Milestones in the Evolving Theory of Evolution

344 Pages·2020·6.091 MB·English

by  David Wool

Milestones in the Evolving

Theory of Evolution

David Wool
Department of Zoology

Tel Aviv University

Israel

Language Editor
Naomi Paz

Department of Zoology

Tel Aviv University

Israel

Illustrations by
Dr. Leonid Friedman
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Cover credit: Lost forever: “Lonely George”, the sole survivor of the Pinta tortoises, Galapagos, died in 2012.
Photographed at the Darwin Station, Santa Cruz, Galapagos:
David Wool 1987. Inset in the upper right corner: the same animal, photographed in 2010 by
R. Lamb & Patricia MacKay, 2010
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Library of Congress Cataloging-in-Publication Data
Names: Wool, David (Professor of zoology), author.
Title: Milestones in the evolving theory of evolution / David Wool,
Department of Zoology, Tel Aviv University, Israel.
Description: Boca Raton, FL : CRC Press, [2020] | Includes bibliographical
references and index.
Identifiers: LCCN 2020019400 | ISBN 9780367903336 (hardcover)
Subjects: LCSH: Evolution (Biology)--History.
Classification: LCC QH361 .W66 2020 | DDC 576.8--dc23
LC record available at https://lccn.loc.gov/2020019400
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Dedication
In fond memory of my beloved grandparents,
Michael and Golda Einbinder,
and my parents,
Avraham (Avrum) and Esther (Tunia)

