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! (^_^)

(note: comments do not require an email. citizens against altruism!)

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an over-simplification

the following is an oversimplification, but oversimplified models can be useful in thinking about complicated things, even if they don’t exactly represent what they’re supposed to be representing. i’ve been wanting to draw this out for myself for a while now, ’cause i’m a very visual thinker and i like pictures — even tho i’m not very good at drawing them myself. (~_^)

so … here is an (oversimplified) model of some altruism genes and inclusive fitness.

there are two altruism genes (alleles) here: a and A. (the capital letters here don’t signify dominance.) since humans (mostly) have two copies of every gene (allele), there are three possible combinations of these altruism genes that any individual might have: aa, aA, and AA.

ok. so far so good. now here comes the oversimplification…

in my little model, in the case of a famine, an individual with the aa combination does not share any of his food with his siblings. ever. so aa individuals always survive a famine, but their siblings do not. an individual with the AA combination always gives away all of his food to his siblings, keeping none for himself; so he always perishes in a famine, but his siblings always survive. aA individuals give away all of their food in half of the famine situations, so they perish half of the time while aiding their siblings’ survival half of the time.

obviously, genes don’t work this way. like i said … oversimplification.

anyway, so here’s what an average family in an average, randomly mating population would look like (edit: i guess i should’ve made two of them daughters. i’ll fix that later.):

what happens to the inclusive fitness — direct + indirect fitness — of each of these types of individual over the course of, say, ten famines?

first aa individuals. aa individuals never share any of their food with their siblings, so over the course of ten famines, the ten aa individuals will survive but none of their siblings will. so the direct fitness (not counting kids) of aa individuals = 10aa. indirect fitness (only counting siblings) = 0.

AA individuals always share all of their food with their siblings in a famine, so over the course of ten famines, the ten AA individuals will die, but all of their siblings will survive. so their direct fitness = 0. indirect fitness = 20aA (they each save one aa sibling and two aA siblings, but the aa siblings don’t count towards their inclusive fitness since they have no A alleles).

aA individuals share their food in 50% of the famines, so half of the time they survive, and half of the time their siblings survive. so their direct fitness over the course of ten famines = 5aA (themselves); indirect fitness = 5aa + 5aA + 5AA.

what does this all mean? well, if you add it up, it all looks like this:

– aa individuals ultimately save 20 of their own genes (10aa = 20a).
– AA individuals ultimately save 20 of their own genes plus 40 other genes (10aa + 20aA = 40a + 20A).
– aA individuals ultimately save 40 of their own genes (5aa + 10aA + 5AA = 20a + 20A).

so, maybe it pays (in a randomly mating population) to be somewhat altruistic, but not altruistic 100% all of the time? that makes sense. if you sacrifice yourself 100% of the time — well, does anybody do that? no. seems like that would be pretty quickly de-selected for (if that’s the right way of putting it). unless your familiy’s inbred, of course. that changes the situation. i’ll look at that in my next post.

also, take a look the AA individuals again. over the course of ten famines, they save twice as many alleles that they don’t even have as they do their own alleles. remember that. i’m gonna come back to that in another post.

that is all! (^_^)

previously: four things and technical stuff

(note: comments do not require an email. world’s most beautiful camel — descended from the camels of allah. she is kinda sweet! (^_^) )