Bisexual Superiority and the Genetics of Homosexuality
In attempting to argue for the evolutionary value of homosexuality, scientists are confronted with an apparent paradox: if homosexuality is a valuable trait, it should have a genetic basis—yet how can a gene that doesn’t lead directly to reproduction continue to be passed on from one generation to the next? Perhaps, some have suggested, because the putative gene for homosexuality does not operate on its own, but rather is acting in tandem with another gene to promote reproduction. An often-cited analogy involves the genetics of sickle-cell anemia and malaria resistance in humans. People who receive a sickle-cell gene from one parent and a regular hemoglobin gene from the other parent are resistant to malaria; those who receive two sickle-cell genes (one from each parent) succumb to sickle-cell anemia, while those who receive two regular hemoglobin genes are more likely to succumb to malaria. Thus, genes that (on their own) can potentially decrease an individual’s reproductive capacity continue to be passed on because they are beneficial when combined with each other. Scientists have suggested that this might also be the case with homosexuality, as follows: Suppose there were one gene that predisposed an individual to homosexuality, and another that predisposed an individual to heterosexuality. Those individuals who receive two homosexual genes (one from each parent) would be exclusively homosexual; others would receive two heterosexual genes and be exclusively heterosexual; while those receiving one of each would be bisexual. If individuals who have one homosexual and one heterosexual gene were somehow more successful at reproducing, then the gene for homosexuality would confer an advantage and would continue to be passed on, even though it would sometimes result in individuals who do not reproduce (those who receive a homosexual gene from each parent). 20
At first glance, this hypothesis seems counterintuitive: regardless of the genetic mechanism involved, why should bisexual individuals be superior at procreating or have a reproductive advantage? On the contrary, one would expect individuals with two heterosexual genes—those who are exclusively or “doubly” heterosexual, as it were—to be more successful breeders than bisexuals. Nevertheless, this hypothesis accords surprisingly well with a number of aspects of animal homosexuality that remain puzzling under other accounts. First of all, as noted previously, bisexuality is widespread in the animal kingdom. Unlike other theories about the evolutionary value of homosexuality, this hypothesis recognizes that many individuals who participate in homosexual activity may also be involved in heterosexual behavior, and therefore capable of reproducing and passing on their genes. Additionally, the incidence of bisexuality within populations is often high: in a number of animals such as Bonobos, Japanese Macaques, Bottlenose Dolphins, Mountain Sheep, Giraffe, and Kob, for instance, virtually all members of the species (or of one sex) participate in both same-sex and opposite-sex interactions (either concurrently or at different points in their life). Again, this hypothesis predicts that such situations should exist, since it argues for the maximization of bisexuality in a population—that is, if bisexual individuals are more successful breeders, they should tend to make up the majority of a population.
Even more startling, in a few species bisexual animals actually do appear to be more successful than exclusively heterosexual individuals at reproduction, heterosexual mating, and/or attracting members of the opposite sex. As we have already discussed, pairs of male Black Swans, who can father cygnets by associating temporarily with a female and then raise the resulting offspring on their own, are generally more successful parents than heterosexual pairs. In part, this is because such same-sex pairs are more aggressive than male-female pairs and are therefore able to acquire larger and better-quality territories, which are essential for successfully raising cygnets. They may also have an advantage because both males contribute to incubating the eggs, whereas in heterosexual pairs males may take part in less of the incubation duties. Over a three-year period, 80 percent of male pairs in one study were found to be successful parents, while only about 30 percent of heterosexual pairs successfully raised offspring (unsuccessful parents either deserted their clutches, lost them to predators or other hazards, or ended up having their cygnets die). Homosexual pairs constituted up to a quarter of all successful parents even though they made up only 13 percent of all breeding pairs or associations in the study population. 21
Animals who participate in homosexual activity are also sometimes more successful at attracting members of the opposite sex, or participate more often in heterosexual mating. For example, male Ruffs who display with and mount male partners on their courtship territories attract females for mating more often than males who display by themselves. Because of their superior strength and courage, as well as their high rank in the flock, Greylag Geese in gander pairs or other homosexual associations are also sometimes attractive to the opposite sex. Females may associate themselves with a gander pair and eventually form a bisexual trio, mating with one or both of the males and raising their goslings together. In Pukeko, breeding groups in which homosexual interactions take place between males are also the groups in which the most intense heterosexual copulatory activity occurs. Adolescent Guianan Cock-of-the-Rock males who participate in the most visits to adult males’ display territories, during which homosexual courtship and mounting often occur, sometimes acquire their own territories at a younger age. With earlier access to heterosexual mating opportunities, this may give them a “head start” on breeding. Likewise, female Oystercatchers in bisexual (as well as heterosexual) trios may have an advantage in acquiring their own breeding territories and heterosexual mates in subsequent years. 22
A number of studies have also shown that animals that are the most active heterosexually are sometimes also the most active homosexually. In specific populations of Sociable Weavers, Bonnet Macaques, and Asiatic Elephants, for example, the top two males in terms of heterosexual mountings and other behaviors also participated in the most homosexual activities. Some of the most complete male homosexual behavior in Japanese Macaques, including full copulations with ejaculation, was exhibited by “one of the most vigorously heterosexual males in the troop,” while in another study the one female in a troop who failed to form any homosexual consortships also did not participate in any heterosexual consortships. 23And as mentioned in the preceding chapter, in a number of birds such as Common Murres, Laysan Albatrosses, and Swallows, most individuals who participate in homosexual copulations are in fact breeders who have heterosexual mates, rather than nonbreeders who are heterosexually inactive.
In spite of these rather unexpected confirmations, however, the bulk of the evidence does not actually favor this hypothesis and in fact disconfirms many of its predictions. Most of the examples cited above that seem to support the idea of “bisexual superiority” are misleading because they are based on anecdotal, rather than quantitative, information, and because they only look at a few individuals at a single point in time (or, at most, over the span of a few breeding seasons). To assess whether bisexual animals are more successful at reproducing, what is actually needed is a long-term study of large numbers of individuals that tracks them over their entire lifetimes, comparing the total number of offspring produced by bisexual animals to the total number produced by heterosexual individuals. Needless to say, this would be a huge and difficult undertaking, complicated by the logistics of keeping track of hundreds or even thousands of animals over many years and potentially large geographic areas, tabulating not only the reproductive output of each individual but also his or her entire sexual history to determine which animals are bisexual and which are exclusively heterosexual. Not surprisingly, few longitudinal studies of this type have been conducted, and those that have rarely involve species in which homosexual or bisexual activity is prominent (or else they do not take into account such behavior when it is present).
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