Jody Hey                  Evolutionary Genetics

  Professor    -     Department of Genetics     -   Rutgers University

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A Reduction of "Species" Resolves the Species Problem ----- Jody Hey, January 1997


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THE ROLE OF SEX

In a gene tree view of the history of a sample of DNAs, sex is synonymous with recombination, and can be defined as any process that causes different portions along the sequence of a set of DNAs to have different gene tree histories. In the absence of sex, the gene tree history of a sample of DNAs is the same for all parts of the sequence. With sex, it is possible that the speed of genetic drift for one portion of a DNA is different from another portion. If there is a high recombination rate, then a large sample of DNAs will have a history of many different gene trees, perhaps as many as there are base pairs in the sequence.

For sexual organisms, a genetic species is the same as a Mendelian population, as defined by Dobzhansky:

A Mendelian population is a reproductive community of sexual and cross-fertilizing individuals which share in a common gene pool. . . .The smallest Mendelian populations are panmictic units (Wright, 1943), which are groups of individuals any two of which have equal probability of mating and producing offspring. (Dobzhansky, 1950)

Thus by definition, organisms within a Mendelian population share in a probabilistic process of reproduction, and all pairs of organisms are equally subject to reproductive failure and equally likely to reproduce. Within a Mendelian population, each generation occurs with some distribution of reproductive success among the component organisms. The shape of this distribution may vary across generations, but at any point in time the particular pattern of reproduction is a major determinant of the gene tree for all portions of the genome. A sample of DNAs for a short region of the genome will have a particular history, while a different genomic region will have a different history; yet all of these histories must run through the same historical procession of organisms, with a different group of reproductives each generation. Thus a Mendelian population carries genomes with numerous gene trees that were all shaped by a common birth and death process.

GENETIC DRIFT AND NATURAL SELECTION

From a genetic perspective, natural selection can be defined as variation in reproductive success caused by genotypic variation (Lewontin, 1970), and it is often cast as a directed force of evolutionary change in contrast to the random force of genetic drift. However at the level of DNA where there is linkage, natural selection on functional DNA sequence variation contributes to the genetic drift that occurs among linked sequences. In a genetic species of asexual organisms, a mutation that changes a DNA sequence and causes natural selection, also causes a new pattern of genetic drift among organisms that carry that mutation. In effect, a new genetic species is created by the mutation; although one of the species will probably be replaced by the other. For the DNAs of organisms with recombination, the acceleration of genetic drift by natural selection depends on the degree of linkage, the number of sites of functional variation, and the strength of natural selection on the functional variation (Hill and Robertson, 1966; Felsenstein, 1974).

Natural selection on functional genotypic variation may play a major role in the formation of new genetic species. However, shared genetic drift, and not natural selection, is the appropriate description of the essence of genetic species. Genetic species will share in the process of natural selection on functional DNA sequence variation, and thus will share adaptations. However this process proceeds both concomitantly with, and as a contributor to, genetic drift. Furthermore, from a genealogical perspective (Fig. 2), genetic drift proceeds even in the absence of DNA sequence variation and in the absence of natural selection caused by DNA sequence variation.



 

 



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© 1997 Jody Hey