human biodiversity and culture and history

william hamilton, considered to be one of the — if not the — greatest evolutionary theorists since darwin, had this to say:

“The incursions of barbaric pastoralists seem to do civilizations less harm in the long run than one might expect. Indeed, two dark ages and renaissances in Europe suggest a recurring pattern in which a renaissance follows an incursion by about 800 years. It may even be suggested that certain genes or traditions of pastoralists revitalize the conquered people with an ingredient of progress which tends to die out in a large panmictic population for the reasons already discussed. I have in mind altruism itself, or the part of the altruism which is perhaps better described as self-sacrificial daring. By the time of the renaissance it may be that the mixing of genes and cultures (or of cultures alone if these are the only vehicles, which I doubt) has continued long enough to bring the old mercantile thoughtfulness and the infused daring into conjunction in a few individuals who then find courage for all kinds of inventive innovation against the resistance of established thought and practice. Often, however, the cost in fitness of such altruism and sublimated pugnacity to the individuals concerned is by no means metaphorical, and the benefits to fitness, such as they are, go to a mass of individuals whose genetic correlation with the innovator must be slight indeed. Thus civilization probably slowly reduces its altruism of all kinds, including the kinds needed for cultural creativity (see also Eshel 1972).”

so hamilton clearly thought that biology and human biodiversity strongly influence culture and history, including the broad movements of history like renaissances or maybe even reformations or enlightenments, etc. and he thought that outbreeding, specifically too much outbreeding (i.e. panmictic populations), and presumably inbreeding too, relate to the selection for altruistic behaviors…and, therefore, certain aspects of cultures and history, etc. (remember that there’s more to hbd than just iq. (~_^) ) i dunno, maybe i and other hbd-ers are crazy (if so, we’re in GOOD company!), but this just makes intuitive sense to me. as john derbyshire said [15:00]:

“…if dimensions of the individual human personality are heritable, then society is just a vector sum of a lot of individual personalities.”

i like the big, probably impossible to answer fully questions: where does culture come from? where do institutions come from? where do renaissances come from? i don’t have the answers to those questions. nor am i under any illusions that i’ll ever be able to answer them. but am i very certain that they cannot be answered without taking into consideration human biology and biodiversity along with more conventional explanations drawn from history, economics, etc., and so i like to periodically bring them up.

so, if you happen to be new here, if you don’t like questioning — on every level — or biological explanations applied to The Big Questions, i’m afraid you’ve come to the wrong place. sorry. for my fellow hbd-ers — see you back here later in the week! (^_^)

previously: renaissances
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p.s. – btw, my vacation has been extended by a week (long story), so i’ll be back properly next week. sorry for the delay! (*^_^*)

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there and back again: shame and guilt in ancient greece

william hamilton wondered if renaissances/enlightenments happened in places roughly 800 years after some hardy altruism genes were introduced by barbarians into panmictic (really outbred) populations. i wonder instead if what happens is that renaissances/enlightenments occur after ca. 500 years or so of outbreeding which results in nepotistic altruism (or clannishness) being reduced or even mostly eliminated which, in turn, leads to greater cooperation and reciprocal altruism within the populations — conditions i think you might need to have a renaissance at all (see also here).

where intensive outbreeding (and manorialism) happened in medieval europe — and there is a lot of good, strong evidence for it — certainly seems to match well with where the european renaissance occurred. after some fits and starts in the 500s to 700s, the practice of avoiding close cousin marriages really took hold in exactly the areas where the renaissance/reformation/scientific revolution/enlightenment later happened — i.e. core europe — in short: england, france, the netherlands, germany, and northern italy. scandinavia a bit, too. oh…and the lowlands of scotland.

the evidence for outbreeding in ancient greece is much more tenuous. it appears fairly certain that the upper classes outbred during the archaic period in greece (800-480 b.c.). whether they outbred during the entire time period or began the practice sometime before or after 800 b.c., i don’t know. it may also be, judging by something hesiod said, that the lower classes followed suit, but it’s impossible to know for certain going by just one comment from one ancient writer.

