in Understanding Human History, michael hart did a real nice job of explaining how kin selection or inclusive fitness works and how “genes for altruism” could be selected for [pgs. 37-38]:
“For about a century after Darwin proposed his theory of evolution, the origin of altruistic behavior in animals remained a puzzle. It was not until the 1960s, when William D. Hamilton proposed his theory of kin selection, that a satisfactory explanation was given. That theory can perhaps best be explained by an example:
“Suppose a man sees his identical twin drowning in a river, and estimates (correctly) that if he were to jump in and try to save his brother the probability of success would be 80%, while the probability that he would die in the attempt would be 20%. Consider these two alternatives:
“a) Some of the man’s genes strongly dispose him to rescue his brother, and he therefore jumps in and tries to save him (‘altruistic behavior’).
“b) The man does not have genes that dispose him to rescue his brother, and he therefore stays on the shore and lets his brother drown (‘selfish behavior’).
“In case (b), exactly one copy of the man’s genes survives, and may later be replicated. However, in case (a), if the rescue attempt is successful, two copies of the man’s genes survive (one in his own body, one in his brother’s). As this will happen 80% of the time, on average 1.6 (= 0.80 × 2) copies of the man’s genes will survive. In this situation, therefore, genes that dispose a person to altruistic behavior will — on average — have more surviving copies than genes that dispose a person to act selfishly and will be favored by natural selection.
“Now consider a slightly different example. Suppose that the man on shore is a brother — but not a twin — of the person who is drowning. Case (b) will still result in one copy of his genes being preserved. However, since ordinary siblings share only 50% of their genes, if the man on shore succeeds in rescuing his brother then (on average) 1.5 copies of the man’s genes will survive. Since 80% of the attempts will be successful, case (a) will on average result in 1.2 (= 0.80 × 1.5) copies of the altruistic genes surviving. Since 1.2 is greater than 1.0, the altruistic genes will be favored by natural selection in this case too.
“Suppose, however, that the two men were not brothers, but merely first cousins. First cousins, on average, share only one-eighth of their genes. In this case, altruistic behavior results in only 0.9 (= 0.80 × 1.125) copies of the man’s genes surviving, and natural selection will therefore favor the genes for selfish behavior.
“The upshot is that a gene that disposes its bearer to behave altruistically toward a close relative can have a selective advantage over one that disposes its bearer to act completely selfishly. Furthermore, this can occur even though the relative never returns the favor, and even if the survival of the relative does not increase the group’s chances of survival. It is not necessary that either reciprocal altruism or group selection operate for kin selection to result in the spread of genes that dispose their bearer to act altruistically toward close relatives.”
what’s missing from these examples is, of course, inbreeding. and depth of time.
take michael’s second example up there…
“Suppose that the man on shore is a brother — but not a twin — of the person who is drowning.”
…but let’s add that the parents of these brothers were first-cousins. that makes these two guys: brothers AND second-cousins (i.e. the children of two first-cousins). so they probably share not only 50% of their genes in common as brothers, but also 3.13% of their genes in common as second-cousins. so the “push” to jump in the water to save the brother/cousin must be somewhat stronger in the inbred pair than for the brother to save just a plain ol’ brother.
now let’s take this example of michael’s…
“Suppose, however, that the two men were not brothers, but merely first cousins. First cousins, on average, share only one-eighth of their genes.”
…but let’s make them double first-cousins rather than just first-cousins. what happens then?
well, while first-cousins probably share 1/8th or 12.5% of their genes in common, double first-cousins share … well, double that! … or 1/4 or 25% of their genes in common.
what happens to michael’s calculation then?
“In this case, altruistic behavior results in only 0.9 (= 0.80 × 1.125) copies of the man’s genes surviving, and natural selection will therefore favor the genes for selfish behavior.”
in the case of double first-cousins the calculation becomes 0.80 x 1.25 = 1.0. that’s just breaking even using michael’s example, but what if the odds of saving the cousin from drowing are better than 80%?
or what about the depth of time i mentioned above? what if the family of my double first-cousins has been inbreeding for a very long time. a very, very long time. like for fifty generations or more. then the relatedness between all the family members, including these double first-cousins, will be even closer. natural selection ought, then, to favor such double first-cousins jumping in to save each other.
as wade and breden showed (see also previous post), inbreeding can help to accelerate the rate of the evolution (or frequency in a population) of altruism genes [pg. 846]:
[T]he increase in matings between homozygous parents decreases the genetic variance within families, because these matings produce genotypically homogeneous arrays of offspring.”
repeated inbreeding in a family reduces the diversity (whoa!) of the allele types within that family, and if we’re talking about “genes for altruism” here, then the variety of those must get reduced within inbred families, too. in a population that consists of, say, ten inbreeding families, the one that has super-duper altruism genes that lead all of its family members to help each other out more than the members of the other families will have the advantage (provided selection favors that advantage for whatever reasons). and those super-duper altruism genes will no doubt eventually spread to the other families since, in reality, no family groups inbreed 100% of the time anywhere — there will pretty definitely be gene flow between families. so then you’ll get a whole population of super-duper family altruists (note that these people are NOT altruistic to unrelated individuals).
the human populations on earth today that inbreed most closely (within patrilineages) and often practice double first-cousin marriage — AND have been doing this for prolly at least a couple of thousand years (time depth) — are the arabs (who later spread these mating practices to the maghreb, the mashriq and far off places like iraq and afghanistan and all the other ‘stans) and some peoples in the levant like the druze. i think that, because of their long-standing mating practices, they are the prime human examples of wade and breden’s accelerated evolution of altruism thanks to inbreeding.
previously: inbreeding and the evolution of altruistic behavior and more on inbreeding and the evolution of altruistic behavior
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