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WM:- I have to start out by thanking Guy for a nice explanation of why meiosis negates the gene as a unit of selection >snip< JE:- Once real epistasis (non additive genomic gene associations) and not just phoney epistasis (additive genomic gene associations) which appear to typify every single genomic gene _fitness_, enters selective reasoning, then genes cannot be independently selected no matter if Hamilton's rule suggests that they can. If a gun fails to shoot then it is always more parsimonious to suggest the gun is faulty rather than the laws of nature have suddenly changed... > GH:- > OK. Here is my quick answer to why it does not make sense. I think > the most concise way to explain it has been frequently argued by John > Edser. [Note that I have always agreed with John on this aspect of his > arguments, while disagreeing with other aspects.] Individuals are not > even close to merely the additive effects of genes. JE:- All real gene fitness is epistatic. Thus real gene fitness is entirely dissolved at meiosis, leaving only the organism level of fitness for selection to act on. Thus all independent gene fitnesses that are supposed to be independently selectable at Hamilton's proposed gene level of selection are fiction, leaving only one level of selection within _testable_ evolutionary theory. A simplified model is simply, not a theory and is misused if it is allowed to compete and win against any theory it was simplified from (as if such a thing was not 100% obvious). GH:- > ..it is still rather inconceivable to me how natural selection > can operate when the reproducing agents exist at a scale of > organization different from that where fitness is manifested (others > might prefer to word "evaluated" here), I hope that you (or anyone) > can explain it to me. I am not aware of any logical explanation being > provided by Dawkins, and IMHO there is no such logic. JE:- All you seem to be asking is how can a multilevel have different levels such that one level "acts" while another produces all the "fitness". This is possible when a lower level is only selectable at a higher level, as is the case with gene fitnesses. Dawkins just has everything in REVERSE. Since no independent gene level exists then genes are actors for individual fitnesses that incorporate 100% of gene fitness epistasis. Amazing as it may seem, Darwin's suggestion that Darwinian fitness is heritable is more accurate than all the algebra used by population geneticists re: fitness at the gene level because Darwin's remark includes all epistatic gene fitness. The challenge is to reformulate genetics and population genetics so that non additive gene relationships become "heritable" and not just "inherited". Fisher's model is so entrenched nobody is even willing to think outside his (tiny) square. Regards, John Edser Independent Researcher PO Box 266 Church Pt NSW 2105 Australia [EMAIL PROTECTED]
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