THE

LIFE ORGANIC
The Theoretical Biology Club and the Roots of
Epigenetics

Erik L. Peterson

UNIVERSITY OF PITTSBURGH PRESS

 Published by the University of Pittsburgh Press, Pittsburgh, Pa., 15260
Copyright © 2016, University of Pittsburgh Press
All rights reserved
Manufactured in the United States of America
Printed on acid-free paper
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Cataloging-in-Publication data is available from the Library of Congress

ISBN 13: 978-0-8229-4466-9

ISBN 10: 0-8229-4466-9

Cover art: Joseph Needham, Dorothy Needham, and Jean Brachet at the Hopkins Building on Tennis Court
Road, Cambridge, 1938. Photograph reproduced with permission from the archive of the Department of
Biochemistry, University of Cambridge, with thanks to Robin Hesketh.
Cover design: Michel Vrana

ISBN-13: 978-0-8229-8198-5 (electronic)

 For G & W,
my organisms
The way you can go
isn’t the real way.
The name you can say
isn’t the real name. . . .

Two things, one origin, but different in name, whose identity is mystery.
Mystery of all mysteries!
The door to the hidden.
Lao Tzu, Tao Te Ching

“If one insists on putting the question in those terms,” said Frost, “I think
Waddington has given the best answer. Existence is its own justification. The
tendency to developmental change which we call Evolution is justified by the
fact that it is a general characteristic of biological entities.”
C. S. Lewis, That Hideous Strength

Insistence that the study of organisms requires principles additional to those of

the physical sciences does not imply a dualistic or vitalistic view of nature.
G. G. Simpson, “Biology and the Nature of Science”
 CONTENTS

Acknowledgments

Introduction

1. The First Generation of Organicists

2. Needham’s Revival of Mechanism

3. Socrates and the Principia Biologæ

4. The Tipping Point

5. Waddington and the Organizer

6. The Original Theoretical Biology Club

7. Large Plans versus the Ultimate Littleness of Things

8. As Many Opinions as There Are Men

9. “Off in all directions like an expanding universe”

10. Mechanism Reduced to Molecules

11. The Lysenko Morality Tale and the Epigenetic Landscape

12. Ernst Mayr, Neo-Darwinism, and Beanbag Genetics

13. History of Science Is Written by the Laureates

14. The 1960s Reincarnation of the Debate

15. The Conventional Wisdom of the Dominant Group, or COWDUNG

Conclusion: A Third Way after Waddington?

Epilogue: Is Modern Epigenetics Organic?

Notes

Bibliography

Index
 ACKNOWLEDGMENTS

I am humbled by the number of people that need to be thanked for a single, no

doubt imperfect, enterprise such as this. It was by no means through my own
efforts that the Cambridge Theoretical Biology Club gets to reunite on these
pages.
Working like an archaeologist, from the upper strata downwards, we begin
with Abby Collier, the enthusiastic and clear communicator, and her team at
University of Pittsburgh Press. I am grateful for the comments of Marsha
Richmond and the reviewers who gave copiously of their time. Their work was
rigorous and insightful throughout, and they corrected my errors of commission
and omission with a light hand, which I appreciated to no small degree.
Frank Bowles, Robin Hesketh, John Moffett, Hans-Joachim Niemann,
Marjorie Senechal, and Gary Werskey selflessly gave of their time to help solve
mysteries, locate images, and spur me onward. I hope that this result will
contribute in a tiny way to their own important work. Tom Cornford and his
family—Louisa Polak, Sam, Jacob, Dora, Conrad Steel, and Charlotte Sewell—
welcomed a complete stranger on some quixotic (pun intended) quest into their
home for tea; how can I repay their effusive kindness? The Honourable N.
Avrion Mitchison offered warm memories of Sir Peter Medawar and the postwar
Oxford-based Theoretical Biology Club. Many thanks to the archivists and
librarians across the United Kingdom and United States who answered a
fusillade of questions, often with a smile. George T. Thompson has been a
stalwart advocate and a sage. Emma Wilson helped conceptualize the
information in the epilogue. D. Jay Cervino worked the last-minute magic that
saved me from (some forms of) mental illness.
At a still more foundational level, Margaret Peacock and Janek Wasserman
helped whip this unruly animal into some semblance of discipline. My debt to
them will have to go on a payment plan. Brian K. Hall and David Depew saw
this project in an early form and inserted their needed perspectives. Dan
Nicholson has been a stalwart co-laborer on this organicism stuff. Charles Wolfe
opened a number of doors; I still owe him.
Deeper still, Greg Macklem and Elise Crull offered sympathy and some
candies. Daniel J. McKaughan, A. Brant Cook, Brian Estabrook, and Brad Pritts
patiently endured late nights attempting to wrangle these ideas. Phillip Sloan,
Thomas Stapleford, and Christopher Hamlin have acted as foster parents for this
research for a number of years now. Bob Bulkeley has always been my target
audience and provided wind in the sails repeatedly. Barbara Peterson continues
to be my greatest teacher. Brooke W. B. Peterson has long done the things only
bedrock can do.

In every hair, there are an infinite number of lions, and in addition all the single hairs, together with
their infinite number of lions, in turn enter into a single hair. In this way, the progression is infinite,
like the jewels in Celestial Lord Indra’s net.
Fa-tsang (ca. 643–712)
 INTRODUCTION

