategies can take account of observable tags, such as scent. Tags differ from signals (Spence 1974; Grafen 1990) by being inflexible expressions of an individual’s genotype rather than subject to individual control (Hochberg et al. 2003). To allow competition for scarce resources, the population is viscous, and the population size is fixed. Because the tags and strategies are not linked, the model allows for the possibility of ‘‘cheaters’’ who can be free riders in the group whose tag they carry. The resulting agent-based model is based on a model previously developed to study ethnocentrism in humans (Axelrod 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.
http://deepblue.lib.umich.edu/bitstream/handle/2027.42/72180/j.0014-3820.2004.tb00465.x.pdf?sequence=1&isAllowed=y
David Queller and others in 2006:
Which is the best: kin selection or group selection?
We view this as an empty question. There are three different ways of partitioning social selection: (i) the inclusive fitness extension of individual selection; (ii) the direct fitness model of individual selection; (iii) and the within-and-between group selection model [6,7]. Fletcher et al. spend most of their time advocating the second (a form of kin selection theory) but then conclude that group selection is best [1]. In reality, all three models are important and useful tools for investigating and modeling social evolution and, if applied carefully, will give the same answers [6–8].
https://lirias.kuleuven.be/bitstream/123456789/81568/1/foster_etal_tree_2006b_nothing+wrong+with+inclusive+fitness.pdf
Besides Henry Harpending, Alan Rogers and Vincent Sarich were involved in checking the math in Salter's book.
E.O. Wilson: "On Genetic Interests is a fresh and deep contribution to the sociobiology of humans."
Nor is Salter the first person to discuss "genetic interests". The phrase appears many times in the writing of Richard D. Alexander, for example.
On your political preconceptions: I never said you were a Marxist. I think your level of conditioning is probably if anything below that of the typical Westerner (and certainly typical academic) today. But it's still there.
Since you have failed to explain how an allele that encodes the aforementioned behavior (and therefore the behavior itself) could actually arise from low frequency (Challenge 1), I will assume that you cannot (as I said I would). If you are an honest scientist, this should worry you: you are unable to explain to a competent person why you believe what you believe.
ReplyDelete"Salter says it would be adaptive for the Englishman. Not for a "novel allele" in the Englishman."
Explain to me what "adaptive" means if it does not mean that a genetically-encoded behavior will spread to high frequency from low frequency. As I told Anonymous, I don't care about semantics: talk about what actually evolves. (This sounds, again, like you think loci that do not encode altruistic behavior have something to do with this. They don't.)
"the sorts of traits we're talking about will be massively polygenic"
Yes, the behaviors are polygenic. But all genes have to start at low frequency, and you have not explained how they get to high frequency in very large ethnic groups despite being selected against - except for "getting lucky," which is not adaptive. *Very* lucky, actually: more on this below.
"...can easily drift to high frequencies in small groups. Ethnocentric groups outcompete non-ethnocentric groups. Bands grow into tribes. Tribes grow into nations."
Yes, in small groups. But drift gets weak as the group grows. So unless you have constant turnover of groups recreating between-population variation (which is not the case for large historical ethnic groups), within-group selection kills off the ethnic altruist genes at the large level. See below. (Of course, ethnic altruist genes could be maladaptive relics of the past, when groups were small. But Salter says they are adaptive now. That's what I'm debunking.)
No, I'm not. But it's a tricky subject - I can see why you're confused. Let me explain it for you.
ReplyDeleteWhen altruistic alleles arise anew, they are present in only one person. Since they pay a cost, they are always disfavored in the first generation, and must survive some time to cross a threshold before they become selectively favored. With sib altruism, it's easy: if the allele can survive the first generation, then the rb>c is expected to hold in the next (yes, of course, segregation will mean it's not exactly 1/2 among sibs, but the point is that it becomes adaptive very quickly). For grandchild altruism, it's 2 generations, and so on. So, if the altruistic allele can stick around for just a bit (that is, if the cost is not too strong), it will pass the threshold, and be selected upward. (And that means the behavior evolves, which is what we care about.)
