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A Short History of Mathematical Population Dynamics

Nicolas Bacar

A Short History of Mathematical Population Dynamics

Nicolas Bacar IRD (Institut de Recherche pour le Dveloppement) Bondy France nicolas.bacaer@ird.fr

Whilst we have made considerable efforts to contact all holders of copyright material contained in this book, we have failed to locate some of them. Should holders wish to contact the Publisher, we will make every effort to come to some arrangement with them ISBN 978-0-85729-114-1 e-ISBN 978-0-85729-115-8 DOI 10.1007/978-0-85729-115-8 Springer London Dordrecht Heidelberg New York British Library Cataloguing in Publication Data A catalogue record for this book is available from the British Library
Mathematics Subject Classication (2010): 00A09, 01A05, 92D10, 92D25, 92D30, 92D40 Springer-Verlag London Limited 2011 Apart from any fair dealing for the purposes of research or private study, or criticism or review, as permitted under the Copyright, Designs and Patents Act 1988, this publication may only be reproduced, stored or transmitted, in any form or by any means, with the prior permission in writing of the publishers, or in the case of reprographic reproduction in accordance with the terms of licenses issued by the Copyright Licensing Agency. Enquiries concerning reproduction outside those terms should be sent to the publishers. The use of registered names, trademarks, etc., in this publication does not imply, even in the absence of a specic statement, that such names are exempt from the relevant laws and regulations and therefore free for general use. The publisher makes no representation, express or implied, with regard to the accuracy of the information contained in this book and cannot accept any legal responsibility or liability for any errors or omissions that may be made. Cover design: deblik Printed on acid-free paper Springer is part of Springer Science+Business Media (www.springer.com)

To Aili, Nina and Sophie

Preface

Population dynamics is the area of science which tries to explain in a simple mechanistic way the time variations of the size and composition of biological populations, such as those of humans, animals, plants or microorganisms. It is related to, but still quite distinct from, the more descriptive area of population statistics. One common point is that they make extensive use of mathematical language. Population dynamics is at the intersection of various elds: mathematics, social sciences (demography), biology (population genetics and ecology) and medicine (epidemiology). As a result it is not often presented as a whole, despite the similarities between the problems met in various applications. A notable exception in French is the book Mathematical Population Theories1 by Alain Hillion. But it presents the subject from the point of view of the mathematician, distinguishing various types of model: discrete-time models (t = 0, 1, 2...) and continuous-time models (t is a real number), deterministic models (future states are known exactly if the present state is known exactly) and stochastic models (where probabilities play a role). The book then considers logically discrete deterministic models, continuous deterministic models, discrete stochastic models and continuous stochastic models. In the present book I have tried to discuss the same subject but from a historical point of view. Research is explained in its context. Short biographies of scientists are included. This should make the book easier to read for those less familiar with mathematics and can usually help in understanding the origin of the problems under study. But this book is not just about history. It can also serve as an introduction to mathematical modelling. It seemed important to include the details of most computations so that the reader can really see the limitations of the models. Technical parts are emphasized in grey boxes and can be skipped at rst reading. The last chapter focuses on the numerous contemporary problems in population dynamics that one can try to analyze from a mathematical point of view. For those who would like to know more, the lists of references at the end of each chapter also include web sites from which original articles may be downloaded.
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Presses Universitaires de France, Paris, 1986. For English readers, Quantitative Methods in Biological and Medical Sciences by H.O. Lancaster (Springer-Verlag, 1994) gives a good historical overview without the mathematical details.

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Preface

It was not possible in a book of this length to give a complete picture of all the work developed until now or to talk about all the scientists who have contributed to the subject. The choice made necessarily contains an arbitrary component, particularly for the most recent decades. I hope nevertheless that the sample chosen is representative enough, and that people active in the eld whose work is not mentioned will not be upset. The ideal audience for this book would include: High school and university students wondering what links may exist between the mathematics courses they have to attend and the world around them, or students preparing personal work on a theme related to population dynamics. Mathematics teachers trying to make their course more attractive. The knowledge of the four elementary operations is enough to understand most of Chapters 1, 2 and 5. Chapter 3 can serve as an introduction to the applications of logarithms. This book also covers: recurrence equations in Chapters 1, 3, 8, 11, 14, 21, 23, 24; differential equations in Chapters 4, 6, 12, 13, 16; partial differential equations in Chapters 20, 25; an integral equation in Chapter 10; and applications of probability theory in Chapters 2, 7, 8, 9, 15, 16, 17, 18, 19, 22. People already familiar with demography, epidemiology, genetics or ecology and willing to compare their favourite area with others which may involve similar mathematical models. Readers interested in the history of science. This book is essentially a translation of the French edition published by Cassini Editeurs (Paris) in 2008 under the title Histoires de Math ematiques et de Populations. Some chapters have been reorganized or rewritten. Four gures have been added. A few misprints have been corrected. The lists of references at the end of each chapter have been extended and updated. These lists include web sites showing the original works. A reference followed by a URL means that it can be easily found at least in part by searching on the World Wide Web (e.g. through Google Books). A number of people have made remarks on various versions of the book, provided references and pictures or discussed copyright issues: Andr e and Catherine Bella che, Bernard Bru, Joe Gani, Geoffrey Grimmett, Franc ois Hamel, Nikolai Nikolski, Carel Pretorius and Niklaus Salzmann. Thanks to all of them. I also thank my mother for helping with the translation. Marrakech, August 2010 er Nicolas Baca

Contents

1 2 3 4 5 6 7 8 9

The Fibonacci sequence (1202) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Halleys life table (1693) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

1 5

Euler and the geometric growth of populations (17481761) . . . . . . . . . 11 Daniel Bernoulli, dAlembert and the inoculation of smallpox (1760) . 21 Malthus and the obstacles to geometric growth (1798) . . . . . . . . . . . . . . 31 Verhulst and the logistic equation (1838) . . . . . . . . . . . . . . . . . . . . . . . . . . 35 Bienaym e, Cournot and the extinction of family names (18451847) . . 41 Mendel and heredity (1865) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 Galton, Watson and the extinction problem (18731875) . . . . . . . . . . . . 49

10 Lotka and stable population theory (19071911) . . . . . . . . . . . . . . . . . . . 55 11 The HardyWeinberg law (1908) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59 12 Ross and malaria (1911) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65 13 Lotka, Volterra and the predatorprey system (19201926) . . . . . . . . . 71 14 Fisher and natural selection (1922) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77 15 Yule and evolution (1924) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81 16 McKendrick and Kermack on epidemic modelling (19261927) . . . . . . 89 17 Haldane and mutations (1927) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97

Contents

18 Erlang and Steffensen on the extinction problem (19291933) . . . . . . . 101 19 Wright and random genetic drift (1931) . . . . . . . . . . . . . . . . . . . . . . . . . . 105 20 The diffusion of genes (1937) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111 21 The Leslie matrix (1945) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117 22 Percolation and epidemics (1957) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121 23 Game theory and evolution (1973) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127 24 Chaotic populations (1974) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133 25 Chinas one-child policy (1980) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141 26 Some contemporary problems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149 Figures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 155 Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157

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