N ew Yorkers had been anticipating his visit for months. At Columbia

University, where French intellectual Henri Bergson (1859–1941) was to give
twelve lectures in February 1913, expectations were especially high. When first
approached by officials at Columbia, he had asked for a small seminar room
where he could directly interact with students and faculty—something that fit
both his personality and his speaking style. But Columbia sensed a potential
spectacle. They instead put him in the three-hundred-plus-seat lecture theater in
Havemeyer Hall. That much attention, Bergson insisted, would make him too
nervous to speak in English without notes. Columbia persisted. So, because
rhetorical presentation was as important to him as the words themselves,
Bergson delivered his first American lecture entirely in French. 1 Among the
standing-room-only throng of professors and editors were New York journalists
and “well-dressed” and “overdressed” women, all fumbling to make sense of
Bergson’s “Spiritualité et Liberté” that slushy evening. Between their otherwise
dry lines of copy, the reporters’ incredulity was nearly audible as they recorded
how hundreds of New Yorkers strained to hear this “frail, thin, small sized man
with sunken cheeks” practically whisper an entire lecture on metaphysics in
French. 2
That was only a prelude. Bergson’s “Free Will versus Determinism” lecture
on Tuesday, February 4th—once again delivered in his barely audible French—
caused the academic equivalent of a riot. Two thousand people attempted to
cram themselves into Havemeyer. Hundreds of hopeful New Yorkers were
denied access; long queues of the disappointed snaked around the building and
lingered in the slush. According to legend, motorists on Broadway slowed to a
standstill to observe the spectacle. Between the goose-necking and the
meandering pools of onlookers, Bergson’s lecture instigated the first traffic jam
in the history of Broadway. 3 Though the traffic did not become a problem again,
this level of overcrowding continued through nearly all of his Columbia
University lectures. Bergson clearly touched a nerve in New York. He had nearly
everywhere he spoke.
He claimed, self-deprecatingly, to have no well-articulated philosophical
system, nothing to champion, no reason for his audiences to be this large, this
engaged. His supporters in Europe and North America knew better. Bergson was
a philosopher, of course, but his brand of philosophy felt fresh and new, not just
a rehashing of Kant or Aristotle. Some called it vitalism . Living things, Bergson
believed, possessed an élan vital, some animating force or spirit irreducible to
mere chemistry or physics. Organisms, and by extension humans, were not mere
automatons; according to Bergson, we were special. It was Bergson’s notion of
organic specialness—and the individualistic self-determination that seemed to
come with the vitalism—that resonated with audiences in the early twentieth
century.
Yet the ideas that motivated Bergson and drew his crowds were not originally
his ideas at all. Two decades earlier, in Naples, Italy, a young German named
Hans Driesch (1867–1941) squinted through his microscope at a developing sea
urchin embryo. It had just divided: one egg into two gelatinous cells. He had
decided to repeat an earlier experiment done by Driesch’s well-known mentor,
Wilhelm Roux. Roux had used frogs’ eggs and had lanced one of the two
developing cells with a hot needle, killing it. The second cell from that egg lived
and continued to grow and divide, but it formed only half a tadpole. This result
made sense to Roux. Any complex system that lost half of itself would, at best,
form only halfway. But Driesch saw something else there, and he altered the
experiment. Instead of killing one cell with a needle, he painstakingly teased the
two cells apart, making them float alone. After several trials, the floating cells,
separated from their other halves, grew not into half sea urchins, but full ones
with all of their parts in their right places. What kind of machine, Driesch
wondered, could lose half of itself yet continue to develop into a whole,
functioning system? Only a living thing had the ability to do this. So, he thought,
living things must be exceptional, crucially different from nonliving things, and
this insight led Driesch to vitalism.
The ripples of Driesch’s experiment and his endorsement of vitalism tore
open an older debate that appeared to have been settled once and for all decades
earlier. By the beginning of the twentieth century, Driesch, together with
Bergson and others, stood for a reinvigorated vitalism against what they saw as
the dehumanizing hegemony of mechanism. Theirs were only the first salvoes;
this new battle between mechanism and vitalism would continue for decades.
The mechanism-vitalism debate originated in a set of perennial questions.
Aside from superficial dissimilarities in size, shape, and density, what makes
mountains distinct from mountain lions, stars unlike starfish, or sea anenomies
crucially different than the seawater surrounding them? Do organisms possess
certain attributes—the ability to replicate, for instance, or the need to breathe—
that fundamentally set them apart from nonorganisms? Put another way, what
makes something alive? Is organic life essentially different from the inorganic
and nonliving, or is it just a wetter way of assembling the carbon, hydrogen,
oxygen, and so on that chemists find in the earth and the earthworm alike?
Traditionally, mechanists believe there is no real line between life and nonlife;
vitalists believe that without something extra, something outside of physics and
chemistry alone, nonliving material cannot live.
These questions preoccupied some of history’s most influential natural
philosophers. Long ago, Aristotle and Theophrastus wrestled over them while
walking along the Aegean coastline; the Philosopher—Aristotle—insisted living
beings were greater than the cold material causes imagined by Democritus and
the atomists. Five centuries after Aristotle, Galen of Pergamon, surgeon to the
gladiators, insisted that arterial blood carried pneuma , a vital fluid or spirit,
unique to the living. Over a thousand years after Galen, in the seventeenth
century, Britain’s William Harvey corrected Galen’s antique theories of anatomy
but retained an almost commonsense faith in the need for an extra fluid or spirit
to account for the complexity of life—that hallmark of vitalism echoed in Henri
Bergson’s élan vital. Harvey’s seventeenth-century contemporary René
Descartes, by contrast, maintained that organisms were little more than automata
—mechanical devices. The modern conflict between mechanism and vitalism
was born.
It continued to involve some of the most important thinkers in seventeenth-,
eighteenth-, and nineteenth-century Europe. Gottfried Wilhelm Leibniz weighed
in; so did John Turberville Needham, Lazzaro Spallanzani, Georg Ernst Stahl,
William Hunter, and many others. The debate died down in the mid-nineteenth
century due to, among other things, a decisive advance from four prominent
German mechanists. Emil Du Bois-Reymond, Carl Ludwig, Ernst Brücke, and
Hermann von Helmholtz vowed to explain life itself and every function of an
organism solely through physicochemical means. Wilhelm Roux’s work was in
the lineage of these mechanists.
Surprisingly, given the effectiveness of the “Helmholtz school,” vitalism was
reinvigorated in the wake of Hans Driesch’s embryological work. But it settled
little. Once again, an aggressive mechanistic opposition met the new vitalism,
this time through the work of Jacques Loeb (1859–1924). Like Driesch, Loeb
was an adroit experimentalist who emigrated from his native Germany to better-
funded American laboratories.
Sparked by Driesch, Bergson, and Loeb, the mechanist-vitalist dispute
continued on into the new century. Between the First and Second World Wars,
individual biologists, including Nobel laureates, would declare the defeat of one
or the other position. Decades later, Francis H. C. Crick (1916–2004) reminded
his audience that his work in molecular biology contributed to the final conquest
of vitalism. But in the 1960s, another group of accomplished biologists gathered
in Alpbach, Switzerland, to claim that Crick had overstated the strength of his
mechanistic position. Some biologists took an even stronger stand for vitalism.
Sewall G. Wright, founding father of the neo-Darwinian synthesis, for instance,
held that some extramaterial substance or principle was required to explain the
vast complexity of life and evolution. 4 Clearly, the conflict was still very much
alive in the middle of the twentieth century.
Like any conflict, the mechanism-vitalism debate has gained a lot of attention
from historians and philosophers over the years. This makes sense: The battle
lasted so long and involved some of the most memorable names in biology. But,
important as that debate is, this is not a book about it.

