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TOPIC: group fitness Evolution Beyond Selfishness: Inclusive Fitness
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group fitness Evolution Beyond Selfishness: Inclusive Fitness
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/ hpindda:ca.environment /
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(Alan McGowen) / 9:13 am Jun 10, 1991 / To: <
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Subject: Evolution beyond selfishness Message-Id: < 910609215654 70701.140 DHJ50-1 @CompuServe.COM The Law of Natural Selfishness Evolution by natural selection operates by changing gene frequencies. But exactly what does selection *select*? It does not generally select individual genes, but traits of whole organisms. Genes do not face the world as individuals, but only as partners in a complete genotype whose _expression_ is the set of traits of an organism, living and reproducing in an uncertain environment. Accordingly, the classical theory of evolution focused upon individual organisms as the units of selection. Fitness, according to this idea, although it may be reductively defined for individual *alleles* (gene variants), is biologically increased or decreased at the level of the whole *organism*. The fitness of an allele results from its effect, positive or negative, on the fitness of the entire organism. The fitness of the organism is its overall success in increasing its genetic representation in the future of the population. This is calculated as *individual fitness*, the relative reproductive success of a whole organism, rather than of a single gene variant. One can formulate a `law of natural selfishness' which states that organisms act so as to increase their own individual fitnesses. This `law' can be `derived' by arguing that it is the stable outcome of evolution: those individuals whose genotypes lead them to obey it are more favorably represented (almost by definition) in the next generation than those who do not. Selfishness therefore waxes in the world, and unselfishness wanes, quite inevitably as the outcome of selection. Much of organic life and behavior can be understood from the standpoint of this `law'. The philosophy of Social Darwinism accepts the validity of this `law' and extrapolates it to human societies, arguing that social policies which attempt to circumvent its operation (usually tacitly and illegitimately redefined as applying to subjective utility rather than to Darwinian fitness) are hopeless attempts to violate a fundamental truth of biology. There are however some very notable violations of this law. The most drastic example of all is the evolution of sexual reproduction, one of the greatest problems in all biology. Individual asexual organisms place 100% of their genotype into their descendents, but sexual organisms only 50%. Sex immediately reduces individual fitness by a factor of two. How then could it evolve? It does no good to point to all the good things which happen for a species as a whole once the scheme is adopted. From the standpoint of the selfish interests of the asexual organism, there is a reduction of profit in it: any step in that direction should be selected against. So how did it happen? The law of natural selfishness prohibits sex. But the world has been celebrating it for more than a billion years. Another, less all-encompassing but still extreme, example of violation, is provided by sterile worker castes in social insects. How can *sterile* individuals evolve? No matter how much good they might do for the hive as a whole, any organisms who took steps in the direction of sterility would be violating the `law of natural selfishness'. According to that law, sterile workers cannot exist. But exist they do. Altruism A trait
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group fitness Evolution Beyond Selfishness: Inclusive Fitness
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Alan McGowen: The most drastic example of all is the evolution of sexual reproduction, one of the greatest problems in all biology. Individual asexual organisms place 100% of their genotype into their descendents, but sexual organisms only 50%. Sex immediately reduces individual fitness by a factor of two. How then could it evolve? There are a number of mechanisms at the level of single-celled organisms which are analogous to sexual behavior in larger species. There are, in fact, asexual creatures which engage in sexual activity. I'm not sure where this idea came from that the evolution of sexual reproduction is one of the greatest problems in all biology , but perhaps it is not suprising that all the references given were social science references, not biology texts. And, to be blunt, I found the sociology in the post tedious and stupid.