Preface
The present book emerged from my series of lectures on the history of the Theory of Evolution,
delivered freely since my retirement in 2002, once a week for one or two semesters every year, at
the School of Zoology, Tel Aviv University. In the audience were scientists, university students, and
members of the general public.
I am a biologist, not an historian. I have taught courses related to evolution for almost 30 years.
My book, “The Driving Forces in Evolution”, was published in Hebrew in 1985 – and an extended
version appeared in English in 2006. In the present book I have attempted to describe, at least
briefy, the ideas that have contributed to the theory that changed the course of thinking about the
biological world and impacted so many issues in human society, as these ideas have been expressed
during the last 200 years.
My motivation to switch from active biological research to history – besides the need to give
up my laboratory upon retirement – derived from a gift given to me of an 1874 copy of Darwin’s
book “The Descent of Man”, by my late friend and colleague, Prof. Boaz Moav, following the
publication of the Hebrew version of my book in 1985. The following year, when on sabbatical
leave at Michigan State University, in East Lansing, I saw a large display of Darwin’s books at the
entrance to the library – and realized how little I knew about the contributions of Darwin to science
(in particular, I was ignorant of Darwin’s botanical books). In a used-book bookshop across the
street from the university I purchased the two volumes of Darwin’s book on domestication, as well
as his book on the expression of emotions in animals and man, which fascinated me and stimulated
my desire to collect more of his works – forming the nucleus of my personal library, which now
includes most of Darwin’s books as well as those of his supporters and other contemporaries.
The selection of events, and of scientists (below I shall use the term “savant”, often used by
Darwin and his contemporaries to refer to educated and thinking men) whose ideas are part of
the history of the Theory of Evolution – and which are described in this book – are undoubtedly
affected by my own impressions, pre-conceived ideas, and limits of knowledge. While preparing the
material for my lectures, as well as for the present book, I devoted myself to reading and referring
to the original works of each of the savants that I considered important. Until most of the material
had been written, I avoided secondary materials apart from the biographies of these savants, which
I gathered from sources such as the Encyclopedia Britannica and from databanks publicly available
on the internet.
When refecting on the contribution of these savants to the evolution of the Theory of Evolution,
I wanted to illustrate how different their ideas were from the contemporary, accepted interpretation
of the world at the time they had proposed them. I did not attempt to judge how closely their ideas
agreed with the Theory of Evolution as accepted today. I did not intend to offer a progressive picture
of the development of the theory evolution; but, rather, to evaluate the ideas according to the erratic
chronological order in which they had been published.
History, like evolution, is a continuous process, and one that can be surveyed only in retrospect.
The Theory of Evolution, however, seems to have been developed by saltations (to use Darwin’s
expression), marked by new points of view expressed by a savant – not entirely in agreement with
the currently accepted opinion – or even directly opposed to it.
Chronologically, it is possible to mark the dawn of the 19th century as a reference point, when the
great majority of persons who gave the matter any thought believed that the world had been created
vi Milestones in the Evolving Theory of Evolution
as we see it now, and as it has remained ever since1. Before that time, the biological world had
been regarded as a chain or ladder, leading from the highest point – Man, the height of perfection
created “in His image”, down to the lower, degenerate, other animals – or, conversely, from simple
organisms up to the higher and more complex, as argued by J.B. Lamarck in 18092. The next
landmark is 1859, the year that Darwin’s book “The Origin of Species” was published and posited
that the world had evolved gradually, with some forms becoming extinct and being replaced by
others by natural selection. The chronological path from 1800 to 1859 – and then onwards to the
“modern synthesis” and molecular evolution towards the end of the 20th century, was not paralleled
by a linear progression of the theory. Lamarck’s suggestion in his “Philosophie Zoologique”3 that
animal species were not fxed but have changed (“transmutation of species”) and evolved gradually
from simple to more complex organisms, was an idea that was later incorporated into Darwin’s
theory – but, at the time, Lamarck’s eccentric ideas were ignored or ridiculed even by his colleagues
at the French Academy of Sciences. In the 50 years that followed, biological thinking revolved
around geology and fossils. The fossils, interpreted as the remains of extinct animals, were accepted
as indications of previous, extinct forms of life – which had disappeared due to catastrophic changes
on the face of the earth4. At the same time, the British geologist Lyell insisted that – given enough
time – the normal, observable forces of erosion and volcanic activity were suffcient in themselves
to account for the changes observed in the fauna between successive geological strata, and it was
unnecessary to assume catastrophes to explain these changes.
Darwin adopted the ideas of transmutation and common descent5 and offered a new mechanism
which could explain how the transmutation was effected – Natural Selection. It took years before
most biologists converted to the new theory, and some prominent ones even reverted to Creationism6.
A signifcant advance occurred in 1900, when a paper on crosses of plants, written by Gregor
Mendel in 1865, was rediscovered and began the era of genetics7. The modern Theory of Evolution
came to life when genetics became amalgamated with the Darwinian theory.
In the absence of a linear trend, I decided that the best way to arrange the book would be in a
series of four circles. The centers of the circles move by saltations along the temporal axis from
1800 to the end of the 20th century. Chapter 1 deals with theories of the origin of life on earth:
clearly, no biological evolution was possible before there was life on the earth. The four circles
commence in Chapter 2. The center of each circle is dedicated to a dominant fgure. Around the
centers, the work and ideas of distinguished savants are arranged irrespective of their more or less
“advanced” views of evolution. Diverging ideas, sometimes contradicting each other, foat within
the circle – occasionally a “satellite” idea leaps away from the perimeter.
As the center of the frst circle I have selected William Paley, a British clergyman who published,
in 1802, a book called “Natural Theology”8. In this book – which Darwin claimed he knew
almost by heart – Paley presented his outlook of the world as created by a Supreme Power – an
interpretation continuous with that which had prevailed before him. These ideas still remain today
as the fundamentals of Creationism, which rejects evolution. This outlook can be regarded as a “null
hypothesis” in the context of the present work.
The center of the second circle moves from England to France. After the French Revolution
in 1789, the French Academy of Sciences and the Museum of Natural History in Paris became