Instead, this book maps a comparatively understudied network of scientists who

shared two features in common. The first is that they were not vitalists like
Bergson and Driesch; the second is that they were not mechanists like Roux and
Loeb, either. What they were instead of these two things was hard to define,
even for them. Using their own idioms, each claimed that they were pioneers of
a “third way,” a path that cut across the mechanism-vitalism debate. They
advocated a new manner of seeing —the way that, when given a fourth point and
a triangle, one might create a pyramid instead of a trapezoid. It was a way of
seeing life, even the universe, organically; advocates often called their approach
the “philosophy of organism,” “organic philosophy,” “organismalism,” or
“organicism.”
Their concepts require unpacking, not because they are counterintuitive or
even unfamiliar, but because their “third way” was much more complex than the
mechanism or vitalism it was intended to displace. There were more threads to
hold in tension. Stated as a positive definition, the philosophy of organism was
the belief that a whole organism is “as essential to an explanation of its elements
as its elements are to an explanation of the organism .” 5 More frequently,
however, advocates presented their organic philosophy simply as a negative—a
rejection of the other two ways. As philosopher Morton Beckner pointed out in
the middle of the twentieth century, this “third way” “may be described as an
attempt to achieve the aims of the murky organismic-vitalistic tradition, without
appeal to vital entities.” 6 These “third way” biologists commonly insisted that
highlighting the functional dependence of individual organs and elements (e.g.,
cells) on whole organisms was not an argument for vitalism, it was merely an
argument against the mechanists’ tendency to explain only parts, with the
implication that the explanation of parts sufficed to define the wholes of which
they were a part. As it turns out, this strategy—to define and attack the
weaknesses of the opposition without being able to offer a positive replacement
for that position—has been practiced so often in the history of science that we
might envision it as the ordinary manner of theory change. 7
The Life Organic , then, is an attempt to trace this sometimes murky “third
way” tradition through its derivations and renegotiations. It is also an attempt to
trace a network of individuals —often outsiders for political or cultural reasons
as well as scientific ones—who were advocating for the acceptance of this “third
way.” Aside from a set of scientific concepts, organicists approached the world
as richer and more complex than mechanists, less mysterious and inscrutable
than vitalists.
I make four main arguments in The Life Organic . First, I argue that the
century-long history of the organic philosophy, including the scientists who
defined and refined it, deserves a place at the table with the rest of the history of
biology—with the mechanism-vitalism debate, for instance, over which much
ink has already been spilled. Secondly, I argue that the subfield of epigenetics as
originally conceived was a product of “third way” thinking—an observation that
has important consequences for the headline-making epigenetics of the twenty-
first century. Third, I maintain that the organic philosophy did not die out, as has
been claimed by some historians and scientists, but has continued to find
prominent supporters among scientists of all stripes through the twentieth
century and into the present. Finally, I argue that the discipline of biology visible
through this historical lens seems not to conform to three widely referenced
models for how biological concepts have changed: the models introduced by
Julian Huxley, Karl Popper, and Thomas Kuhn. Let me explain each of these
arguments in more detail.
First, why does this century-long history of the “third way” matter? Excellent
accounts have been written on the origins and development of cell theory, the
neo-Darwinian or “modern” synthesis, molecular biology, the race for the double
helix of DNA, theoretical population genetics, and so on. Considerably less
attention has been drawn to areas outside of these important foci of modern
research, despite the fact that a half century ago, Herbert Butterfield implored
historians of science to look beyond “the emergence of the views that we now
regard as right.” 8 As The Life Organic shows, organicism, though shoved to the
margins over the last portion of the twentieth century for being not “right,” has
been a guiding philosophy of a significant number of scholars on both sides of
the Atlantic Ocean for over a century. It motivated members of multiple
disciplinary subfields, and it impacted both their experimental practices and the
reflective pieces they later wrote about their work. In the 1970s, for instance,
commentators spoke about a crisis in the life sciences regarding the persistence
of biology as a field autonomous from biochemistry and, ultimately, physics.
One could witness this crisis playing out, as philosopher J. Ronald Munson
noted, just by observing the research pursuits biologists chose: “If a biologist is
convinced that it is only a matter of time before we have physical explanations
for all biological phenomena, then he will be inclined to choose research
problems that are amenable to treatment by the methods and theories of physics
and chemistry. Rather than studying the functions and interrelations of cells and
groups of cells, for example, he will concentrate on the chemical processes that
take place within cells.” Commitments to these theoretical concepts have been
“instrumental in directing the future course of biology,” thought Munson. 9 For
this reason alone, perhaps, we should include organicism in our standard
histories as one of those core concepts that has influenced the course of the life
sciences.
But it is not enough to argue that the history of organicism should be included
in order to address a lacuna in the historical literature. 10 When we step back and
take a broader look at the story of organic philosophy over the course of the late
nineteenth and twentieth centuries, it is clear that—my second argument—it
played a central role in the creation of epigenetics .
One need only listen to the morning news or glance at a popular science blog
to learn that epigenetics is the hot new approach sweeping across the life
sciences. These reports radically exaggerate its novelty, however. Epigenetics
did not originate from a set of laboratory experiments in the 1990s or 2000s. It
emerged from discussions in the 1930s and early ’40s surrounding the definition
of the “third way.” Moreover, epigenetics was rooted in a specific place and in a
specific social context: a group of scholars drawn from across England and the
Continent known as the Theoretical Biology Club. The identities and trajectories
of some of these figures comprise the core narrative of this book.
Commentators present twenty-first-century epigenetics as an amendment to
standard genetic accounts of inheritance. Processes most often mentioned in
twenty-first-century epigenetics, such as DNA methylation, insert a scintillating
hint of Lamarckism into the orthodox story. 11 But in its original context,
epigenetics offered something far more complex than occasional breaches of the
central dogma: a way to conceptualize the organism as it develops from
genotype to phenotype. 12 Unpacking the black box of the developing organism
was a central concern of the original formulators of epigenetics. Ironically, lost
amid the hyperbole regarding twenty-first-century epigenetics is the reason why
development was so important to its mid-twentieth-century founders—namely,
the role of development in evolution. They believed that the contours of the
evolutionary landscape are written not in the language of DNA alone, as it is
often misleadingly implied, but via the poorly understood syntax by which
genotypes develop into phenotypes and phenotypes reciprocally alter genotypes.
My third argument is a historiographical one that flows out of the second
historical point: Historians and scientists have claimed that the “third way” faded
into obscurity after World War II, with historians suggesting that the concept was
so tightly tied to fascism in the 1930s that the defeat of the Third Reich also
signaled the demise of organic holism. If and when fascination with the organic
whole did percolate upward in the second half of the twentieth century, they
have claimed, it was often in service of New Age groups embracing a back-to-
nature movement or some species of anti-intellectualism. 13 Yet the long history
of the “third way” outlined in this book does not correlate with either of these
cultural-political trends. Quite often, scientists on the political left—communists,
Fabians, even Christian Socialists—found organicism compelling. And rather
than being the purview of anti-intellectual or anti-establishment radicals,
Cambridge dons and Stanford physicists championed the organic philosophy
through the middle of the twentieth century—hardened experimentalists with
solid records of empirical research.
By my lights, this history reveals something more profound than what other
scholars have suggested. Instead of self-marginalization or adoption by fringe
groups, here we witness a sustained aversion toward nonmechanistic thinking by
a central group defending biological orthodoxy (or, at least as often, wholesale
apathy about theoretical biology). It was this tendency on the part of notable
scientists that pushed the “third way” to the margins, even though it did not
extinguish the philosophy entirely. Perhaps the newfound popularity of
epigenetics in the twenty-first century will reverse this trend, spurring renewed
interest in theoretical biology and an increase in the status of theoretical research
among important life scientists. Unfortunately, recent statements by respected
science popularizers who discount the relevance of philosophy to science make
this scenario seem both desperately needed and terribly unlikely.
Finally, my fourth argument: An examination of the century-long time line on
offer here raises significant questions about how we should model the
development of the discipline of biology. Since the middle of the twentieth
century, the models most often applied to concept development in biology
include the synthesis model championed by Julian Huxley, the Popperian
conjecture-refutation model, and Thomas Kuhn’s model of wholesale paradigm
shifts. While The Life Organic makes no claims to be a detailed contribution to
the history of the philosophy of biology, it seems worth noting that none of these
three models fit the narrative presented here. Each model is predicated on the
notion of replacement —some theory takes the place of another one. Huxley
showed how neo-Darwinism replaced old Darwinism by synthesizing it with
Mendelism. Popper stressed that concepts are floated until some experiment or
set of experiments falsifies that concept. But it is Kuhn’s language of paradigm
shifts that historians and social scientists most often invoke to explain broad-
spectrum concept change in science. 14
Ordinarily, thought Kuhn, scientists are involved in solving various small
puzzles in their fields, and they typically adopt the dominant terminology and
methodology in which they were trained to address those puzzles. Over time,
anomalies develop—unexpected results, puzzles that persistently resist solutions.
When enough of these anomalies accumulate, the time is ripe, believed Kuhn,
for scientists to reject the entire collection of concepts, the language, methods,
and even the worldview in which they were trained. In a sweeping revolution
that Kuhn compared to a gestalt switch, these scientists would adopt a new
paradigm. Admittedly, the revolution might last for a while—perhaps until the
old guard of scientists died off—but eventually it would culminate with a new
paradigm replacing the old one. Naturally, that new paradigm solved the old,
seemingly unsolvable anomalies. But it also helped with puzzles unrecognized in
the old paradigm. In this way scientific progress could be preserved in the
historical account.
Though Kuhn insinuated that his model applied to all natural sciences, he was
particularly interested in what he called the “Copernican revolution.” In his 1957
book with that title, Kuhn carefully illuminated the path by which sixteenth- and
seventeenth-century European natural philosophers overturned the millennia-old
geocentric model of the solar system. Such a revolution, Kuhn showed, required
new concepts of motion, the composition of elements, mathematics, and even the
way we acquire knowledge, as well as an alternate cosmology. The Copernican
revolution and the more recent revolution in physics and astronomy inspired by
Albert Einstein’s work strongly suggested to Kuhn that such change was
unidirectional. It would be ridiculous, for instance, to return to a pre-Copernican
system. That would involve rejecting not only the whole of modern astronomy,
but modern physics as well. In other words, according to Kuhn’s model of
scientific change, once the revolution has happened, there is no going back.
Instead of resolution or synthesis, revolution or replacement, the long history
of organicism presented here reveals recycling and coexistence—a messier,
back-and-forth, almost pendular action. Mechanism has seemed at times to be on
the verge of Kuhnian replacement by some anti-mechanistic challenger. But
mechanism always returns. Surprisingly, the reverse is also true: Despite popular
accounts that celebrate the ultimate dominance of mechanism—Crick’s Of
Molecules and Men (1962) is among the most enduring—mechanism never
successfully silences its opposition either. Instead, mechanism and anti-
mechanism perpetually dance around each other, a yin and a yang. One is
dominant for a time, the other dismissed as useless knowledge of the past. Then
the once-discarded, discredited concept buoys up to the surface of biology again,
reincarnated, old wine poured into new wineskins. 15