some circumstantial evidence that might offer further support to the outbreeding-in-archaic-greece theory is that, in the 400s to 200s b.c., there was a shift in kinship terminology in ancient greece. the distinctions in the greek language between the paternal and maternal sides of the family began to disappear — for example, uncles on both sides came to be called just “uncle,” rather than there being specific words for paternal vs. maternal uncle, and so on and so forth. the same sort of linguistic shift happened in medieval europe. in germany, for instance, that shift happened between the 1100s and 1400s. at the end of the day, all cousins came to be called simply “cousin” rather than “father’s brother’s cousin” or “mother’s brother’s cousin.” the lesson seems to be: change the kinship structures and the long-term mating patterns in a society, and it shouldn’t be surprising that the kinship terminology will also change. no need to specify different sorts of cousins if all of them are off-limits as marriage partners.

michael mitterauer points out that there was a time lag in the linguistic shifts in medieval europe — the terminology changed ca. 300 to 600 years after the mating patterns began to change. perhaps something similar happened in archaic greece — the linguistic shift happened in ca. the 400s to 200s b.c. so perhaps we can infer that the mating patterns had changed to a more outbred form a few hundred years earlier. maybe right around the end of the greek dark ages and the beginning of the archaic period. dunno. complete speculation.

now i’ve come across another piece of circumstantial evidence that outbreeding may have been happening in archaic greece and that is that there was a(n incomplete) shift in the society during the time period from being a shame culture to being a guilt culture. i’m getting this from The Greeks and the Irrational, a book originally published in 1951 and written by classical scholar e.r. dodds (who was kicked out of oxford for supporting the easter rising — troublemaker! (~_^) ). presumably there have been works criticizing dodd’s thesis written since the 1950s, but i’m afraid i haven’t read any of them yet. i’m just going to run with dodd’s idea for now, but, please, consider this a sort-of thought experiment. more speculation.

first of all, in shame cultures, bad behavior is checked by the fear of being caught — of being shamed and embarassed. in guilt cultures, bad behavior is checked by one’s inner voice — feelings of guilt occurring before any action is taken. these are behavioral traits that must have been variously selected for in different human populations. secondly, shame cultures are all tied up with honor — especially family honor. japan — with its meiwaku and seppuku — is the classic example of a shame culture, but china with its confucian filial piety is not far behind. the arabized populations are definitely shame cultures with their honor killings and all their talk of respect. even european mediterranean societies are arguably more honor-shame cultures than guilt cultures [pdf].

if you’ve been reading this blog for any amount of time, you’ll recognize all of those shame cultures as having had long histories of inbreeding: maternal cousin marriage was traditionally very common in east asia (here’re japan and china); paternal cousin marriage is still going strong in the arabized world; and cousin marriage was prevelant in the mediterranean up until very recently (here’s italy, for example). it’s really, once again, the outbred northwest “core” europeans who are unique here with their guilt culture (although perhaps there are other guilt cultures out there as well). my guess is that long-term inbreeding tends to result in shame-honor cultures, while long-term outbreeding leads to guilt cultures. i’ve said so before.

back to dodd, his thesis is that ancient greece went through something of a transition from a shame to a guilt culture, but that shift was incomplete. the trend may even have reversed in classical athens. dodd points to several thematic shifts in greek literature from the iliad to the writings of plato including: a move away from blaming human failings on atē or the direct, external influences of the gods to more personal “demons,” often seen only by the individual person; the gradual adoption of the idea that individual humans have “souls” or independent “personalities”; a move away from the idea that people’s failings are due to a lack of knowledge (again coming from outside the person) as opposed to, perhaps, their own culpability; that zeus over time becomes more and more a dispenser of justice rather than just a being who capriciously interferes in human affairs (justice being important in guilt cultures as opposed to revenge in shame-honor cultures); and that philosophers and thinkers increasingly complained that the inheritance of guilt down through a family line was unjust. here from dodd on that last point [kindle locations 669-671]:

“Solon speaks of the hereditary victims of nemesis as άυαίτιοι, ‘not responsible’; Theognis complains of the unfairness of a system by which ‘the criminal gets away with it, while someone else takes the punishment later’; Aeschylus, if I understand him rightly, would mitigate the unfairness by recognising that an inherited curse may be broken.”

