‘Fitness’, in evolutionary terms, simply measures how much of your genome you manage to pass on. At first blush, you’d think this would be 1/2 the number of offspring (because each offspring of a sexually-reproducing organism carries half that organism’s genes). But this misses a big part of the picture. Each of the organism’s siblings also carries half its genes (and so, in genetic terms, is equivalent to its offspring). When a sibling reproduces, its offspring carry 1/4 the organism’s genes; more distant relatives carry smaller parts. In a given generation, the organism’s inclusive fitness is half its offspring, plus one quarter its siblings’ offspring, plus one eighth its first cousins’ offspring, and so on. So, the organism can still show evolutionary fitness without itself reproducing.
Worker bees are a dramatic example. Darwin and others were at a loss to explain these bees’ apparent selfless devotion; they suggested some vague group-selectionist altruism at work (‘for the good of the hive’, so to speak.) As it turns out, worker bees are (genetically speaking) very selfish indeed. W.D. Hamilton showed why. Bees and their relatives are haplodiploid. Drones (males) develop from unfertilised eggs, while females (workers, mostly, plus the occasional queen) develop from fertilised eggs. A worker bee’s siblings share more (75%) of its genes than would its own offspring. If you consider only individual fitness, the workers’ ‘altruism’ makes no sense. But their inclusive fitness is greater if they work to maximise the queen’s reproductive success than it would be if they themselves reproduced.
Humans (and most other animals) aren’t haplodiploid. But inclusive fitness can still be at work. An organism (it can be human if you like, but that doesn’t particularly matter) that for whatever reason cannot or does not wish to reproduce can still ‘reproduce’ indirectly, through its siblings, cousins etc. If the organism in some way helps the siblings etc. have greater reproductive success than they would without the organism’s assistance, the organism increases its own inclusive fitness.
This is where same-sex pairings enter the picture. An organism with an exclusively same-sex orientation is unlikely to reproduce personally (it happens, of course, though it’s likely to be at a much lower rate than in its opposite-sex-oriented relatives). If there is a genetic basis for the orientation (this doesn’t mean that genes rigidly determine the orientation, merely that there is some genetic factor to it), then if you didn’t know about inclusive fitness, you’d expect that genetic factor to disappear pretty quickly. But if that genetic factor also helped its bearer to increase its inclusive fitness, it wouldn’t disappear; it would be passed on through the offspring of close relatives, and, since it added to overall fitness, would keep expressing itself in individuals with the same-sex orientation. (Note that any single same-sex oriented individual with this factor wouldn’t necessarily have actually to act in a way that increased its inclusive fitness; what matters is the overall tendency among all individuals with this factor.)
So far, all this is just what evolutionists, in tribute to Kipling, call a ‘just-so story’ – a plausible explanation, but one you could come up with sitting in an armchair; it might have no connection at all with reality. (And I should note that inclusive fitness is not the only possible explanation of how a genetic factor for homosexuality, if there is one, would survive despite the apparent paradox, i.e., how could a tendency not to reproduce sexually be passed on to new generations?). To firm it up, you’d need to predict (for example) that non-breeding female gulls who form a lesbian relationship tend to get more of their genes passed on than non-breeding females who don’t; and then you’d have to go out to the field, band a bunch of gulls, and start counting.
It’s a lot easier to use this hypothesis to explain same-sex behaviour in gulls (and even more so in insects – yes, there are insects with same-sex sexual behaviours) than in humans. The behaviour of most animals, especially the so-called ‘lower’ animals, tends to be stereotypical. That of humans is much more plastic (and almost certainly determined by genes to a far smaller extent than in other animals). But there are ways to test the hypothesis that increased inclusive fitness helps explain the success of a genetic factor for human homosexuality (again, if there is one). Among some Native American groups, certain males would assume a female-like role, even to the point of marrying another male. Far from being despised, these individuals enjoyed high status, and raised the status of their immediate families. In most societies, high status has correlated with high reproductive success. Gathering and processing the data would be difficult (if possible at all), but if it could be shown that families that produced these individuals consistently tended to enjoy greater overall reproductive success than families that did not (after adjusting for any other factors that contribute to reproductive success), that would lend credence to the inclusive-fitness hypothesis.
You’ll probably have spotted one difficulty right away. These Native Americans had a same-sex orientation, but would they have defined themselves the same way that gay people do in modern western society? Who knows, but there’s a good chance they wouldn’t have. Perhaps they saw themselves as female, only with male genitals; perhaps they saw themselves as a third sex altogether. (However they understood their difference to other males, the difference was important; they were viewed by the whole tribe as endowed with sacred powers). Some women from the Nuer people in the Nile basin would marry another woman; the wife would bear her female husband children with the assistance of a helpful male (who had no role in the family thereafter). These women seem to have viewed themselves, and to have been viewed by others, as simply men with a woman’s body. (And yes, they did add to their extended family’s status – the children were legally their offspring, not the male proxy’s, and the birth of a son brought a dowry to the female husband’s family.) Ancient Greek males who enthusiastically formed same-sex pairs would also enthusiastically marry and have kids; the two categories seem to have been completely separate for them, and I doubt they’d have identified as gay in our modern sense. Same-sex oriented people have understood themselves in many different ways at different times and in different societies. Even if there is some genetic component to same-sex orientation, I don’t think that has anything to tell us about the lives, experiences or self-understanding of people who have that orientation. I really mean it when I say that the question (i.e., is there a genetic component or not) is of great interest in an evolutionary context, but of none whatever in any other context.
As interesting as this question may be for an evolutionist, your instinct is correct. The far bigger question is why there is any sex in the first place. The earliest organisms were asexual (and many still are). Sex had to evolve, and it’s an apparent paradox that it could do so. After all, it immediately halves the number of genes an organism passes on, and that number decreases by half in each generation thereafter. An asexual organism, by contrast, can go to its grave content that its descendants unto the nth generation will still bear (except for any mutations) 100% of the genes it carries itself. How could sex possibly compete with the stellar fitness rates of asexuality? Forget any hypothetical gene for homosexuality; a gene for sex at all should have disappeared in the blink of an evolutionary eye; yet it didn’t. Hamilton, probably the most important evolutionary theorist of the 20th century, spent most of the latter part of his career puzzling over this. His theory – called the ‘parasite red queen’, after the Red Queen in Alice who noted that one had to spend all one’s time running just to stay in the same place – was that sex represented a major new weapon in the eternal arms race between organisms and the organisms that parasitise them (‘parasites’ being understood in a broad sense, to include microbes etc., not merely ticks and the like). Despite the apparent vast drop-off in fitness caused by sex, Hamilton hypothesised that sexual organisms are, over the long term, far better able to adapt to parasite attack than are asexuals. It doesn’t matter that only half the sexual organism’s genes go forward in each offspring; half of a hundred is fitter than 100% of three. And there have been field studies suggesting Hamilton was right. This is immensely important for evolutionary theory; but really, it’s not the first thing on my mind when I am, emm, doing my bit against the parasites.
Well, I’d intended for this to be a brief response, and now look at what you’ve set off. Sorry; but evolutionary theory has an endless fascination for me, and my buttons are easily pushed. Do have a look at the Olivia Judson book if you have a chance. It tells some truly amazing stories and is immensely entertaining to boot. If you read it, you’ll see that we organisms – humans as well as our cousins the elephants, fish, spiders, tapeworms etc. – are stranger than anything a science fiction writer could dream up.
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