Why, if this history is as important as I claim, have historians paid so little

attention to it? A number of reasons come to mind. First, the main advocates for
“third way” thinking were always a minority and often at the edges of the
mainstream of their discipline during their lifetimes. Secondly, they generated
less heat, and therefore drew less attention, than those who squarely argued for
mechanism or vitalism. Thirdly, organicists often had a difficult time explaining
their alternative in any light other than its rejection of mechanism, which goes
some way toward explaining why historians, when they discuss these scholars at
all, have lumped them with more vociferous, more straightforward vitalists.
Finally, even the notion that biologists should be worried about long-standing
questions regarding the ontology and epistemology of their field is controversial.
As we will see at multiple points in this account, during the course of the
twentieth century the practice of biology became increasingly segregated from
the business of thinking about what biology means. Unlike the trajectory of
physics, for instance, theoretical biology was not always crucial to the training
and interests of life scientists later in the century. Advocates for an alternative to
mechanism and vitalism could be safely ignored, even derided as mere
philosophers .
By contrast, much earlier in the twentieth century, biologists often extolled
the theoretical aspect of the field as their raison d’être. When scientists and
philosophers took positions on the mechanism-vitalism debate, especially once
joined by Bergson, the standoff began to seep onto the pages of popular
newspapers and magazines. By this point, a generation of biologists in the UK
and United States were already engaged in formulating the “third way”
alternative—they will be the focus of chapter 1 . Still, it was not until the advent
of the First World War, and the palpable triumph of mechanized death over any
romantic notion of organic life, that a critical mass of English-speaking scholars
turned their attention toward examining the debate. The most concentrated
public forum for that examination took place at the first American Philosophical
Association (APA) meeting held after the 1918 Armistice. By the second decade
of the twentieth century, physics had already undergone a revolution in its most
basic concepts beginning with the work of Einstein and Bohr. Biology seemed
close behind. Cambridge-mathematician-turned-Harvard-philosopher Alfred
North Whitehead persuasively argued that the universe was not constructed
primarily of isolated particles or mechanisms but relational wholes or organisms
. He called his approach the philosophy of organism, and it resounded through
European intellectual circles. (Indeed, ripples of their holistic alternative
permeated that field where the word organic still connotes a similar meaning:
agriculture. A persuasive early call for organic farming made by Oxford’s Walter
James, 4th Baron Northbourne, entitled Look to the Land [1940], borrows
heavily from both Whitehead and a convinced Whiteheadian statesman, Jan
Christiaan Smuts. 16 )
The story did not end there; the philosophical shift signaled by Whitehead
was not yet a fully experimental approach. Intending to sharpen and extend the
work of Whitehead and the first generation of organicists, one group in particular
met repeatedly over the 1930s to sketch out what this resolution could mean in
the laboratory and for society. Called by some of its members the “Theoretical
Biology Club,” this network of polymathic, transdisciplinary British men and
women (with occasional prominent visitors from the Continent and North
America) worked throughout the 1930s to clarify what exactly was meant by
concepts like organic . 17 Consistent members of this group included biochemists
Joseph and Dorothy Needham, biomathematician Dorothy “Dot” Wrinch,
biophysicist John Desmond “Sage” Bernal, embryologist-geneticist Conrad H.
“Wad” Waddington, and—the glue that held the group together—Joseph Henry
“Socrates” Woodger. The interactions of the members of this club (detailed in
chapters 2 through 8 ) incubated some of the most fruitful organicist work. Most
significantly, it was in the context of his work with the Theoretical Biology Club
that Waddington took on the project that would make him best known:
epigenetics. Thus, epigenetics was brought to life through the search for the
“third way” embodied in the discussions of the Theoretical Biology Club and the
work that Waddington conducted with members of the club. Epigenetics, as
originally conceived, was organic. What that meant, exactly, would have to wait
to be spelled out until after the Second World War.
In retrospect, the 1930s were the early zenith of the organic philosophy.
While the pendulum had swung away from the vitalism exemplified by Bergson
and Driesch, it did not cease swinging. Instead, the war and the ensuing geo-
political tension between the United States and the USSR as highlighted by the
Lysenko story served as a backdrop for the steady re-entrenchment of
mechanism. 18 From the new analytical philosophy of science to the growing
insistence that DNA was at the heart of nearly every important biological issue,
the organic philosophy counted fewer open adherents. Despite the overwhelming
popularity of epigenetics in the twenty-first century, Waddington’s approach,
first formulated in the 1930s and ’40s, did not attract a large and devoted
following in the decades after the Second World War. Funding and training were
devoted to molecular biology. By the time of the deaths of the key members of
the Theoretical Biology Club in the 1970s and ’80s, popular and more
aggressively mechanistic concepts like sociobiology and selfish genes further
muted the influence of organicism. The marginalization of the “third way” in the
second half of the century, and Waddington’s perpetual struggle for its
recognition against conventional wisdom regarding gene-centric mechanism, is
at the heart of chapters 9 through 15 . Finally, an epilogue reexamines the
present standing of epigenetics in light of its roots in organicism using the
analytical tools of the digital humanities.
Given the status of epigenetics today, one might believe The Life Organic
details a triumphal story of professional recognition and scientific celebration—a
“comedy,” as Hayden White once styled it. I should say up front that this is not
that kind of story. Organicists did not join the pantheon of great biologists
alongside Darwin, Pasteur, Medawar, and Crick. 19 Nevertheless, I believe this
historical account enlightens our present conception of the living world. By the
beginning of the twenty-first century, at the very moment that a new version of
epigenetics was drawing significant popular attention and funding,
developmental biologist Scott Gilbert and philosopher Sahotra Sarkar argued
that the organicism of the early twentieth century needed to be dusted off and
brought back into the conversation. 20 They join a cadre of other scientists and
philosophers who together assert that there is something about the organic
philosophy that deserves reclaiming in the twenty-first century. 21 The Life
Organic , then, is their story as well.
 1

THE FIRST GENERATION OF ORGANICISTS

E dmund Duncan Montgomery (1835–1911) died mostly alone and mostly

unsung, except by the laconic, grinding chorus of cicadas among the oaks on his
ex-plantation outside of Hempstead, Texas. His neighbors—if they thought about
him at all—likely regarded him as an eccentric, a hermit, and a foreigner to boot.
They knew he had a background in medicine, a strange accent, and a passing
interest in local politics, but other than this, they knew little about his past and
seemed to care even less. His late wife, on the other hand, was more interesting.
She had been a famous artist, Elisabet Ney (1833–1907)—the only touch of
European cosmopolitanism to pass through this remote village. Sadly, her
elegance had departed from Hempstead even before her death: She lived in
Austin, over a hundred miles west of her husband, for most of the closing years
of their lives. 1 (Remnants of Ney’s Austin studio became the core of Texas’s
first true art museum.) As they laid Montgomery in the ground next to Ney,
under oaks they planted decades earlier specifically for this funerary purpose, his
neighbors could not realize that Montgomery had slowly, patiently, almost
single-handedly chiseled through one of the hardest issues in the history of
biology—What separates living from nonliving things? From this out-of-the-way
corner of America, Montgomery would pioneer a worldview that deviated from
Bergson’s and Driesch’s and Loeb’s—neither vitalism nor mechanism, a “third
way.” His views would echo through biological science into the twentieth
century, rippling through even the conversations of the Theoretical Biology Club
in the 1930s.