the idea that only the transgressor should be punished (as in guilt cultures) as opposed to additional or all of his family members (as in shame-honor cultures) doesn’t actually occur to these writers, so they haven’t quite arrived fully into a guilt culture, but they do seem to have been on the way there. much more so than earlier writers anyway. again, dodd emphasizes that [kindle locations 587-588]:

“[M]any modes of behaviour characteristic of shame-cultures persisted throughout the archaic and classical periods. There is a transition, but it is gradual and incomplete.”

the transition may have been incomplete — in fact, may have even gone into reverse — because inbreeding (cousin marriage) became increasingly common in classical athens (see here). from “Agnatio, Cognation, Consanguinitas: Kinship and Blood in Ancient Rome” in Blood and Kinship: Matter for Metaphor from Ancient Rome to the Present [pgs. 24-26], we saw in a previous post that while “aristocrats in early [archaic] Greece…married beyond the limits of their *patris*”, in classical athens “members of the *anchisteia*, the legally defined kinship group including first cousins once removed, were the preferred marriage partners.” the ancient greeks might’ve gone from being a (presumably) inbred/shame culture in the dark ages, to an outbred/quasi-guilt culture in the archaic period, and back to an inbred/shame culture over the course of the classical period. maybe. Further Research is Required^TM.

(yes, i know. it’s all very tenuous. i told you it was speculative!)

in any case, evolution is not progressive. (heh! i’ve just been dying to say that. (~_^) ) there’s nothing to say that evolution cannot go in reverse, although perhaps it wouldn’t go back down the exact same pathway it came up. there’s no reason why we — or, rather, our descendants — couldn’t wind up, as greg cochran says, back in the trees^*.

i think the way to think of the evolution of behavioral traits like nepotistic and reciprocal altruism in humans — especially perhaps in recent human evolution — is like a big simmering cauldron of stew where bubbles of certain behaviors rise up in some places only to sometimes pop and deflate and almost disppear again. outbreeding appears to have occurred many places, although whether or not over the long-term is not always clear: archaic greece (maybe), ancient rome, the bamileke of cameroon, the igbo of west africa, the turkana of east africa, the semai of malaysia, the bushmen of southern africa (aka The Harmless People), and europeans since the early medieval period — especially northwest europeans. the ancient greek experiment seems to have run out of momentum and collapsed on its own; the roman example probably popped thanks to the barbarian invasions; and the northwest european one is…currently ongoing. for now.

previously: renaissances and the transition from shame to guilt in anglo-saxon england (and “core” europe) and archaic greek mating patterns and kinship terms and ελλάδα
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_{*“Many were increasingly of the opinion that they’d all made a big mistake in coming down from the trees in the first place. And some said that even the trees had been a bad move, and that no one should ever have left the oceans.”}

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inclusive fitness

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

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

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

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

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

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

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

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

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

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

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

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reverse renaissance?

trigger warning: the following post contains much that is speculative. in fact, the entire post is one long speculation. if the thought of speculating when it comes to human biodiversity/sociobiology makes you queasy or fills you with existential angst, this might not be the blogpost for you. no, really. you might want to pass the time in some other way.
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i wrote about this once before, and since i’m extremely lazy, i’m just going to cut-and-paste from the previous post:

“in Innate Social Aptitudes of Man: An Approach from Evolutionary Genetics [pdf], william hamilton suggested that, perhaps, one gets a renaissance by (re-)introducing barbarian altruism genes into a too outbred population, letting the mixture ferment for ca. 800 years or so, and then enjoying the fruits of everyone’s labors. he’s talking here, of course, about the european renaissance of the fourteenth to seventeenth centuries…and classical greece/athens after the dorian invasion of ca. 800 years earlier? i *think*. if it happened at all (link inserted by me):