Now, what happens in large groups - say, 60 million, like 2015 England? Well, in the first generation, the allele is at frequency ~0, which means it has F_ST ~0 among groups. So neither between-group nor within-group selection are helping it at this point (actually, the latter is disfavoring it). Same is true in the second generation. And third. And fourth... In fact, the allele will continue to be disfavored (both within-group and globally) unless it can manage to drift to the frequency such that the FST Price equation holds for group selection to work (at which point it is globally favored). Note that it doesn't just need to *survive* for many generations, like the kin selection case, because "just surviving" means staying at the same frequency! It actually needs to *increase* in frequency relative to other alleles - despite being selectively disfavored! If you think this is a likely outcome, I have some swampland in Florida for you - a terrific investment opportunity.
For fun, let's imagine that miracles do happen, and the allele manages to drift to high frequency. Problem solved, right? Well, no. Selection will still be killing it off within the group (and any other group it happens to be in), so unless it can stay lucky forever (or if groups reshuffle a lot: see next paragraph), it's going to die off in every group, and therefore globally. The end result is that F_ST ->0 for these alleles, under these circumstances.
This is why groups cannot be allowed to be stable if group selection is going to work in the long term. If the groups are stable (i.e., they exist with high integrity for many generations), then the allele will go extinct in every single one of them. What needs to happen is that membership constantly reshuffles. When groups are quite small, this constant reshuffling will maintain high F_ST indefinitely, because some groups will happen to get many more copies of the allele than others. It *won't* work for large groups (e.g., 60 million - not that modern countries constantly reshuffle anyway), because the law of large numbers will ensure that basically all of them have the same frequency - so F-ST->0. So, for these kinds of alleles to be selected today, England would have to include a few dozen adults, and not exist for very long. They might even have to bump elbows with Bantus. Scary thought, I know.
So, I do get what I'm saying. And I take the time to explain it, too, instead of just bolding sentences in quotes. Give it a try, some time.
(That previous post should have been preceded by your quote: "When you imagine a novel allele and assert that if it is not favored within a population subdivision it must inevitably be selected against, you are denying the possibility of group selection whether you realize it or not.")
ReplyDelete"based on a model previously developed to study ethnocentrism in humans (Axelrod and Hammond 2003)"
Remember that I didn't ask you to gauge their opinions on group selection, but on Salter's work. (Remember that I also resimulated this model with dispersal and recombination, and it didn't work. But that's a separate mechanism entirely, and, to be fair, I didn't share the code with you.)
I don't think E.O. Wilson really gets mathematical evolutionary theory that well, to be honest.
The phrase "genetic interests" is a pretty simple one. I've used it before - that doesn't mean I agree with Salter, obviously.
Alan Rogers and Henry Harpending are smart. If they actually believe the Salter examples I've listed here, I'll be surprised (and will direct them to this debate). I won't just believe them, though: I don't defend my beliefs by quoting other people.
"I think your level of conditioning is probably if anything below that of the typical Westerner"
Not really sure what this means, but, again, I don't think you have any claim to know what my political beliefs are.
Will be offline for a few days. Hope to see something good when I get back. Hope you enjoyed the free pop gen lesson above.
RCB,
ReplyDeleteWhat are your thoughts on the arguments for the EGI concepts made here:
http://www.counter-currents.com/2011/04/ethnic-genetic-interests/
http://www.counter-currents.com/2011/04/the-ethics-of-racial-preservation-frank-salters-on-genetic-interests/
http://www.counter-currents.com/2011/04/why-was-the-understanding-of-ethnic-genetic-interests-delayed-for-30-years/
RCB,
ReplyDelete"you are unable to explain to a competent person why you believe what you believe."
I'm trying to be patient here, but I don't know how much better I can explain this. You either get it, or you don't. If you don't, you're not as competent as you believe; you either don't really understand the basic concepts we're talking about or have mental blocks when it comes to applying them to humans, and this is something you need to deal with yourself.