Many years earlier, in a very different part of the world, Montgomery was a bold
young man of Scottish aristocratic upbringing busily sculpting an illustrious
career in medicine. After receiving university training at Heidelberg, Berlin,
Bonn, and Würzburg under some of the most prominent names in mid-
nineteenth-century European science, including Karl Vogt and Hermann von
Helmholtz, Montgomery acted as the resident physician at London’s German
Hospital. In 1860, he earned an appointment as Demonstrator in Morbid
Anatomy at St. Thomas’s Hospital. 2 There were the usual obligations of such
positions, yet Montgomery found the time to begin studying protozoa on the side
—an auspicious choice given the growth in interest in cells following the work
of Schleiden, Schwann, and Virchow in the 1840s and ’50s. By the early 1860s,
Montgomery had successfully leveraged his work on protozoa into a network of
scientific relationships through British scientific society. He even conducted a
physiological demonstration at the London Zoo, where he met a curious Charles
Darwin, who had recently published On the Origin of Species . 3
Montgomery’s star began to fall almost as quickly as it had risen, however. In
1863, he learned he had contracted tuberculosis. This diagnosis initiated a nearly
decade-long period of wandering around Europe seeking relief. He traveled to
Funchal on the Portuguese Atlantic island of Madeira, then to Mentone, Italy,
then to Rome, then Munich. He was able to practice medicine in each case. He
even managed to publish a few scientific articles when his health improved. In
truth, however, his health was only one factor for the wandering; his path was
equally dictated by his desire to be near Elisabet Ney. So began a four-decade
relationship that waxed and waned between intense and romantic on the one
hand, sedate and platonic on the other. When it appeared by the mid-1860s that
Montgomery’s condition had worsened, Ney traveled to Madeira. The two
hurriedly wed. As soon as Montgomery began to recover, however, Ney slipped
away to pursue her burgeoning art career. To outsiders, their relationship never
felt more settled or definite than this: Through most of their marriage, she
continued to go by “Miss Ney,” she traveled abroad for long stretches without
Montgomery, and she possibly became pregnant in 1870 by Ludwig II of
Bavaria, whose image she was sculpting. 4 Montgomery never shared whatever
thoughts he had about these matters.
In the midst of his wandering phase, in 1866, Montgomery was invited to
London’s Royal Society to deliver the paper that became his entrée into Anglo-
American biology, “On the Importance of So-Called Cells in Animal Bodies.”
His essay—sprinkled with allusions to Vogt—confronted some of the
implications of the cell theory so much in vogue at the time. Despite all the
hoopla surrounding them at that moment, cells themselves had no vital powers;
they appeared very much like wet crystals, in fact. Moreover, argued
Montgomery, they could be artificially manipulated into altering their forms
from one crystalline formation to another. They were certainly nothing unique.
The reception to his paper was mostly positive. Richard Owen, the eminent
anatomist and occasional nemesis of Charles Darwin and T. H. Huxley, declared
the essay an “important contribution to the philosophy of physiology.” 5 But at
least one American reviewer balked, “These are startling statements, and should,
at least, be carefully verified before they are accepted.” 6
Another, more important audience also remained unconvinced: Though
sources reveal little directly about their conversations in the early 1860s, we can
guess that Montgomery could not win over Charles Darwin. 7 Since the
publication of Origin , Darwin had searched for a concept of inheritance and
variation that would support natural selection. He was hesitant about the
purported power of cells to explain any sort of biological phenomena, yet his
skepticism never extended beyond vague doubts. In his two-volume The
Variation of Animals and Plants Under Domestication (1868), Darwin
tentatively explored pangenesis—an old eighteenth-century idea accounting for
how organisms such as starfish and earthworms could regenerate after being cut
into pieces. In the book’s second edition, Darwin responded to a growing
number of critics of the concept, including Montgomery. 8 Though understanding
their difficulties with pangenesis—and after mulling over Montgomery’s concept
that “cells and tissues of all kinds may be formed, independently of pre-existing
cells”—Darwin nevertheless settled on the notion that organisms must be
comprised of “a multitude of organic units, all of which possess their own proper
attributes,” independently functioning and able to develop into unique units
because of “some inherent power which the cells possess and not to any external
agency.” In other words, when prodded into a defense, Darwin found support for
his pangenesis hypothesis from a “cellular doctrine,” ignoring Montgomery’s
critique of it. 9 After reading Darwin’s conclusion in the new edition of Variation
, Montgomery moved even more decisively against cell theory, publishing
articles in The Quarterly Journal of Microscopical Science and the Transactions
of the Pathological Society of London , among others.
Everything changed in December 1870. Ney’s artistic career took an
unspecified but clearly unfortunate turn after she became pregnant while far
from Montgomery in Ludwig II’s court. Montgomery again complained that his
health was on the decline. So the couple followed the suggestion of their
erstwhile friend Baron Carl Vicco Otto Friedrich Constantin von Stralendorff,
who had moved recently to an experimental tuberculosis colony in Thomasville,
Georgia (United States), and had married an American woman, Margaret. In
January 1871, Montgomery and Ney took with them their servant Crescentia
“Cencie” Simath and fled Bavaria, leaving behind all of Ney’s most prized
pieces. They hoped to find a simpler, healthier life with von Stralendorff in the
small Reconstruction-era resort town.
Sadly, nothing worked out as planned. Humidity did little to help the two
European consumptives. Cencie contracted malaria. Then Ney’s child, Arthur,
contracted the disease. She survived; he did not. After only two years the von
Stralendorffs retreated back to Europe. Vicco perished soon after their return.
Montgomery and Ney contemplated following the Stralendorffs back across the
Atlantic, but Elisabet was pregnant with her second child, and Montgomery was
still desperate to settle in a drier climate. The couple moved farther west, settling
at “Liendo,” a recently abandoned Texas cotton plantation halfway between
Houston and Austin. There, in that unlikely spot, they lived out the remaining
four decades of their lives, sometimes together, sometimes not. 10
Montgomery resumed scientific work on protozoa in Darwin-like seclusion.
His findings quickly began to sharpen. In the 1860s, in that Royal Society essay
—subsequently republished in journals across Europe—Montgomery had
appeared to be a run-of-the-mill anti-vitalist. But after his move to the United
States, he began arguing just as strongly against the Vogt-like mechanism he
once seemed to support. Montgomery’s conversion was not to vitalism but away
from the growing faith in the power of aggregated mechanical units. “The
distinct morphological divisions of higher animals are, indeed, integrant, not
constituent parts,” Montgomery insisted. This is what makes them exceptional:
“They are specialized and segregated from a pre-existing whole and are in no
way discrete and independent units joined together in the composition of a
complex totality. . . . The whole is here in all reality antecedent to its parts. The
organism is prior to its tissues, the tissues prior to their supposed elements. The
centralized organism is not, as universally assumed, a multiple of ultimate units
but is, on the contrary, itself one single individuality.” 11 In this two-part article,
“The Unity of the Organic Individual” (1880) and in subsequent publications,
such as “Are We ‘Cell-Aggregates’?” (1882), Montgomery argued that the
executive level of an organism could not possibly be the solitary element or unit.
These small parts always functioned as subordinate components of tissues,
insisted Montgomery, and tissues always functioned as part of still larger
systems. Only when in the form of integrated organisms could individual units
resist the buffeting of the external world by regulating their own internal
environments. In other words, whole organisms became exceptions to the
ordinary rules of physics and chemistry—even if their constituent parts were not
imbued with some sort of vital property.
Montgomery’s most direct impact was on European biologists. His work
appeared in major European journals of science and philosophy, including Mind ,
International Journal of Ethics , Jahresberichteder Anataomie und Physiologie ,
and Monist , to name just a few. He cogently attacked Darwin’s theory of
pangenesis and gained the respect of a number of prominent British biologists. 12
He became well known to cutting-edge German physiologists, including Jacques
Loeb and, especially, Hans Driesch, with whom the aging Montgomery
corresponded until his death. In fact, Driesch’s 1907 Gifford Lectures, later
published as Science and Philosophy of the Organism , was a specific response
to charges leveled at his vitalism by Montgomery. 13 American scientists, by
contrast, did not typically mention Montgomery. Perhaps his absence from
discussions of vitalism and mechanism in the United States was due merely to
timing. Only five years after the publication of Montgomery’s Philosophical
Problems in Light of Vital Organization , and less than one since Montgomery’s
death in Texas, Jacques Loeb published The Mechanistic Conception of Life
(Chicago: University of Chicago Press, 1912). 14 The following year, Columbia
University welcomed Henri Bergson with great fanfare. Had Montgomery lived
through these moments in the second decade of the twentieth century—and had
he felt comfortable enough to leave his Texas plantation to address Loeb and
Driesch from a stage in, say, Boston or New York—historians might speak of a
multiway debate between mechanists, vitalists, and organicists in the early
twentieth century rather than the binary debate we ordinarily discuss.
Edmund Montgomery may have been a John the Baptist, crying from the
Texas wilderness that mechanistic explanations of organisms in terms of cell
parts could not be sufficient, but he was far from the only prophet of the “third
way.” A number of scholars on both sides of the Atlantic comprised a first
generation offering alternatives to mechanism and vitalism. A rough
contemporary of Montgomery’s in France, Claude Bernard, had touched on
some of these issues. 15 But the first real efflorescence of “third way” scholars
emerged during the last few decades of the nineteenth century and the first few
of the twentieth. These included Charles Otis Whitman and Lawrence J.
Henderson in the United States and Conwy Lloyd Morgan, John Scott Haldane,
and Edward Stuart Russell in the United Kingdom. This list is far from complete
—many more scholars attacked both mechanism and vitalism in the first decades
of the twentieth century—but it is representative of organicism prior to the
Theoretical Biology Club of the 1930s. 16 Like Montgomery, Whitman and E. S.
Russell stressed the inability of aggregated parts to explain whole organisms.
Lloyd Morgan contributed the twin notions of “gappiness” and “emergence”—
apparent gaps between levels of complexity are not imaginary products of
accumulated tiny alterations; rather, different levels might play by different
rules. Henderson and Haldane elaborated the internal regulatory and external
buffering abilities exclusive to whole organisms. What follows is a brief survey
of each of their contributions.
First, though, permit an aside regarding past portrayals of these figures.
While Whitman, Russell, Lloyd Morgan, Henderson, Haldane, and others
reflected that era’s skepticism regarding the belief that mechanistic science
would lead to perpetual social progress, they embraced neither the lionization of
irrationalism nor the post-Schopenhauer pessimism associated with the fin-de-
siècle period, sometimes called the romantic period. 17 In other words, they did
not see themselves as particularly romantic biologists. 18 Collectively, their work
prepared the intellectual soil for the “third way” biology that followed,
influencing the Theoretical Biology Club and, eventually, Waddington’s
epigenetics.