“‘The incursions of barbaric pastoralists seem to do civilizations less harm in the long run than one might expect. Indeed, two dark ages and renaissances in Europe suggest a recurring pattern in which a renaissance follows an incursion by about 800 years. It may even be suggested that certain genes or traditions of pastoralists revitalize the conquered people with an ingredient of progress which tends to die out in a large panmictic population for the reasons already discussed. I have in mind altruism itself, or the part of the altruism which is perhaps better described as self-sacrificial daring. By the time of the renaissance it may be that the mixing of genes and cultures (or of cultures alone if these are the only vehicles, which I doubt) has continued long enough to bring the old mercantile thoughtfulness and the infused daring into conjunction in a few individuals who then find courage for all kinds of inventive innovation against the resistance of established thought and practice. Often, however, the cost in fitness of such altruism and sublimated pugnacity to the individuals concerned is by no means metaphorical, and the benefits to fitness, such as they are, go to a mass of individuals whose genetic correlation with the innovator must be slight indeed. Thus civilization probably slowly reduces its altruism of all kinds, including the kinds needed for cultural creativity (see also Eshel 1972).'”

william hamilton — probably the greatest evolutionary theorist since darwin and an evil, evil speculator! not to mention crimethinker.

anyway…my own speculation re. the biological substrate of renaissances is that it’s not populations which experience an injection of barbarian altruism genes that wind up having a renaissance, but rather that populations which outbreed (i.e. quit marrying close relatives) for ca. 400 to 800 years (egs. medieval/renaissance europe and archaic/classical greece?) undergo a sort-of wikification of their society which drives intellectual openness and curiosity and sharing — the kinds of behavioral derring-do that you need in order to have a renaissance at all. see the previous post for more on all of those speculations.

today’s speculation is that perhaps the arabized world underwent a reverse renaissance process thanks to the introduction by the arabs of the most inbred form of cousin marriage — father’s brother’s daughter (fbd) marriage — to the populations of the middle east/maghreb (not to mention the introduction of arabs, themselves, who had probably been inbreeding closely for up to nine hundred years before their expansion).

the islamic golden age lasted for a good six hundred years or so, but instead of the scope of islamic philosophy and science and law widening over the time period — instead of a wikification process — the tendency was for thinking in the arabized world to narrow. ijtihad (“independent thinking”) was gradually replaced by taqlid (“imitation”). this narrowing of thought was already widespread in the muslim world by the twelfth century — just about 400-450 years after the arab conquests. (braudel puts the beginning and end dates of the islamic golden age as 813 and 1198, the beginning of al-ma’mun’s caliphate and the death averroes respectively. – pg. 202.)

irfan habib points out that the islamic golden age in science was very much founded on long-established traditions of free inquiry in the near east, from greece to persia [pg. 69 — link added by me]:

“[T]his particular phase in Islamic history was marked predominantly by the Mu’tazilite school of philosophy, which was based on freethinking and rationalism. It was an ecumenical setting for science, where savants of nearly all creeds and origins worked towards a common purpose. And this was not something new, it was a long established pre-Islamic tradition in the Near East, where translation of scientific and philosophical texts from Greek to Syriac took place….“

i wonder if what happened was that, with the establishment of the caliphate and all the civilized elements that went with it — good communications over long distances, (relative) peace within the realm, an excess of wealth — a “renaissance” was quickly established. however, that golden age — which happened in the early part of the era of the caliphs — was really a late flowering of whatever had been going on the region previous to the arabs (especially in persia). this renaissance was then reversed — stunted, really — as a result of the centuries of close inbreeding of the populations in the middle east and maghreb thanks to the introduction of fbd marriage by the arabs.

like i said — pure, unadultered speculation! (~_^)

previously: renaissances

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renaissances

in Innate Social Aptitudes of Man: An Approach from Evolutionary Genetics [pdf], william hamilton suggested that, perhaps, one gets a renaissance by (re-)introducing barbarian altruism genes into a too outbred population, letting the mixture ferment for ca. 800 years or so, and then enjoying the fruits of everyone’s labors. he’s talking here, of course, about the european renaissance of the fourteenth to seventeenth centuries … and classical greece/athens after the dorian invasion of ca. 800 years earlier? i think. if it happened at all (link inserted by me):

