ANIMAL BEHAVIOR R. H. Wiley 28. Sexual selection Why are females choosy and males competitive (in most species)? -- females (by definition) produce nutritive gametes (eggs), which are metabolically expensive to replace -- their reproductive success is usually limited by how many eggs they can produce (or young they can nourish) -- male reproductive success is usually limited by how many mates he can obtain -- production of gametes or nourishment of young does not usually limit male reproduction -in some species, a male's contribution to parental care (building nests, feeding young, incubating eggs) does limit his reproductive success -- in these species males are choosy about mates (seahorses that brood their mates' eggs in a pouch, sticklebacks that build and tend a nest for eggs) Natural selection results from differences in reproduction or survival of individuals (phenotypes) with different genes in a particular environment sexual selection is a special case of natural selection -- it results from differences in reproduction as a result of differences in access to mates -- it differs from other forms of natural selection in a crucial way -- it is self-accelerating -- once started it tends to snowball Consider two loci (for simplicity) -- one affects female preferences for mates (P results in a preference for males with some trait, like a long tail, and p results in no preference) -- one affects the male trait (T results in a long tail, t does not) -- females with allele P tend to mate with males with allele T, females with allele p mate at random, males with allele t have less chance for mating than those with allele T -- in the next generation more individuals will have allele T -- note what happens to allele P -- because females with P tend to mate with males with T, their progeny (both male and female) tend to have both alleles (T alleles might not be expressed in females, nor P alleles in males, but both male and female progeny carry them) -- tendency of two separate alleles to occur in same individuals is called genetic correlation As generations pass, alleles T and P continue to occur together in the same individuals -consequently when females expressing allele P choose a mate with allele T, they usually are also choosing a mate carrying allele P -- the more P alleles there are in the population, the greater the mating advantage of males with T alleles, the faster allele T spreads, and the faster allele P spreads (because of genetic correlation) -- so the P allele spreads itself -- in other words, spread of P alleles is self-accelerating (produces positive feedback) ANIMAL BEHAVIOR R. H. Wiley Sexual selection (continued) Sexual selection can produce two paradoxical results (1) Female preferences result in evolution of male traits that reduce their survival -higher mortality of males with preferred traits is balanced by higher success in mating -experiments have compared male barn swallows with shortened or elongated or unmanipulated tail feathers -- birds with experimentally shortened tails do better than unmanipulated males -- they are more likely to survive the winter and they bring larger (more nutritious) insects to feed their young -- those with experimentally lengthened tails do worse (2) Self-accelerating feature of sexual selection allows purely arbitrary female preferences to spread -- arbitrary preference is one for a male trait that has no advantages for male or female other than access to mates On the other hand, sexual selection is eventually stopped either (1) by sufficiently high mortality of males with the preferred trait or (2) by any net costs to the female of choosing Female preferences cannot evolve (spread in a population) if females incur costs as a result of expressing a preference -- for instance, preferences can take time and involve risks of predation and consequently might reduce chances for reproduction or survival Consequently, female preferences for advantageous traits (rather than arbitrary traits) are more likely to spread -- two kinds of advantages for female preferences -- direct advantages (because preferred males provide more parental care or better territories or because choosing them involves less risk of predation or disease for the female) -indirect advantages or good genes (because preferred males have genes for greater resistance to disease which offspring inherit) Male barn swallows with longer tails are better parents (provide larger insects to feed young), have greater genetic resistence to insect parasites of nestlings, and are more likely to survive the following winter -- evidence for both direct and indirect advantages of mate choice