Charles Otis Whitman (1842–1910) . In 1975, the aged former wartime foe of
the United States, Emperor Hirohito, stood on the shores of Cape Cod,
Massachusetts. He bowed with the graceful stiffness that belongs only to
dignitaries with years of practice as he accepted a small gift from his hosts. It
was a photo of the main building of the Woods Hole Marine Biology Laboratory
(now Woods Hole Oceanographic Institution), named after Charles Otis
Whitman. On the surface, this was an odd gift for the once-deified leader of the
Empire of Japan. Yet it was Hirohito himself who initiated the trek to Woods
Hole as part of his tour of the United States. That side trip meant far more to him
than the more publicized visit with President Ford at the White House or his
excursion to California’s Disneyland: The emperor had spent much of his life
developing an expertise in marine biology. He first became acquainted with the
field as a boy at Gakushuin (then The Peers School). This is where the otherwise
strange connection between Hirohito and Whitman lies. His instructors at
Gakushuin had become engaged in the subject decades earlier when a visiting
professor from the United States established marine biological research at the
University of Tokyo. That professor was Charles Otis Whitman. 19
Like Charles Darwin, Whitman found his way into biology via birds, which
he studied for much of his life. But by 1873, he had fallen under the spell of
Darwin’s Swiss opponent at Harvard University, Louis Agassiz, who convinced
Whitman to attend Agassiz’s new summer school in marine biology at the
Penikese Island station in the Elizabeth Islands, off Cape Cod. There, Whitman
met E. S. Morse, one of Agassiz’s influential students, who lured Whitman to
Tokyo in 1880. Because of Whitman’s experience in Tokyo; at the Naples, Italy
marine biology station; and at the Allis Lake Laboratory in Milwaukee,
Wisconsin, Whitman was chosen by United States Fish Commissioner Spencer
Fullerton Baird to head the new Marine Biological Laboratory at Woods Hole.
By all accounts, his leadership established Woods Hole as the premiere
marine biology station in the eastern United States. 20 As Woods Hole director
(1893–1908) and as zoologist at the newly founded University of Chicago
(1892–1910), Whitman came into contact with a large portion of the practicing
zoologists in the country and thus produced a particular “style” of zoology that
accompanied his style of management at Chicago and Woods Hole. 21 He
directly impacted many of his students’ research projects during and after their
student years. Some, like Ralph and Frank Lillie, continued to pursue the same
questions of development and evolution that motivated Whitman.
Whitman vehemently disagreed with Roux’s mosaic concept of development,
wherein different portions of the organism possessed separate, nonredundant
developmental programs; it was especially problematic when wedded to cell
theory. According to Whitman, the adherents of cell theory, Schleiden and
Schwann included, regarded “organization” as reducible to cellular structure,
“ontogeny” as merely cell creation. But Whitman, like Montgomery, remained
unconvinced. Perhaps our confidence in the power of the cell was a product of
the intensity of our search.

Our microscopes resolve the organism into cells, and ontogeny shows that the many cells arise
from one cell; hence the organism seems to be the product of cell-formation and the cleavage of the
germ seems to be a building process. . . . All the search-lights of the biological sciences have been
turned upon the cell . . . it has been searched inside and out, experimented upon, and studied in its
manifold relationship as a unit of form and function. It has been taken as the key to ontogeny and
phylogeny, and on it theories of heredity and variation have been built. 22

Certainly, there was a great deal of attention being paid to the cell. But
Whitman, much like Montgomery, could not be convinced that cells truly were
the architects of all organic form and function. Some other processes begin their
existence only when collections of cells congregate in a particular state of
organization. Turn away from what Whitman derided as the “cell-doctrine” or
“dogma” and one would begin to see the “organism-standpoint”—certain
organic components can change when and how they function in a global sense.
For example, a fragment of hydra might, if separated from the whole,
nevertheless reproduce a whole organism; it could act as if it was a whole even if
it was only a part. Should a pair of blastomeres in the developing amphioxus (a
bottom-dwelling warm-water lancet) become separated from the rest of the
developing organism—recall Driesch’s experiments on sea urchin embryos—the
isolated portion of blastomeres would at first continue to produce cells only
corresponding to their original position. After a brief time, however, this isolated
portion would reconfigure itself to produce a whole organism. From these
examples, Whitman drew the lesson that form and function are both independent
of the number of component cells. Cells multiply, but the organ remains the
same throughout. So far as homology is concerned, the existence of cells may be
ignored.

[A]n organism is an organism from the egg onward. . . . [C]ontinuity of organization would be the
essential thing, while division into cell-territories might be a matter of quite secondary importance.
. . . Continuity of organization does not of course mean preformed organs, it means only that a
definite structural foundation must be taken as the starting-point of each organism, and that the
organism is not multiplied by cell-division, but rather continued as an individuality through all
stages of transformation and sub-division into cells. 23

Whitman feared it was too easy to create reductive explanations relying upon
the smallest detectable particles. He contrasted the particle-centric approach with
the organism perspective that focused not on the static cell but on the process of
development. Development, too, must be the way by which individuality is
maintained even while the body fluidly rearranges its structure. After all, the
adult butterfly is every bit an individual as when it was chrysalis, caterpillar, and
egg even if its bodily shape, eating habits, behavioral patterns, and every one of
its cells have been replaced by something different in both form and function. If
cell division, cell interaction, and the cell itself were the wrong places to look,
where should biologists focus their attention? Though he toyed with a neologism
—“idiosome”—Whitman could neither define nor specifically locate these
hereditary determinants of structure that, in turn, delineated function.

Conwy Lloyd Morgan (1852–1936) . Across the Atlantic Ocean, a contemporary

of both Whitman and Montgomery struggled with similar problems. Lloyd
Morgan trained in philosophy at a young age, but mining and metallurgy seemed
like more lucrative paths. He could not escape his earlier interests, however. As a
student at the London School of Mines, Lloyd Morgan stumbled across T. H.
Huxley’s “The Physical Basis of Life” and found it troubling. Interestingly, he
also felt reciprocal shame at being troubled . After all, why shouldn’t even
complex phenomena—human cognition, for instance—be determined by
physics? As he admitted, thinking and feeling seem to be greater activities than
the mere physical state of molecules in the brain, but are they? It was the
beginning of a lifelong passion, addressing the question, “Just how did a science
of mind link up with this science of the physical basis of mind?” 24
By a twist of fate, Lloyd Morgan was seated next to T. H. Huxley himself at
an annual student and staff dinner in 1873. True to his usual form, Huxley
questioned and cajoled the young mining student over the clinking of plates and
glasses; he talked with Lloyd Morgan long after Huxley completed his own
address. Though awed by the presence of “Darwin’s Bulldog,” Lloyd Morgan
handled himself well enough that, at the close of the evening, Huxley suggested
that Lloyd Morgan study under him for a year.
The year with Huxley passed. By then Lloyd Morgan felt well enough
equipped to tackle the large biotheoretical questions that percolated through
Victorian society in the mid-1870s. But unlike many of his peers, Lloyd Morgan
was not from a family of means. He needed to find steady employment. He tried
his hand as a mining assayer, but his heart was no longer in it. After spending a
few years lecturing on the side, he finally quit his mining job in 1878 to accept a
full-time lectureship at the Diocesan College (better known as Bishops) in the
small town of Rondebosch, South Africa, on the fringes of the British Empire.
Despite his training, Lloyd Morgan taught courses far outside his expertise,
including British Literature and Constitutional History. He enjoyed the
experience. The Bishops schedule allowed him plenty of time in the evenings
and on holidays to read and think about science and the philosophy of science.
While it is certainly conceivable that Lloyd Morgan had already come to the
attention of the English university system due to his connections with Huxley,
that is not what brought him back to England. His return journey from South
Africa began on the pages of Nature in 1882. There Lloyd Morgan, prep-school
tutor from a remote corner of the empire, critiqued George Romanes, the well-
heeled, brilliant disciple of Charles Darwin, on Romanes’s favorite subject: the
concept of animal intelligence. Romanes took no offense at the challenge from a
young outsider. In fact, Romanes magnanimously backed the publication of
Lloyd Morgan’s The Springs of Conduct . 25 Lloyd Morgan used that work to
leverage a lectureship in geology and zoology at a startup undergraduate
institution in Bristol, England. This new school would eventually grow into
University College Bristol (now Bristol University).
His challenge of Romanes paid additional, completely unexpected dividends.
Upon Romanes’s untimely death in 1894, it fell to Lloyd Morgan to shepherd the
final uncompleted volume of Romanes’s magnum opus through publication,
contributing more than half of the content to complete the multivolume Darwin,
and After Darwin . 26 This, too, was an auspicious move. By the turn of the
century, Darwin and most of his ardent supporters had passed on. By chance
then, C. Lloyd Morgan, an ex-mining assayer and former prep school jack-of-all-
trades, ascended to be a major sculptor of the debates surrounding mechanism,
vitalism, and evolution at the turn of the twentieth century.
Lloyd Morgan published in Monist , the philosophy journal also favored by
Montgomery. In two essays in particular, “The Doctrine of Auta” (1893) and
“Three Aspects of Monism” (1894) Lloyd Morgan charted the direction this new
“third way” would take in biology and psychology. 27 As he envisioned it,
monism referred to the following three conceptions. First, it denied that subject
and object could be understood completely independently of each other. We have
one experience that contains two aspects, flip sides of one coin. Though the
author , a subject, writes the manuscript , an object, the two different aspects are
intimately interwoven in the moment of writing, the experience . Acknowledging
monism in the realm of knowledge means that we have to regard the entirety of
the act as of seamless importance. In any given moment, said Lloyd Morgan,
there is no clear distinction between a seer , the process of seeing , and the thing
seen . We only produce clear subjects and clear objects upon reflection. This
point would be developed soon after in the work of Alfred North Whitehead and
taken up again by C. H. Waddington decades later.
Lloyd Morgan also recognized what he called monism of “natural descent,”
which meant that any feature observed at any organismal level must be a feature
shared at levels below it. (This feature of Lloyd Morgan’s thought would be
echoed in Theoretical Biology Club meetings as well.) Dualists, at least since
Descartes if not the ancient Greeks, regarded mind and matter as separate.
Monists, according to Lloyd Morgan, regarded “mind” not as extranatural or
superadded in any way but as a product of evolution. All living things, in other
words, must exhibit mind to a degree; the biological and the psychological must
be conjoined. By working directly in animal behavior, Lloyd Morgan could
justifiably say that he had authority to speak about the barest, most basic aspects
of life as well as human psychology—these phenomena were different in degree,
not kind. Finally, he introduced “analytic monism,” though Lloyd Morgan
struggled to define what he meant by this final tenet. At best, he could say that
analytic monism was neither mechanistic materialism nor “psychism.”
Mechanistic materialism must classify consciousness and sentience as illusory or
epiphenomenal, a not-real property of our incorrect perceptions. Psychism, an
equally ancient idea drawing on gnosticism (though Lloyd Morgan only traced
the idea to the Reverend Charles Kingsley, the famous admirer of Darwin and
author of the wildly popular Waterbabies ), emphasized that matter was just the
gritty, temporary shell to be sloughed off upon death. Neither mechanistic
materialism nor “psychism,” thought Lloyd Morgan, captured what it was to be a
living thing. For Lloyd Morgan, we simply must believe that monism is true and
regard all attempts at segregating life/mind/spirit from body/material as
destructive oversimplifications. 28
His monistic path charted, Lloyd Morgan looked for ways to test it—or,
rather, to apply it in experimental form. He was deeply committed to the totality
of common descent with modification; that is, to the evolution of mind in
tandem with body. So he adopted and cared for different types of animals in the
hopes of discovering behavioral traits that prefigured those in higher organisms.
In his Introduction to Comparative Psychology (London: W. Scott, 1894) and
subsequent books, he mentioned working with insects, birds, and small
mammals. 29 One imagines a house full of crickets, mice, dogs, and sparrows,
with Lloyd Morgan seated in their midst, field notebook in hand. Late in life, he
could still be seen walking across windy Clifton Downs, near Bristol, notebook
and old ball in hand, dog by his side, ready to conduct observations.
Surrounded by organisms, Lloyd Morgan soon found a flaw in Darwin’s
account of gradual descent with modification. Common descent did not mean we
should imagine that less complex organisms exhibited the same traits as more
complex ones but on a shrunken scale. The spaces between the traits appeared
large, abrupt, full of gaps. The gaps in mental constitution between amoebas and
grasshoppers and mice and chimpanzees and humans, for instance, seemed
titanic. The more he compared birds and mammals and insects to one another
and to human behavior, the more alien each way of sensing and organizing the
world seemed to be from the others. Even in physiologically similar organisms,
psychological gaps manifested themselves. Physiological common descent
Lloyd Morgan could continue to regard as an a priori truth. Behavior gaps
between organisms, however, could not be dismissed so easily. But how could
we explain these apparent gaps? In answering this question, Lloyd Morgan
struck upon the idea of emergentism that, together with his commitment to
monism, became a foundational premise of the “third way.” 30
Emergent Evolution , the book derived from his early-1920s Gifford Lectures
at the University of St. Andrews, remains a touchstone of first-generation
organicism. For Lloyd Morgan, emergence was an intuitive, not to say simple,
notion. Imagine three levels of natural events, A, B, and C. On level B, there
exists a structure of relationships, even a manner of relating , that does not exist
in level A. The same is true of C. If one had some experience on the B level, one
could not predict the characters of level C; truly, the characteristics of the C level
have not come about yet because the C way of relating has not yet come about in
B. The thing that cannot be predicted from B is the new kind of relatedness, the
new way of fashioning relationships even between already extant phenomena.
That comes into being—that emerges—at level C. Perhaps the concept would be
more concrete if we name A “quantum,” B “molecular,” and C “cellular.” The
rules whereby relationships work at the quantum level tell us little or nothing
about how relationships between components will work at the chemical or
biological level. It’s not that the quantum level doesn’t operate at the cellular
level, it’s that the quantum level doesn’t tell us much of relevance—it is a
problem of our knowledge. Emergent evolution, then, is merely the recognition
that new phases of the same underlying stuff have radically different,
unpredictable characteristics, and that these traits have come about seemingly de
novo over the course of time. 31
Lloyd Morgan understood how his audience might view notions of erupting
phases—new configurations arising seemingly from scratch—and he tried to
head off criticisms from both extremes. Afraid emergent evolution would be
cataloged with vitalism, Lloyd Morgan insisted that