the center of all sciences in early 19th-century Europe. Three great scientists made their mark on
1 “Milton’s hypothesis”: Huxley 1876, Science and the Hebrew Tradition. Lectures on Evolution I.
2 See Chapter 6.
3 See Chapter 6.
4 As advocated by Cuvier, Chapter 4.
5 See Chapters 8-9.
6 Chapter 10 on Gosse and Agassiz.
7 Chapters 20-25.
8 See Chapter 2.
Preface vii
the biological sciences: Georges Cuvier9, Geoffroy Saint-Hilaire10, and in particular, Jean Baptiste
Lamarck11. In the same circle I have included the British (actually Scottish) geologist Charles Lyell12.
The third circle, centered on Charles Darwin, encompasses a wide perimeter – dealing with the
birth and publication of the “The Origin of Species”, the central issues of Darwin’s theory of natural
selection13 and his later books “The Descent of Man” and “Sexual Selection”14. It includes the
contributions of Darwin’s allies and supporters, Alfred Russel Wallace15, Thomas Henry Huxley16,
and Ernst Haeckel17. The struggle of the new theory to survive against the opinions of the Church
and other critics (Samuel Wiberforce, Robert Jenkin, and St. George Mivart in particular) is treated
at length18.
Towards the end of the 19th century, as if in anticipation of the interaction of heredity with
evolution, we fnd the work of August Weismann19. Connecting the 19th with the 20th century we
fnd Darwin’s nephew, Francis Galton20 and his friend and colleague Karl Pearson21, laying the
foundation of the science of statistics.
The widest circle is the fourth. It properly begins in the year 1900, with the discovery of the
paper written 35 years previously by Gregor Mendel22, and is dedicated to the contribution of
genetics to evolution. The perimeter includes the American geneticist Thomas Hunt Morgan23
– and a retrospective discussion of the inheritance of acquired characters, with social ideology
and political interference causing a scientifc rift between East and West24. It includes the three
great founders of the mathematical Theory of Evolution – Ronald A. Fisher, John B.S. Haldane,
and Sewall Wright25, preceded by an introduction to population genetics theory26. The perimeter
includes the great formalizers of the Modern Synthesis of evolution – Theodosius Dobzhansky and
Ernst Mayr27 – who put it all together, despite objectors like the “heretic” Richard Goldschmidt28.
The following chapters discuss the study of natural selection in feld populations (“Ecological
Genetics”) and Motoo Kimura’s neutrality theory of molecular evolution29, which dominated the
study of evolution towards the end of the 20th century.
In this book I have attempted to provide readers who are interested in evolution with some
knowledge – however brief and incomplete – regarding the major contributors to the theory. The fact
that the book was originally written as a series of independent lectures necessitated my inserting in
later chapters, notes referring to preceding ones, for the sake of continuity. I realize that the texts I
referred to, some of them published over 100 years ago, are not readily available, and have included
many quotations from the original works – if only to avoid misinterpretation.
My decision to publish this book was reinforced by the comments of colleagues:
9 See Chapter 4.
10 See Chapter 5.
11 See Chapter 6.
12 See Chapter 7.
13 See Chapters 8 and 9.
14 See Chapters 15 and 16.
15 See Chapter 11.
16 See Chapter 12.
17 See Chapter 19.
18 See Chapter 14.
19 See Chapters 18-19.
20 See Chapter 21.
21 See Chapter 22.
22 See Chapter 23.
23 See Chapter 25.
24 See Chapter 26.
25 See Chapters 28, 29, 30, respectively.
26 See Chapter 27.
27 See Chapters 32 and 33, respectively
28 See Chapter 34.
29 See Chapter 35.
viii Milestones in the Evolving Theory of Evolution
As many teachers of science have noted, scientifc problems are normally better understood from
studying their history than their logic30.
The history of science forces us to think more than science alone can reveal31.
Finally, I cannot but sympathize with David Quammen32, who expressed his disappointment that
a PhD in evolutionary theory may be obtained at universities without the required reading of even
one sentence of Darwin’s works. What a shame, and what a loss for the students!
30 Sapp 2001, p. 10.
31 J.K. Nester, cited by Wood and Orel, 2001, p. IX.
32 Quammen 2006.
Introduction
Science is a human activity, motivated by hope, cultural prejudices, and the pursuit of glory, but
stumbling in the erratic path towards a better understanding of Nature33.
The American geneticist and evolutionist, Theodosius Dobzhansky, once wrote: “Nothing in
biology makes sense except in the light of evolution”34. Darwin’s introduction of the theory in 1859
made it a cornerstone in the history of human culture and a turning point in the approach to the
study of nature, with important consequences for other sciences as well as for human society.
Like every other achievement in science, the present Theory of Evolution is not the product of a
single active mind, Charles Darwin’s, but has emerged from a background of ideas contributed by
many savants; it also has not remained in the same form as that introduced by Darwin in 1859. The
Theory of Evolution has evolved – and many great biologists have contributed to its evolution, as I
seek to show in this book.
The Starting Point: Evolution versus Creation
Clearly, there could be no evolution before the appearance of life on earth. A living organism, as the
name implies, is an ordered, organized system – be it as simple as a single cell or as complex as a
man. The creation of order from disorder is opposed to the general tendency of things (entropy) and
requires the investment of energy.
The biblical book of Genesis describes the creation of order from chaos. The process of creating
the world, from disorder to the living world – man included, was accomplished in only six days.
To accomplish this in such a short time certainly would require the immense energy of a Supreme
Power. The Theory of Evolution suggests, in contrast, that the biological world may not have
required an extraordinary source of energy if it had taken place gradually over a very very long
time. The point of departure for the study of evolution is that the animals and plants, with which we
are familiar, evolved gradually over a very long sequence of years, from some unknown non-living
matter.
The Theory of Evolution has encountered, since the beginning, the aggressive opposition of
those who considered the biblical stories as historical truth. The Creationists believe that organisms
were created by the Almighty, in their present form, and that that form has not changed since
Creation. An extreme example of the extent to which that Creationist belief can render a person
blind to the facts, may be the book “Omphalos” [Greek: navel], published in 1857 by Philip Henry
Gosse35. Gosse was a British biologist (he had lived for a few years in Canada as a young man) who
published research articles on insects and marine organisms. In his book, published just two years
before Darwin’s “Origin”, Gosse describes – in conventional terms – the geological evidence of the
gradual accumulation of particles to form the layers of rocks, one on top of the other, a process that
seemingly required millions of years. He describes the presence of different fossils in different rock
layers, as indications of changes of the fauna in different ages. He describes the “annual rings” in the
trunks of trees, with one or two being added every year as the tree grows – and similar well-known
biological phenomena. The reader is taken by surprise when, at the end of each section, without
33 Gould 1982a.
34 Dobzhansky 1964: American Zoologist 44: 449 (413-452).
35 Gosse 1998.