Table Of ContentGENE AVATARS
THE NEO-DARWININAN
THEORY OF EVOLUTION
Pierre-Henri Gouyon
Jean-Pierre Henry
Jacques Arnould
Translated by Tiiu Ojasoo
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PREFACE
France holds a rather unusual position in the field of evolutionary biology.
Whereas French naturalists from Buffon, Cuvier and Lamarck onwards made
great discoveries in centuries past, French biologists missed the turning when
it came to genetics. Until the 1970s, most French biologists were convinced
that genetics was not as interesting as developmental science (some "rare
species", for example R. Chandebois, still hold this view). For them, the
general principles of heredity resided in the cytoplasm rather than in the
genome. This was not, however, as great an obstacle as one might have
thought. For example, it did not prevent Malécot from being one of the most
outstanding theoreticians of population genetics, L’Héritier and Teissier from
inventing population cages for Drosophila, or Monod and Jacob from
discovering messenger RNA or, for that matter, from conceiving major works
such as “Chance and Necessity” or “Evolutionary tinkering”. In fact, it might
even have been an advantage because, in France, the centralized vision of
heredity rooted in Mendelian genetics was strongly tempered by the
importance traditional biology conferred to the cytoplasm. Not unsurprisingly,
therefore, it was in France that a mechanism whereby thecytoplasm regulates
genetic information was discovered. It was in France also that the genetics of
non-Mendelian elements like mitochondria and the ecological genetics of
nucleo-cytoplasmic interactions (e.g. male-sterility, see Chapter 4) were
developed.
Until the 1980s, evolutionary genetics was chiefly the preserve of Great
Britain and the USA. These countries published all the scientific journals in
the field(Evolution, American Naturalist, Heredity, Genetics...). Continental
Europe, however, built up enough strength to found a European Society of
v
vi Gene Avatars–The neo-Darwinian Theory of Evolution
Evolutionary Biology and edit a society journal (“Journal of Evolutionary
Biology”) which, with time, has acquired a good reputation and substantial
impact. The first editor, of American origin, worked in Basel (Switzerland);
the second was the first author of this book (the editorial office is now based
in the Netherlands). A major reason for creating the journal was to
counterbalance the dominance of Anglo-American culture in the field. Not
that our colleagues were unfriendly. On the contrary, they helped us a lot in
developing evolutionary genetics in France. We are deeply indebted to
scientists like Bob Allard, Janis Antonovics, Herbert Baker, Arthur Cain,
Deborah Charlesworth, Theodosius Dobzhansky, Chris Gliddon, John
Harper and Ledyard Stebbins - among many others - for their advice and
collaboration. The trouble was that not enough account was taken of the
“cultural” differences between “Anglo-Americans” and Europeans in a field
of study - evolution - with such close links to ethics and politics. (We shall
not discuss the ambiguity of the position of Great Britain which belongs to
both worlds).
Evolutionary biology tells us that isolation, by preventing a group of
individuals from being drowned in the average mass, can foster novelty.
Protection from mainstream genetic and evolutionary thinking has certainly
helped French biologists retain their strong interest in development. In
France, it was obvious - maybe too obvious - that a synthetic theory of
evolution had to take ontogeny and non-Mendelian genetics into account
long before Gould and Lewontin made such a claim. The original French
edition of this book was written in order to acquaint the French reader with
the theory of evolution. To the English-speaking reader its interest lies not in
any new theory but in the rather unfamiliar emphasis.
The authors’ interest in evolutionary genetics was awakened by Georges
Valdeyron's course on population genetics at the “Institut National
Agronomique” in Paris. This book is dedicated to him. The French edition
benefited from the ideas of Anne Atlan, Denis Couvet and Isabelle Olivieri
and was improved by suggestions from Beatrice Albert, Irène Till-Bottraud,
and many others. The authors are also grateful to John Maynard Smith and
Andy Dobson for their judicious comments.