on the evidence, not only atoms and molecules, but organisms and minds are susceptible of
treatment by scientific methods fundamentally of like kind; that all belong to one tissue of events. .
. . In other words the position is that, in a philosophy based on the procedure sanctioned by progress
in scientific research and thought, the advent of novelty of any kind is loyally to be accepted
wherever it is found, without invoking any extra-natural Power (Force, Entelechy, Elan , or God)
through the efficient activity of which the observed facts may be explained. 32

No extra-natural force was necessary because the emergence of novelty itself
should be understood as a fully natural process, a possibility in the initial
conditions of the universe. Yet Lloyd Morgan argued just as forcefully against
his critics that this emergent evolution was not mere mechanism in a fancy
wrapping:

It is pretty certain that the interpretation of nature I put forward will, in some quarters, be . . .
spoken of as “the mechanistic dogma.” The odd thing here is that the whole doctrine of emergence
is a continued protest against mechanical interpretation, and the very antithesis to one that is
mechanistic. It does not interpret life in terms of physics and chemistry. It does not interpret mind
in terms of receptor-patterns and neuron-routes. Those who suppose that it does so, wholly
misapprehend its purport. 33

A committed monist who regarded life and thought as interlaced, real features of
the world—not just the steam created by the molecules of nervous tissue in the
brain shifting in space—would then be led to bite the proverbial bullet, admitting
that life and mind are impregnated into the very fabric of the universe. What we
recognize as conscious life, sentience, even logic, evolved as reconfigurations in
the way the underlying mind-stuff relates to itself. The typical mechanistic
depiction of life and mind regarded them as mere aggregations and regroupings
of physicochemical events, algebraically parsable. Vitalism demanded a new
force or substance superimposed on ordinary inert material from outside of
nature. Lloyd Morgan insisted that life and mind were unified with material and
emerged through natural, though unforeseen, departures in the evolutionary
passage of physicochemical events.
In his own day, C. Lloyd Morgan’s attempt at defining the middle way
between mechanism and vitalism went unrealized. Opponents saw his work as
too overtly philosophical to also be scientific. In the either–or intellectual
environment of the early twentieth century, Lloyd Morgan’s notion of gaps was
often read as support for vitalism. His obituary practically begged readers to
ignore his “metaphysics” in order to consider fairly his comparative psychology.
Nevertheless, the effects of Lloyd Morgan’s work in philosophy of biology
continue to be felt today.

