Mathematical Structures In Population Genetics

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Mathematical methods have been applied successfully to population genet ics for a long time. Even the quite elementary ideas used initially proved amazingly effective. For example, the famous Hardy-Weinberg Law (1908) is basic to many calculations in population genetics. The mathematics in the classical works of Fisher, Haldane and Wright was also not very complicated but was of great help for the theoretical understanding of evolutionary pro cesses. More recently, the methods of mathematical genetics have become more sophisticated. In use are probability theory, stochastic processes, non linear differential and difference equations and nonassociative algebras. First contacts with topology have been established. Now in addition to the tra ditional movement of mathematics for genetics, inspiration is flowing in the opposite direction, yielding mathematics from genetics. The present mono grapll reflects to some degree both patterns but especially the latter one. A pioneer of this synthesis was S. N. Bernstein. He raised-and partially solved- -the problem of characterizing all stationary evolutionary operators, and this work was continued by the author in a series of papers (1971-1979). This problem has not been completely solved, but it appears that only cer tain operators devoid of any biological significance remain to be addressed. The results of these studies appear in chapters 4 and 5. The necessary alge braic preliminaries are described in chapter 3 after some elementary models in chapter 2.

This work reflects sixteen hours of lectures delivered by the author at the 2009 St Flour summer school in probability. It provides a rapid introduction to a range of mathematical models that have their origins in theoretical population genetics. The models fall into two classes: forwards in time models for the evolution of frequencies of different genetic types in a population; and backwards in time (coalescent) models that trace out the genealogical relationships between individuals in a sample from the population. Some, like the classical Wright-Fisher model, date right back to the origins of the subject. Others, like the multiple merger coalescents or the spatial Lambda-Fleming-Viot process are much more recent. All share a rich mathematical structure. Biological terms are explained, the models are carefully motivated and tools for their study are presented systematically.

Population genetics is the mathematical investigation of the changes in the genetic structure of populations brought about by selection, mutation, inbreeding, migration, and other phenomena, together with those random changes deriving from chance events. These changes are the basic components of evolutionary progress, and an understanding of their effect is therefore necessary for an informed discussion of the reasons for and nature of evolution. It would, however, be wrong to pretend that a mathematical theory, depending as it must on a large number of simplifying assump tions, should be accepted unreservedly and that its conclusions should be accepted uncritically. No-one would pretend that in the event of disagreement between observation and mathematical prediction, the discrepancy is due to anything other than the inadequacy of the mathematical treatment. The biological world is, of course, far too complex for the study of population genetics to be simply a branch of applied mathematics, so that while we are concerned here with the mathematical theory, I have tried to indicate which of our results should continue to apply in a context wider than that in which they are formally derived. The difficulties involved in the joint discussions of mathematical and genetical problems are obvious enough. I have tried to aim this book rather more at the mathematician than at the geneticist, and for this reason a brief glossary of common genetical terms is included.

The present monograph develops a versatile and profound mathematical perspective of the Wright--Fisher model of population genetics. This well-known and intensively studied model carries a rich and beautiful mathematical structure, which is uncovered here in a systematic manner. In addition to approaches by means of analysis, combinatorics and PDE, a geometric perspective is brought in through Amari's and Chentsov's information geometry. This concept allows us to calculate many quantities of interest systematically; likewise, the employed global perspective elucidates the stratification of the model in an unprecedented manner. Furthermore, the links to statistical mechanics and large deviation theory are explored and developed into powerful tools. Altogether, the manuscript provides a solid and broad working basis for graduate students and researchers interested in this field.

A basic method of analyzing particulate gene systems is the proba bilistic and statistical analyses. Mendel himself could not escape from an application of elementary probability analysis although he might have been unaware of this fact. Even Galtonian geneticists in the late 1800's and the early 1900's pursued problems of heredity by means of mathe matics and mathematical statistics. They failed to find the principles of heredity, but succeeded to establish an interdisciplinary area between mathematics and biology, which we call now Biometrics, Biometry, or Applied Statistics. A monumental work in the field of popUlation genetics was published by the late R. A. Fisher, who analyzed "the correlation among relatives" based on Mendelian gene theory (1918). This theoretical analysis over came "so-called blending inheritance" theory, and the orientation of Galtonian explanations for correlations among relatives for quantitative traits rapidly changed. We must not forget the experimental works of Johanson (1909) and Nilsson-Ehle (1909) which supported Mendelian gene theory. However, a large scale experiment for a test of segregation and linkage of Mendelian genes affecting quantitative traits was, prob ably for the first time, conducted by K. Mather and his associates and Panse in the 1940's.

A definitive account of the origins of modern mathematical population genetics, first published in 2000.