Pierre-Henri Gouyon
Professor at Paris-Sud University
Lecturer at Ecole Polytechnique
Consulting Professor at INAPG
Director of UPRESA 8079
TABLE OF CONTENTS
INTRODUCTION 1
The Biologist's Dream
CHAPTER 1 5
Waiting for Darwin
From pre-Socratic myths to the transformism of Lamarck
1.Before evolutionism: Metamorphosis, fixism, spontaneous
generation 7
2.The concept of evolution. A French discovery 17
3.Lamarck, founding father of transformism 31
CHAPTER 2 45
The Darwinian paradigm
1.The Darwinian revolution 45
2.The origins of genetics: A story about peas 56
3.Biometricians versus geneticists 60
4.Eugenics and Lysenkoism 68
CHAPTER 3 81
Population genetics and ecological genetics
1.Population genetics 81
2.Heyday and decline of ecological genetics 98
3.Gene polymorphism 111
4.Neutral theory and the molecular clock of evolution 124
vii
viii Gene Avatars–The neo-Darwinian Theory of Evolution
CHAPTER 4 135
The key to genes
1.Male opposition 136
2.Individual, where are you? 141
3.What is life? Erwin Schrödinger's "essentialist" answer 143
4.An update on information 149
5.Genetic information 151
6.Genetic information and avatars 152
7.Individuals: Gene inventions for gene reproduction 155
8.Selection levels 156
9.Rules governing the evolution of information 160
CHAPTER 5 165
Conflict and cooperation
1.The genome, an array of compartments in constant conflict 165
2.Let's play! 175
3.The splendour and woes of sociobiology 180
CHAPTER 6 199
Eros and Thanatos revisited
1.Death at our heels? 200
2.What use is sex? 204
CHAPTER 7 221
What can be done, what can't be done
1.A visit to St. Mark's 221
2.Brachiopod architecture or "Seilacher's triangle" 223
3.Constraints and biological evolution 226
4.Was life wonderful in the Cambrian? 228
5.Struggle for life and a royal race 231
6.Finalism revisited 234
CONCLUSION 243
From an ideal to a tinkered world
MATHEMATICAL APPENDICES 245
MA1.Evolutionary pressures 245
MA2.Inbreeding coefficient 247
MA3.Lethal mutations 248
MA4.Genetic load 248
MA5.Fixation of neutral alleles 249
MA6.Game theory 250
Contents ix
GLOSSARY 255
BIBLIOGRAPHY 263
INDEX 269
GENE AVATARS
THE NEO-DARWINIAN
THEORY OF EVOLUTION
Introduction
THE BIOLOGIST'S DREAM
In the 4th century Saint Augustine wrote: "What then is time? If no one
asks me, I know; but if I am asked and wish to explain, I do not know." 1
Augustine's query about time seems to apply also to life. Can any biologist
claim to know the ultimate definition of life? If he did, he might be offered
an honorary chair on the National Ethics Committee but it is more likely that
he would be in real danger of taking himself for yet another Faust or
successor to Dr. Frankenstein. Would he not have acquired the knowledge
and thus the power held by the gods? Would he not have become God?
These epistemological and metaphysical issues have not stopped
biologists from pursuing their quest to understand living things, nor
prevented the deep changes that have occurred during the last two centuries
in a field of biology - reproduction and heredity - that has always fascinated
mankind. We are far from Montaigne's outcry (Essays, Book II):
What kind of monster is the drop of seed from which we are produced
and that encapsulates the impressions not only of bodily form but also of
the thoughts and inclinations of our fathers? Where does this drop of
water harbour these infinite forms? and how does it pass on those
similarities that progress in so bold and disordered a fashion that the
great-grandson looks like his great-grandfather, the nephew like his
uncle?
Jean Rostand (1894-1977), a French biologist and prolific author,
considered this passage by Montaigne exceptional on many counts. It asks
how not only physical characters are passed on but also mental traits. It
conveys the process' capricious nature and the onlooker's feeling of surprise
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