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group fitness Evolution Beyond Selfishness: Inclusive Fitness
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Inclusive Fitness Social Conclusions: the Bankruptcy of Social Darwinism Consider the ubiquitous human incest taboo. This trait reduces individual fitness by reducing reproductive choice. But it raises the group fitness of the groups which practice it by avoiding the deleterious effects of inbreeding (such as reduction of disease resistance and of fertility). Here we see a ethical principle which may well be a heritable rather than purely acquired trait. Incest is damaging to the indidivdual too. This is because the offspring has a much greater chance of having a lethal gene expressed resulting in the death of the offspring. In the rest of the post you develop a moral theory which you try to _base_ on genetics. This gives new meaning to the phrase morally bankrupt . Your genetics isn't very good either. References: The Selfish Gene , The Extended Phenotype , Richard Dawkins, Oxford university press, 1989
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group fitness Evolution Beyond Selfishness: Inclusive Fitness
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1) The existence of bacterial sex in no way reduces the puzzle of how meiotic sex evolved. Sex (mixis) in bacteria is not immediately related to reproduction. In diploid organisms, as I pointed out in the post, there would appear to be a reduction in fitness by a factor of two over asexual reproduction. 2) A great deal of biological work has been done on the evolution of sex, and it is widely regarded as a major problem of biology, comparable in importance to the origin of coding. It is no accident that all multicellular organisms are meiotic: the genome size required to support multicellularity is not possible with less precise reproduction schemes than meiosis. But meiosis is an immensely complex process, whose evolution is certainly not well understood. 3) All the references were biology references. E.O. Wilson (Sociobiology) is a prominent biologist. I can't imagine how the other references appeared to be sociological. Perhaps the frequent use of words like sociality would give a very careless reader the idea that sociology was under discussion. But the content alone, with its focus on selection and fitness, would surely indicate to an even moderately well-informed person that biological sociality was the subject. 4) A person who makes statements that biology references, one of them an extremely famous work by an extremely famous scientist, are clearly not biology references is certainly not qualified to pronounce on what is or is not a difficult problem in biology. The identification of the post with sociology suggests that he is not qualified to discuss that area either.
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group fitness Evolution Beyond Selfishness: Inclusive Fitness
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Alan McGowen: 1) The existence of bacterial sex in no way reduces the puzzle of how meiotic sex evolved. Sex (mixis) in bacteria is not immediately related to reproduction. In diploid organisms, as I pointed out in the post, there would appear to be a reduction in fitness by a factor of two over asexual reproduction. No. The statement was Individual asexual organisms place 100% of their genotype into their descendents, but sexual organisms only 50%. Sex immediately reduces individual fitness by a factor of two. There was no justification for this statement. Further, a large number of organisms exist whose life-spans are not noticeably reduced by engaging in sex. Or was some different definition of individual fitness being used? In fact, as the original post points out (in mentioning incest taboos), there are definite costs to fitness involved in mono-genetics. The post seems to claim that somehow this does not impinge on individual fitness, even though there are costs, in terms of individual fitness, which are clearly visible. 2) A great deal of biological work has been done on the evolution of sex, and it is widely regarded as a major problem of biology, comparable in importance to the origin of coding. It is no accident that all multicellular organisms are meiotic: the genome size required to support multicellularity is not possible with less precise reproduction schemes than meiosis. But meiosis is an immensely complex process, whose evolution is certainly not well understood. Well, the original article was not about the distinction between genetic mechanisms, but about the existence of gene sharing. As Mr. McGowen has just pointed out, there is a bit more going on here than just gene sharing. 3) All the references were biology references. E.O. Wilson (Sociobiology) is a prominent biologist. I can't imagine how the other references appeared to be sociological. Perhaps the frequent use of words like sociality would give a very careless reader the idea that sociology was under discussion. But the content alone, with its focus on selection and fitness, would surely indicate to an even moderately well-informed person that biological sociality was the subject. Ok, I was out of line stating that all the references given were social science refernces. A closer look indicates that the references did have biological content. 4) A person who makes statements that biology references, one of them an extremely famous work by an extremely famous scientist, are clearly not biology references is certainly not qualified to pronounce on what is or is not a difficult problem in biology. The identification of the post with sociology suggests that he is not qualified to discuss that area either. This is also out of line. A biological theory of social evolution is definitely about sociology. Maybe a brief recap of the original article is in order here... (1) formulation of a ''law of natural selfishness'' which states that organisms act so as to increase their own individual fitnesses. This ''law'' can be ''derived'' by arguing that it is the stable outcome of evolution. problem: The article does not use this basis (organisms which make the best choices tend to live longer) in further discussion. Instead, the non-technical meaning of selfishness is used. (2) The law of natural selfishness prohibits sex. Not as formulated. The law of natural selfishness would say that in the long term, if individuals who engage in sex have some sort of advantage over those who do not, there will be more individuals who engage in sex. (3) Another, less all-encompassing but still extreme, example of violation, is provided by sterile worker castes in social insects. How can *sterile* individuals evolve? No matter how much good they might do for the hive as a whole, any organisms who took steps in the direction of sterility would be violating the `law of natural selfishness'. According to that law, sterile workers cannot exist. But exist they do. Again, this totally violates the formulated principle of the law of natural selfishness In fact, this is a rather clear example of selfishness. I suppose there might be some disagreement as to what constitutes an individual in the case of social insects, but since it is the queen which reproduces, the long-term success/failure of a colony depends on the treatment which the queen receives. The sterile individuals are engaging in selfishness by supporting the queen, and the queen is engaging in selfishness by reproducing more sterile individuals. (3) A trait
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