the news that teh archaeologists may have found the body of richard iii under a parking lot in central england brought to my mind another richard — richard ii…
…and that bill hamilton used him in an example of how altruistic behaviors might be asymmetrical depending upon the degrees of inbrededness and outbrededness of the individuals within a family.
it may be that the more outbred individuals in a family would be more altruistic towards their fellow family members who are more inbred since the inbred individuals would have more copies of whatever altruism genes we happen to be talking about. in other words, helping out one’s relative/s that have a greater number of altruism genes would increase one’s inclusive fitness — at least when it comes to altruism genes anyway.
here’s what hamilton had to say [pgs. 97-100 & 105-106]:
“…all that logic about how knowing that either he or that other are inbred should change, for example, how Henry IV felt about his cousin, Richard II, or how Henry’s son, soon to be Henry V, felt about his cousin (second cousin actually in this cast), Edmund Mortimer, and how Edmund Mortimer about him.
“Usually when explaining to non-biologists how kin are to be valued in the evolutionary calculations, I start with a deliberate vagueness about what might be meant by such a phrase as ‘the proportion of genes in common’ between two relatives and I concentrate on the obvious importance of the probabilities of gene survival. What ‘genes in common’ means it takes too long to explain and perhaps even at the end I don’t completely understand the matter myself … Suppose Harry Plantagenet, Prince of Wales, is inbred and hence has two copies, aa say, of some altruism-determining gene, while his second cousin Edmund Mortimer (who, as it happens, was another very plausible contender for the throne) happens to have one only — his genotype is ab let us say. What proportion at this locus do we say the two have in common? … As I have pointed out, diploids have two copies of every kind of chromosome, one from their mother and one from their father. And in fact this problem about ‘genes in common’ was an old chestnut for me and I had pricked fingers on it from the very earliest days of my ‘altruism’ obsession. I had dealt in outline with questions of relatedness when one or both of two relatives could be inbred in some of my recent papers (see Chapters 5 and 8 of Volume 1) but I was very much aware of various issues still unresolved or remaining unpublished.
“About the time of my first invitation from Rick Michod, Nathan Flesness, for example, had published in Nature a paper entitled ‘Kinship asymmetry in diploids’. He showed clearly that if one relative was inbred and another not or less so, as in my example of the first two pretenders to the English throne above, the coefficient of relatedness that the inbred should use when deciding how to behave towards the outbred relative (a cousin once again, let us say) was not the same as the coefficient that the outbred cousin should apply if making the reciprocal decision. The theory says that the outbred person in fact, to a slight degree, should tolerate the unfairness in the interest of his genes, or, to put it more properly, his genes should even see to such an unfairness coming about: he should behave more generously towards the inbred than he would expect the inbred to reciprocate….
“I thought initially that Henry V, as the more aristocratic, would be more inbred than Mortimer, but this turns out to be incorrect. At least as far as my Encylopedia Brittanica (EB) information goes (1967 edition, ed. W.E. Preece, article on ‘Plantagenets’), both had mothers unrelated to their husbands. Both were fighters in high health.
“For a more interesting comparison I consequently transferred attention to Henry V’s father, Henry IV, and his cousin, Richard II, both grandchildren of Edward III. Using these cousins to address the question hinted, we find that they were indeed both complexly but weakly inbred. Remarkably, none of their four parents was inbred, at least according to my source. Their two mothers, however, were linked (through their father) into the web of European aristocracy of their time. I counted for each seven independent paths by which a gene in a mother could be identical to one in her husbands — this within the limited one-page pedigree provided in EB. Amongst such paths, Richard II had the two shortest (via his ancestors Edward I and Philip III of France: six- and eight-ancestor chains, respectively; I will indicate such chains by 6 and 8. Henry IV had Henry III (father of Edward I) as his most recent ancestor, implying a 9-chain, and, more distantly, has to go back to Blanche, daughter of Alfonso VII of Castile, to find his next-best 13 chain. As to the longer chains, 5 for each, all were quite different and none shorter than 13. Altogether, with his coefficient at F=164/8192 against Henry IV’s F=19/8192, Richard II is the more inbred.
“Correspondingly to our theory Richard II usurps the throne; however, obviously such small Fs … are going to make, actually, a fairly neglibible difference to the regression coefficients. Consequently (in the spirit of my argument and showing the method) we need not be surprised that neither pretender to the throne deferred to the other and they fought out their claims.
“Turning to another ‘pretender’ in this picture, Edmund Mortimer, already mentioned, by my data he has F=0 [i.e. completely outbred – h.chick]. In this light it fits again that (a) he joined the rebellious faction of Hotspur and Glendower as its ‘pretender’ only by their persuasion; (b) after his capture, by Henry IV, he stayed tamely (and was tolerated) for the rest of his life as a semi-prisoner at Windsor; and (c) my EB source (Vol. 15, pp. 867-8) tells us that ‘Edmund seems to have rewarded Henry V with persistent loyality’, including informing him of a plot by others to depose the king and put him (Edmund) on the throne. All this has the expected slant of the theory.”