No matter how many times I explain this, you don't want to get it. Your starting point is that Salter can't be right; so when I explain to you how your reasoning is flawed, regardless of how many times we go through this or how many times I address a particular objection, your response is just to throw out additional confused reasoning, often forgetting things you previously agreed I was correct on and switching back to objections that have already been addressed.
The benefit of the ethnocentric altruism alleles comes from between-group selection, not within-group selection. Even in the first generation, an allele for ethnocentric altruism can potentially boost its odds of representation in future generations by, for example, reducing the chance of extinction of the group it's found in (whether by contributing to avoiding defeat and extermination by rival groups in an ancestral environment, or resisting replacement-level immigration in the modern world).
It makes no difference whether there is enough between-group relative to within-group competition at modern scales to maintain or grow this sort of variation in the long run (and certainly Hamilton speculated that self-sacrificing altruism would be found at higher levels in tribal people than in the long-civilized). In a particular instance of intergroup competition of the sort we're discussing, ethnocentric altruism alleles attuned to Hamilton's rule would by definition be adaptive.
This will always be true, regardless of scale, and regardless of how often the group is actually faced with the threat of intergroup competition. In the face of such a threat, the alleles (ones that enhance group competitiveness in a generalized fashion and are sensitive to cost, benefit, and relatedness) will be adaptive.
Your argument is somewhat analogous to claiming sickle-cell alleles can't be adaptive (or even exist in the first place!) in a malarial environment because their frequency would not increase in the long run in the absence of malaria. Even if malaria is nearly wiped out and the frequency of sickle-cell alleles begins to decline, this does not prove that if a mosquito with malaria does land on you you'd be better off not having a sickle-cell allele.
"Yes, the behaviors are polygenic. But all genes have to start at low frequency, and you have not explained how they get to high frequency"
ReplyDeleteAgain: positive selection will come from intergroup competition. If your argument is that all relevant variants would be snuffed out immediately, leaving no variation on which group-level selection could act, this is of course absurd. Yet again: we're talking about very large numbers of weakly-selected variants, and a large surface on which new mutations can arise.
Crow and Aoki modeled group selection for polygenic behavioral traits. Again, it boils down to Hamilton's rule.
Group selection for a polygenic behavioral trait: a differential proliferation model
http://www.pnas.org/content/79/8/2628.full.pdf
Group selection for a polygenic behavioral trait: estimating the degree of population subdivision
Our general approach is to use molecular markers, which are selected very weakly at most, as neutral indicators of population structure. GST gives us an appropriate description of the relevant aspect of the structure. By using Eq. 3 we can state the maximum value of cost/benefit of a quantitative trait if that trait is to increase in average value or frequency in the population. GST describes the present structure of the population; it does not tell us how it got that way. If this value has been roughly stable in the past, we could expect that traits with c/b up to this value would have increased in the population, assuming of course that such traits exist and have heritability greater than zero.
http://www.pnas.org/content/81/19/6073.full.pdf
Empirically, ethnocentrism exists, and no doubt has since before we were humans. Empirically, ethnocentrism has heritability greater than zero.
Nature, nurture, and ethnocentrism in the Minnesota Twin Study
http://www.researchgate.net/profile/Byron_Orey/publication/229073714_Nature_nurture_and_ethnocentrism_in_the_Minnesota_twin_study/links/0046352fddb446a6a9000000.pdf
Common Heritable Effects Underpin Concerns Over Norm Maintenance and In-Group Favoritism: Evidence From Genetic Analyses of Right-Wing Authoritarianism and Traditionalism
http://ioa126.medsch.wisc.edu/findings/pdfs/1287.pdf
Genetic evidence for multiple biological mechanisms underlying in-group favoritism
http://timbates.wdfiles.com/local--files/cv/Lewis%20Bates%202010%20-%20Genetic%20evidence%20for%20multiple%20biological%20mechanisms%20underlying%20ingroup%20favoritism.pdf
Your issue is with reality, not with Salter or me.
