dubois@uwmacc.UUCP (Paul DuBois) (07/31/85)
I got this message from Paul Dolber in the mail, and he said I could post it if I wished. I think it is worthwhile, so here it is: --- Dear Paul, From: Wilson, Edward 0. 1978. On Human Nature. Harvard U Press, Cambridge, MA, pp. 32-33. "Adaptiveness means simply that if an individual displayed the traits [never mind which ones] he stood a greater chance of having his genes represented in the next generation than if he did not display the traits. The differential advantage among individuals in this strictest sense is called genetic fitness. There are three basic components of genetic fitness: increased personal survival, increased personal reproduction, and the enhanced survival and reproduction of close relatives who share the same genes by common descent. An improvement in any one of the factors or in any combination of them results in greater genetic fitness. The process, which Darwin called natural selection, describes a tight circle of causation. If the possession of certain genes predisposes individuals toward a particular trait... and the trait in turn conveys superior [genetic] fitness, the genes will gain an increased representation in the next generation. If natural selection is continued over many generations, the favored genes will spread throughout the population, and the trait will become characteristic of the species." Note that, except in one instance where I made the insertion, the expression is "genetic fitness," not fitness. I agree with you that Rich's comment, as nearly as I can recall it, was rather obvious: "Natural selection predicts that organisms best able to survive will survive; only survivors can procreate." There's no prediction there. Mike's comment, as nearly as I can recall it, in essence finished the thought: "Under any given set of conditions, the population as a whole will come more and more to resemble those individuals which reproduce the most." Not profound, but at least not silly. "Fitness" is somewhat confusing, particularly in combination with "survive;" one ought to speak instead of "genetic fitness," i.e., the ability to survive long enough to produce many offspring who will also have the ability to survive long enough to produce many offspring, and so on. In your hypothetical example, the slow-reproducing population may have been fitter in some scheme -- e.g., they may have better education, better jobs, more money, and an appreciation for Honegger's symphonies -- but the simple fact is that, if they reproduce less, their *genetic fitness* is lower and the future population will resemble them less and less. One may now ask whether natural selection can make more interesting predictions. Dawkins (in "The Selfish Gene") covers some of this ground vis-a-vis reproductive strategies in insect societies, but it was so complex that I can't shoot it back to you sensibly in less than a megaline. So, let's try the following example from Wilson's book, dealing with work done by Trivers and Willard: 1) In vertebrates (especially birds and mammals), large, healthy males mate at a relatively high frequency while many smaller, weaker males do not mate at all. 2) On the other hand, nearly all females mate successfully. 3) Females in the best physical condition produce the healthiest infants, and these offspring usually grow up to be the largest, most vigorous adults. Now, suppose we are considering the females. According to evolution, natural selection, genetic fitness, et al., what is the best reproductive strategy for females to get the greatest number of offspring who will successfully reproduce, thus maximizing the contribution of their genes to the population? Well, if the offspring of the female are decreasingly healthy as the female's physical condition deteriorates, then it would seem best to have the good-times offspring (which are healthiest) to be males more often than bad-times offspring (since these less healthy offspring would be equally likely as good-times offspring to mate successfully if female, less likely to mate successfully if male). "According to natural-selection theory, genes that induce this reproductive strategy will spread through the population at the expense of genes that promote alternative strategies. It works. In deer and human beings, two of the species investigated with reference to this particular question, environmental conditions adverse for pregnant females are associated with a disproportionate increase in the birth of daughters. Data from mink, pigs, sheep and seals also appear to be consistent with the Trivers-Willard prediction. The most likely direct mechanism is the selectively greater mortality of male fetuses under adversity, a phenomenon that has been documented in numerous species of mammals." I'm only just now reading this book, and I'm not entirely comfortable with it yet; such scenarios are sufficiently complicated that it becomes hard to separate the antecedents from the consequences, to tell whether the outcome is being predicted from the givens, or the givens from the outcome -- or whether this makes any difference. Perhaps this will pass with further reading/thought; in any case, I recommend the book to you. Dawkins's examples involving reproductive strategies in insects are even more complicated, but in both presentations it is noteworthy that the investigators whose work is being discussed first made predictions based on natural selection, *then* went and checked on whether or not the predictions were accurate. And again, note that it is "genetic fitness" which is under discussion: In the above, the progeny in good times were males more often than in bad times not because females are "fitter" than males, but because the genes producing this effect ensure their greater representation in the gene pool. If you find this particularly enlightening, despite being written on the fly, feel free to forward it to net.origins. Otherwise, don't bother, and I'll try to cover the Trivers-Willard business myself later in a more cogent form. While some material which is not in quotation marks is more-or-less quoted, I take full responsibility for any errors or misrepresentations in those parts not in quotation marks. Regards and best wishes, Paul. --- Concerning the last paragraph: even though I *have* posted this, I (and I suspect others) would be interested in more on Trivers-Willard. -- | Paul DuBois {allegra,ihnp4,seismo}!uwvax!uwmacc!dubois --+-- | "Why are you standing on one leg?" | "I'm trying to see if I'm a stork."