“The incursions of barbaric pastoralists seem to do civilizations less harm in the long run than one might expect. Indeed, two dark ages and renaissances in Europe suggest a recurring pattern in which a renaissance follows an incursion by about 800 years. It may even be suggested that certain genes or traditions of pastoralists revitalize the conquered people with an ingredient of progress which tends to die out in a large panmictic population for the reasons already discussed. I have in mind altruism itself, or the part of the altruism which is perhaps better described as self-sacrificial daring. By the time of the renaissance it may be that the mixing of genes and cultures (or of cultures alone if these are the only vehicles, which I doubt) has continued long enough to bring the old mercantile thoughtfulness and the infused daring into conjunction in a few individuals who then find courage for all kinds of inventive innovation against the resistance of established thought and practice. Often, however, the cost in fitness of such altruism and sublimated pugnacity to the individuals concerned is by no means metaphorical, and the benefits to fitness, such as they are, go to a mass of individuals whose genetic correlation with the innovator must be slight indeed. Thus civilization probably slowly reduces its altruism of all kinds, including the kinds needed for cultural creativity (see also Eshel 1972).”

“self-sacrificial daring” is probably the equivalent of greying wanderer’s “aggression”, chris’ “drive”, staffan’s “persistence under negative reinforcement”, and/or my “contrarianism” or independent-mindedness.

the connection between these two renaissances might, indeed, be the reintroduction of some good altruism genes, but i think that maybe what these two “rebirths” have in common — what led to them occur at all — are the ca. 400-800 years of outbreeding which happened right before they began. in medieval europe we have the catholic church banning close cousin marriage around the year 500, and many secular authorities banned close cousin marriage at various points after that. and in archaic greece — the period just before classical greece/athens — we apparently have at least ca. 400 years of outbreeding — amongst the upper-classes most probably — and possibly amongst the lower classes, too (hesiod in his Works and Days recommends that a man — an ordinary man, a farmer — marry a nice girl from the neighborhood — from the kome or village — so, if archaic greeks actually did this, their mating patterns would’ve been quite endogamic, but not necessarily to close cousins — maybe third or fourth cousins or something — see A Companion to Archaic Greece).

i think you need some loosening of the genetic ties in populations — enough to get rid of a lot or most of the “clannishness” — so that you can have a “wikification” of those societies, i.e. societies where individuals are really willing to openly share their ideas with other like-minded people (see, for example, harold’s comment on the scientific revolution in england). but outbreed too much, and you might lose that “self-sacrificial daring” — because as hamilton said:

“…the benefits to fitness, such as they are, go to a mass of individuals whose genetic correlation with the innovator must be slight indeed.”

share your innovative ideas — your scientific inventions — with the entire world, and you might wind up benefitting all of those people more than your own descendents (if you’ve got any).

already at the start of the classical period in greece/athens, the mating patterns began to narrow [pg. 67]…

“[W]ith the emergence of the *polis*, exogamy began to give way in some places to endogamy — to marriage within the community. For the upper classes, this meant marriage within a tight circle of aristocratic families living in the same *polis*.”

…so it’s maybe no surprise that the athenians battled throughout the classical period against various aspects of clannishness (cleisthenes’ reforms are one huge example of this struggle) and that their renaissance didn’t last more than a couple hundred years. europeans, on the other hand — especially northern europeans — have continued to outbreed for something like over ca. 1000-1400 years — which, perhaps, is leading to another sort of problem for society — that it’s simplying fraying away at the seams because the weave is not tight enough.

maybe. dunno. all wild speculation on my part, obviously.

previously: archaic greek mating patterns and kinship terms

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the asymmetry of altruism again