"Of course, ethnic altruist genes could be maladaptive relics of the past, when groups were small. But Salter says they are adaptive now."
ReplyDeleteAgain, see above.
And Salter never claims we are well-adapted to ethnic competition in the modern world. If he'd believed that to be the case, he would have had little reason to write the book. From the introduction:
On Genetic Interests is an attempt to answer the empirical question: How would an individual behave in order to be adaptive in the modern world? I adopt the neo-Darwinian meaning of adaptive, which is to maximize the survival chances of one’s genes. I begin by describing humans as an evolved species and thus as creatures for whom genetic continuity consists of personal reproduction or reproduction of kin. [. . .]
Humans can no longer rely on their instincts
There is nothing immutable or necessarily perfect about adaptations or the understanding, appetites and preferences they organize. Natural selection is constrained by evolutionary history and environment. It shapes bodies and behaviours in small increments by modifying existing species. Much in nature is badly designed, if one examines it from an engineer’s viewpoint. [. . .]
Like adaptations that advance them, proximate interests can be imperfect in promoting genetic interests. The main problem is the slowness of natural selection compared to the rapidity of technological and social change since the Neolithic. The inertia of adaptations can cause them to continue to promote proximate interests that no longer serve fitness. For most of humans’ evolutionary history, adaptations tracked slow-moving environmental change, including technological advances. In the species’ distant hominid and pre-hominid past, proximate interests that reduced an actor’s fitness were valued less and less as the genes that coded for such valuation failed to reproduce. For this reason, at most moments in time proximate interests have correlated with ultimate interests because the environment has changed so slowly that physiology and behaviour could keep track with it. Proximate and ultimate interests have been in equilibrium except where rapid changes in environment occurred. The equilibrium applying to humans has been upset in recent generations, so that we can no longer rely on subjectively designated proximate interests to serve our ultimate interest. We must rely more on science to perceive the causal links between the things we value and formulate synthetic goals based on that rational appraisal.
Proximate interests, often reflected in consciously held values, have become increasingly fallible guides to ultimate interests because modern humans live in a rapidly changing world. Humans evolved in small bands consisting of a few families, sometimes grouped into tribes numbering in the hundreds. For most of their evolutionary history humans made a living by hunting and gathering in largely natural environments. They lacked formal organization and hierarchy. Adults coordinated activities by negotiating simple demographic role specializations — by age and sex — on an egalitarian basis with familiar band members. Most information was common. Humans now live in societies numbering in the millions where the great majority of interactants are strangers or acquaintances. They make their living through a great diversity of occupations resulting in radical asymmetries in information. They live and work in largely man-made urban environments. They are formally organized into states administered by extended hierarchies of rank and resources actuated by authoritative commands, impersonal contracts enforced by the state authority, and powerful forms of indoctrination performed by universal education, centralized media and entertainment.
However, to the extent we are able to correctly reason about what would be adaptive in the context of intergroup competition, and act in accordance with this, we have the equivalent of our generalized ethnocentric altruism alleles.
New thread.
ReplyDeleteCertainly if an Englishman is at personal threat of being killed by invading foreigners, he should probably defend himself! But any costly effort to protect his ethnic group *as a whole* from invading foreigners is not going to be selected for....
ReplyDeleteHow many times do you have to tell this guy that "ethnic genetic interests" has nothing to do with selection?
When altruistic alleles arise anew, they are present in only one person.
EGI has nothing to do with altruistic alleles. The interests do not result from, nor are they preserved by, altruistic alleles. In fact, EGI has nothing to do with altruism at all.
The recognition of and attempt to protect EGI is a consciously selfish endeavor whereby the genetic structures existing within an ethny seek to preserve themselves by preserving the ethny.
Explain to me what "adaptive" means if it does not mean that a genetically-encoded behavior will spread to high frequency from low frequency.
ReplyDeleteTo be adaptive in this sense means "to further survival".