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what would be fun is if somebody went through the royal lines of england/the rest of europe and checked to see if there was a general pattern of more outbred individuals deferring to the more inbred ones. wouldn’t that be cool if it was the case! (^_^)
what i wonder is: could this asymmetrical altruism be misapplied? on a large scale?
we know that altruistic behaviors — being just a set of innate, instinctive behaviors — can be “misapplied” in the sense that they can be directed towards unrelated individuals. the poor bird parents who raise cuckoo chicks (chicks?!) are the prime examples; but even in humans, people adopt kids that are totally unrelated to themselves and raise them as their own (not that there’s anything wrong with that!) — or some ladies just direct all their altruistic behaviors towards cats or even “reborn dolls” (there is something wrong with that…).
could it be that outbred individuals might misapply their asymmetrical altruistic behaviors towards unrelated individuals rather than relatives? specifically, i’m wondering if they could pick up on certain behavioral signals given off by inbred peoples (maybe who show strong familial altruism behaviors?) and then defer to them like edmund (maybe) did to henry iv?
it’s a stretch, i know. just thinking aloud.
previously: hamilton’s unequal cousins or the asymmetry of altruism
(note: comments do not require an email. how richard iii was really killed.)
hbd chick 
I’ve been wondering if human nature doesn’t use some down and dirty proxy to estimated relatedness, such as in having grown up together. Or even being in the same fox hole. What evidence do we have for humans (as opposed to insects) being able to make fine distinctions as to F?
Also, if Arabs are so inbred why do they run away on the battlefield? You don’t see much altruism there? (This obviously contradicts the tenor of my previous question — shows what I don’t know.)
@luke – “What evidence do we have for humans (as opposed to insects) being able to make fine distinctions as to F?”
no, i’m not sure we can make very fine distinctions, either. but there is all that genetic similarity theory stuff that rushton has written about … and genetic sexual attraction … so people can pick up on something that indicates genetic similarity anyway (which might serve as a proxy for relatedness).
iow, i don’t think the whole process is as rough as “who was in your nest” (although that’s undoubtedly important, too) — but i doubt if somebody from an outbred society could pick out a second cousin they’d never met in a line-up. definitely not a third cousin, i’m sure. i’d put money on that.
@luke – “if Arabs are so inbred why do they run away on the battlefield”
do they? if they do, does this happen when they’re in battle alongside just their clan/tribe members, or are you talking about if/when they’re recruited into some national army or something?
@luke – “…as opposed to insects…”
btw, if you’re thinking about insects and relatedness, don’t try comparing ants/bees/other eusocial insects to humans, ’cause they have that weird haplodiploid sex-determination system which results in the sister workers being much more closely related to one another than they would’ve been to any theoretical offspring they would’ve had. crazy stuff!
“”I’ve been wondering if human nature doesn’t use some down and dirty proxy to estimated relatedness, such as in having grown up together. Or even being in the same fox hole. What evidence do we have for humans (as opposed to insects) being able to make fine distinctions as to F?”
Inclusive fitness makes sense so it *ought* to exist if there was a viable mechanism for making it so but i wonder if inclusive fitness programming itself is some kind of universal package or whether it’s a selected trait which might vary within a population like height (maybe not so much like height but vary anyway).
On top of the base programming as you say there’s the question of kin recognition. For it to work you need to be able to recognize kin. I see inbreeding as a way of enhancing familial resemblance (as most of the time people were living among people very similar to themselves) and it’s not exactly an exact science but it seems to me in the context in which it developed – cousin-marrying extended families in relatively low population densities -then i think you could have got a Wang nose or Wu ears and that would have worked more or less.
I do think inclusive fitness is likely to be fuzzy because of the difficulties of recognition but *on average* i can see it working at that scale.
I also wonder if scent is part of it as well and possibly in both directions e.g. you have the kibbutz dormitory children not being attracted to each other and the non-westermarck effect with split adult siblings. I wonder if there’s a relatedness scent that creates “love” behaviors and a suppressing scent that cancels the physical side of that. I also have a weird idea about barracks i.e. that armies that have infantry platoons sleeping in the same room might be more cohesive than armies that start to give them 2 or 4 man rooms. Just random thoughts but i think scent has something to do with it especially scent while asleep.
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“Also, if Arabs are so inbred why do they run away on the battlefield? You don’t see much altruism there?”
Altruism to who? One of the problems here – which i have all the time too – is altruism in the west has gradually come to mean selflessness whereas altruism genetically is completely tied up with relatedness. Running away from a *national* army back to the farm of your *extended family* is altruism if the balance of relatedness in that nation is heavily weighted to close kin i.e. through cousin marriage. If the balance of relatedness shifts to the national scale then the altruism shifts to the national scale also and you leave the farm to defend the nation.
“do they? if they do, does this happen when they’re in battle alongside just their clan/tribe members, or are you talking about if/when they’re recruited into some national army or something?”
Yes and no, precisely as you say. The same thing that (generally) creates bad national armies (unless unified by religion) makes very stubborn guerrilas.
@luke – “if Arabs are so inbred why do they run away on the battlefield”
They need to start partaking of alcohol again, tot of rum, dutch courage, the fuel of war.
I’m not into war novels, in fact I’ve only read about one, Fields of Fire, by Jim Webb (now Senator Webb of Virginia, who’s also written a mighty find book about the Scots-Irish — him are one in fact) and there are some unforgettable foxhole scenes in the book populated by a very diverse, as we say nowadays, set of characters. The bonds formed and the sacrifices made or risked were remarkable and I have little doubt they were based on experience. Somehow that has to be explained.
@luke – “Somehow that has to be explained.”
i agree!
@g.w. – “Yes and no, precisely as you say. The same thing that (generally) creates bad national armies (unless unified by religion) makes very stubborn guerrilas.”
yes, that’s what i had in the back of my mind, although i didn’t think it through all the way to the point about guerrilas. (^_^) lots of familial altruism (or whatever you want to call it) of the arab style produces good hit-and-run camel-raiding squads (or guerrilas). less familial altruism (from greater outbreeding) and you can more easily build a more cohesive, larger military force. or something like that.