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.”
_____

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.)}

the evolution migraine

“Believing in the explanatory power of evolution by natural selection is like a migraine, or perhaps still more like being, as it was in the old days, a ‘wise woman’. The majority of humanity seem to have difficulty accepting that the ‘oddness’ of such a believer can be real — that is, simply an oddness and nothing else. As the migraine sufferer is suspected of malingering, and the woman who is merely literally wise, of witchcraft, so the evolutionist is always suspected of covert agendas unconnected with reality or the search for truth. In despair over the unending bemusement in friends and relatives and over the stream of articles and books that still pours forth stating Darwinism to be wrong, dead, right except for natural selection, superseded by this stale or ridiculous notion or that (all of which, evidently, the public eagerly buy and read, no matter what the competence of the writer or his knowledge of the evidence); puzzled, in short, by resistance to ideas that seem vastly more obvious and intuitive than, say, relativity or quantum mechanics, which every one accepts blithely with or without understanding, the evolution sufferer sometimes comes to believe it must be he who mistaken…. At other times the evolutionist may feel like one of the stranger ‘genetic morphs’ of his own theories — mutant carrier, say, of a fourth intellectual pigment of the retina capable of raising into clear sight patterns of nature and of the human future that are denied to the majority of his fellows, or perhaps just a person bewitched in babyhood to have revealed to him through blind sight, through such X-ray eyes, all the ravishing and foreboding beauty of the world that he now endures.”

– w.d. hamilton. “Narrow Roads of Gene Land, Vol 1.” pgs. viii-ix.

hamilton’s unequal cousins or the asymmetry of altruism

*post updated below.*

now here’s something i hadn’t thought of! [pgs. 97-100]:

“Idle in my bag stayed also the notes I had prepared for the defence of my formulas of relatedness against Rick [Michod] — that is, my struggles yet again through 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? Obviously the question hardly makes sense or at least doesn’t unless one says a lot more first. Had they been haploid like two moss plants who were cousins all would be easy; either plant X would have the altruism gene possessed by plant Y or it wouldn’t: one could give a probability and then simply average such chances for all the gene loci. But notwithstanding the thoughts given in the introduction to Chapter 2, most of the socially most interesting organisms are diploid. 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. The key to my understanding this had come when I realized, a year or two after my paper of 1964, that Sewall Wright’s correlation concept of relatedness hadn’t been quite what I needed for my own calculus of relatedness. For a Machiavellian world of pure gene-controlled behaviour I saw that what Henry IV and Henry V need to do to decide their cousinly status with regard to their respective copretenders to the throne is to predict the genotypes of Richard II and cousin Mortimer with reference to their own, and to do so by the best possible statistical reasoning. Just as a statistically minded agronomist uses a regression coeffficient based on his knowledge of responses to fertilizer to predict as best he can the yield of crop, so must the Henrys use regression coefficients of genes to be predict how genes in common may have descended to the cousin. This, of course, is not an adequate explanation but it gives the idea. The upshot was that I, too, needed the regression coefficient, not the correlation one of Wright.

“Wright had practically provided both of them but the one he himself had used had been the correlation. It is well known that if a bivariate scatter of values in not a straight line, the regression of Y on X is not the same as the regression of X on Y. The simplest ‘regression’ relatedness formula is b_XY = 2r_XY / (1 + F_X) where F_X is a measure of individual X’s inbreeding: this formula had been given in my papers of 1970 and 1971. The consequence for asymmetry of altruism was left implicit in the formula at that time — increase F_X and the value of the formula is obviously reduced.“

so, if your cousin is more inbred than you, you should (would) be more altrustic towards him than he to you because he has more of the genes which you share in common.

heh. i love it!

in a similar vein, fox et. al. found that, due to the differential inheritance of the x- and y-chromsomes, not all grandmas are equally related to their grandkids and that they are, therefore, more altruistic to some of the kiddies than others.

meanwhile, i’ve been calculating the different coefficients of relatedness for family members based on the differential inheritance of the sex chromosomes. how was i supposed to know all i needed was a nice regression coefficient formula? (~_^)

update 10/29: i forgot to give you the punchline to the henry vs. mortimer story! [pg. 106]:

“Edmund Mortimer, already mentioned, by my data he has F = 0 [in other words, he was not inbred – hbd 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 loyalty’, 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. The man to whom he gave loyalty, as noted, was seemingly no more inbred than he was; however, his pattern of behavior was probably set during the reign of his relative’s inbred and doubtless (to the youth) dominating and awesome father.”

update 09/12/12: see also the asymmetry of altruism again

_{(note: comments do not require an email. asymmetrical altruism.)}