Zoology - Theses
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ItemNo Preview AvailableStudies in population geneticsThomson, J. A. (John Alexander), 1934- (University of Melbourne, 1959)Population genetics and. population dynamics have formed two distinct avenues of approach to a field of study which has become increasingly important over the last thirty years. Each has proved highly susceptible to mathematical analysis and to quantitative, rather than merely descriptive, treatment. It is, however, surprising to find that relatively little has been done to unite these two aspects of population biology, while in each theoretical work has tended to outstrip the small foundation provided by the observations so far recorded in the literature. Thus Birch (1957.P.217) was led to say... "I am quite sceptical of the predictive value of such mathematical models as have to date been proposed for natural populations". More serious still is the fact that the attempt to fit slender experimental data to such models has often blinded the experimenter to the possibility that factors of importance other than those considered in his model may contribute to the observed properties of the population. In particular, there has been little attention given to the genetic control of the form and rate of population growth; the work of Buzzati-Traverso (1955) was the first major experimental analysis to produce useful results along these lines. Much of the work on Drosophila populations has been done on small breeding groups which have been assumed to have reached an equilibrium density, although it seems likely that laboratory populations of Drosophila develop in the same way as those of Lucilia (Nicholson, 1954 and earlier papers), in which the population number fluctuates quite widely about a mean, not absolute, "equilibrium" level. Further, the number of individuals in the populations has not usually been determined with accuracy, so that it has seldom been possible to study small changes in population size. Preoccupation with "competition" experiments involving oligogenic markers has led even experienced workers to ignore the importance of modifying influences, particularly polygenic systems, associated with the genetic background. In this connection Buzzati-Traverso (loc.cit.) stated: "The fact is that the change in frequency of a single gene (or chromosome) during a number of generations means that the individuals carrying its allele will produce more or fewer adult offspring in the next generation than the individuals not carrying it. The factor which is decisive for changing frequency is a "productivity differential" involving of necessity the whole genotype and not one gene alone, for the latter will have different survival values in different genetic milies. Some extreme mutants, like those mostly used in laboratory experiments, may affect specifically the productivity of its carriers to such an extent as to make the effects of the rest of the genotype and of interactions of the mutant with it insignificant. But under natural conditions the commonest ease is very likely that of 'small* mutants where the natural selection mechanism probably involves many genes at one time" Even in the case of 'extreme' mutants erroneous or unsupported conclusions have been published (see later discussion of the relative adaptive values of the alleles at the white locus of Drosophila given by Merrell & Underhill, 1956); while in other instances over-emphasis of one particular factor has often prevented a balanced statement of the factors operating in laboratory population experiments. The discovery of selective mating in Drosophila, for example, resulted in the application of mating test results to population cage results without regard to other factors which might be causing the observed gene frequency changes. The work of Nicoletti & Solima (1956) and Morpurgo & Nicoletti (1956), amongst others, has passed largely unnoticed, although these authors adduced considerable evidence to support their view that selective mating is in fact not a controlling factor in laboratory cage competitions even where it can be demonstrated to occur in mating tests of the genotypes involved. In the same way, a sharp distinction has not always been made between the overall adaptive value of a genotype in a particular environment. The differential migration rates reported by liar land & Jackson (1958) under the title �Advantage of the white eye mutant of Drosophila melanogaster over the wild type in an artificial environment" provides an instance of such confusion. The only real measure of "advantage" or "superiority" is the relative contribution of a genotype to the gene pool of the succeeding generation, but these authors mention no breeding experiments at all. It is, however, clear that they have identified one of the factors which might be of importance in determining the frequency of white relative to that of its wild type allele in populations maintained in a particular environment. The main object of the present work has been to investigate further the relation of the genetic structure of a population to its rate and type of growth. Some attention has been given to problems of the sex-ratio and to change in gene frequency; in each case the underlying mechanisms have been studied. Emphasis throughout has been placed on the determination of the relative importance of the possible component factors under conditions of intense competition such as those found in population cage experiments.