John Scott Haldane (1860–1936) . The First World War is often nicknamed “the
war of the chemists.” But when a chemist attacked Britain, Britain turned to a
biologist. When German troops under the direction of future Nobel laureate Fritz
Haber unleashed yellow-green gas at Ypres on 22 April 1915, the British War
Office whisked Scottish physiologist J. S. Haldane to the front lines. Haldane
speedily conducted autopsies of two soldiers who had stumbled off the
battlefield, succumbing to asphyxiation hours later. The metal buttons on their
greatcoats were corroded; their esophagi showed extensive tissue damage. He
immediately suspected chlorine gas. By the time he returned to Britain, the
newspapers were already posting his suspicions: “The Germans have again made
use of this diabolical contrivance,” howled the Times . “The report of Dr.
Haldane . . . brings home . . . beyond the possibility of doubt, the deliberate
resort to this atrocious method of warfare.” 34
John Scott Haldane’s role as a scientific witness at the dawn of modern
chemical warfare was no accident. For decades he focused on the physiology of
human and animal respiration, often using himself and his family members as
experimental subjects. These years of study on what to us might seem like
strictly practical problems—how to build an effective gas mask, for one—
instead led Haldane to deeply philosophical questions. Montgomery looked
through a dusty microscope in Texas; Whitman bred pigeons in Chicago; C.
Lloyd Morgan walked misty Clifton Downs with a dog at his side; Haldane
found the “third way” in the heavy, damp darkness of a coal mine.
He did not set out to study mining, as Lloyd Morgan had, but somehow
Haldane ended up making more contributions to the health and welfare of coal
miners than any mining engineer before him. Born into one of Britain’s
intellectually elite families, Haldane received his MD from the peerless
University of Edinburgh Medical School in 1884, then traveled to Jena to study
rigorous, German-style physiology before taking his first post as Demonstrator
in Physiology at University College Dundee. Haldane was well on his way to
being a scientifically trained physician when his maternal uncle, John S. Burdon-
Sanderson, recruited him to Oxford University in 1887. Burdon-Sanderson
occupied the Waynflete Professorship of Physiology, and he hired his nephew on
as a demonstrator.
The relationship was a rocky one, however, and within three years Haldane
was nearly fed up. “I think my uncle Dr. Burdon-Sanderson and myself could
hardly be more at loggerheads than we are in our ideas about the aims and
results of physiology,” he wrote his betrothed, Louisa Trotter, in October 1890.
“I can’t help thinking and writing often the exact opposite of what he does.” 35
Still, Haldane labored to mend their relationship for a time. Aside from
conducting the experiments on the constancy of animal heat and energy—this
was the sort of physics-heavy physiological work his uncle wanted him to do—
Haldane buried himself in work on animal respiration and the regulation of
chemicals in the body. But Burdon Sanderson took a dim view of his nephew’s
interests, especially as Haldane’s respiration work edged closer and closer to
applied science and public health.
In the summer of 1894, Haldane had the opportunity to put this new
knowledge to use. An explosion occurred at the Albion Colliery at Clifynydd in
south Wales. Though such explosions were frighteningly common in the rich
coal seams, it was difficult to explain with scientific precision why they were so
deadly. Was it simply the violence of the explosion? Did the ignition of the coal
dust burn all of the breathable air out of the mine? Was the fire especially
intense? Haldane plunged down into the tunnels, handled miners’ bodies,
measured gasses, and analyzed injuries. He visited several more mines that year
and into the next. Upon returning to Oxford, he tested every theory carefully—
more often than not running the tests on himself. His verdict: Poisonous carbon
monoxide built up rapidly during a coal-dust fire. Even if unscathed by the
explosion itself, any miners unable to scramble to the surface would soon be
asphyxiated. 36
Haldane’s habit of dashing down into mines and then conducting follow-up
breathing experiments on himself struck Burdon-Sanderson as too cavalier,
certainly not how an elite man of science should conduct himself. 37 Moreover,
Haldane maintained a disturbing aloofness from Burdon-Sanderson’s cutting-
edge mechanistic ideas. In 1898, Haldane published “Vitalism” in Nineteenth
Century , the same journal Romanes, Huxley, and other Darwinists had used to
promulgate their ideas. Paradoxically, the article was not an endorsement of
vitalism, but he did declare his uncle’s mechanistic version of physiology
moribund. “The wave of physicochemical speculation,” announced Haldane,
“seems now to have nearly exhausted itself. 38 ” Haldane did not believe that
physical and chemical explanations proved useless in biology—far from it. Yet
the persistent attempt to leverage this limited success into a comprehensive
worldview required a substantial leap of faith that mechanists like his uncle did
their best to conceal. Haldane found no more solace in vitalism, however. The
philosophies of Bergson and Driesch called for a vital something that allowed an
organism to stave off disintegration by borrowing energy from the surrounding
environment. But this idea didn’t make sense: Vital forces required belief in the
expenditure of energy, “which is at the same time not expended, and may even
be growing with expenditure, as in a developing organism.” 39 In other words,
vitalists wanted to watch their cake grow while eating it. Moreover, the élan vital
did a poor job of maintaining living things against degradation and decay. A tiny
bit of poison that would not affect dead egg albumen at all would seriously
damage a living cell. If mechanism couldn’t sustain itself, then neither could
vitalism.
Haldane concluded “Vitalism” by calling for an anti-mechanistic, anti-
vitalistic alternative. He hadn’t worked out the details, exactly, but he knew that
it would start from the assumption that the function of the whole organism
determined its parts’ functions. How far out beyond the individual organism this
holism extended, he could only guess. For ease of investigation, he argued, we
should progress by moving only a small step beyond the examination of
biological function—his domain of physiology—to the study of structure, to
morphology. Morphologists, Haldane thought, understood better than
physiologists two key features of organisms: First, that a general body “plan” of
the organism persists even in the face of “disturbing influences”; and, second,
that organisms replace, to the degree they are able, parts that have been removed,
and can maintain functionality even without their full complement of parts. 40
This general outline would continue to motivate Haldane through the rest of his
career. Ripples of it would appear in the works of other organicists.
After securing a degree of notoriety for his work understanding respiration in
coal mines, deep sea diving, and mountain climbing, Haldane returned to
develop his “third way” alternative in Mechanism, Life, and Personality and
Organism and Environment as Illustrated by the Physiology of Breathing . 41 He
believed he had found a way out of the mechanism-vitalism conundrum—a way
inspired by Claude Bernard’s “organicism,” claimed Haldane. For Bernard,
organicism provided the resolution by discussing how the internal complexities
of living entities related to nonliving environments. 42 Now Haldane could revive
organicism by examining the regulation of animal respiration under extreme and
variable environmental stress. 43
Much like C. Lloyd Morgan, Haldane appealed to monism (i.e., there is but
one quality of material substance in the world) and emergentism. Haldane meant
that the peculiarities of live organisms—their unpredictability, their ability to
self-regulate within certain constraints, and so on—could potentially be shared
by what we would ordinarily call nonliving materials. In certain contexts,
however, new properties emerge in such a way that we might perceive them. The
objects we perceived as sharing these emergent properties we would probably
call “alive.” That wouldn’t mean that these objects were alive because something
had been impressed upon inert matter from the outside, and nonliving things
weren’t simply lumps of matter that hadn’t been granted alive status. The
distinctiveness between alive and not-truly-alive had everything to do with
human perception. At present, according to Haldane, our perception was limited
—we simply didn’t know how to see emergent properties in all physicochemical
systems. When we did learn how to see properly, we would no longer be enticed
to reduce biology to chemistry and physics. Quite the opposite: The famous
epigram attributed to Haldane was, “If physics and biology one day meet, and
one of the two is swallowed up, that one will not be biology.” 44
While his activity in studying respiration continued unabated, Haldane
returned repeatedly to outline his alternative philosophy in ever more detail,
even in Respiration , his best-known scientific work. 45 After three decades of
excellent physiological work, some of which inspired very practical solutions to
the serious dangers of chemical warfare and mining, accolades began to pile up
from the scientific and technical world. But by the 1920s, Haldane had wandered
out of the proverbial cave and into the penetrating light of philosophy. When by
the end of the decade he turned to retrieve those scientists he regarded as still in
chains inside the cave, he was surprised to see so many fellow biologists already
free. Just before the First World War, Haldane believed he had been the lone
voice interested in the larger philosophical implications of biology. Now
biologists from his generation stood alongside him to articulate the organicist
alternative.

Lawrence J. Henderson (1878–1942) . The Great War itself provided some of

the impetus for revived interest in the mechanism-vitalism debate. Vitalists were
fond of seeing the war as an indictment of the mechanistic world picture, with
Germany and Austria-Hungary engaged in living out the logical conclusions of
that philosophy. 46 By the close of the war, Hans Driesch was pressing his attack
on mechanism, Bergson was continuing his speaking tour extolling vitalism and
free will, and Jacques Loeb was frantically promoting a mechanistic worldview
against the incursions of Driesch and Bergson from his position at the
Rockefeller Laboratory in New York City.
In the summer of 1918, the American Philosophical Association (APA)
convened a special committee of five—three scientists and two philosophers—to
subject both mechanism and vitalism to a detailed, impartial examination.
Harvard’s Lawrence J. Henderson would represent chemistry, Herbert S.
Jennings from Johns Hopkins spoke for biology, Howard C. Warren of Princeton
was the psychologist, and Walter T. Marvin (formerly of Princeton’s philosophy
department but by 1918 a dean at Rutgers University) and R. F. Alfred Hoernlé
(Harvard) headed the philosophy segment. Under the direction of APA president
Mary Whiton Calkins, the five met to configure a list of books and articles from
across the spectrum of history, science, and philosophy that they would use to
guide their discussion and, eventually, shape the official APA position. The
committee placed on their reading list the full gamut of positions: Driesch,
Bergson, and other vitalists, mechanistic opponents like Jacques Loeb and Emil
Du Bois-Reymond, and even those who were attempting to bridge the gap—J. S.
Haldane chief among them. Six months later, at the APA annual meeting in
January, the committee members rose to deliver their verdict one by one:
Vitalism was bad science at best, probably poor philosophy as well.
Of course, this was the expected outcome, and Irwin Edman of Columbia
University, chronicler of the committee address, reported the five speeches with
relish. After all, one Great War had already been won; another—the one against
vitalism—was entering its decisive phase. As Edman gazed over the audience,
he noted the warriors returned from the Continental battlefield, now arrayed in
martial finery to observe the latest intellectual artillery barrage: “Philosophers
seemed to have felt that they had done their bit (the records of the War
Department will bear them out, as did the presence of uniforms at the meetings),
and were entitled now to the glorious dissipation of problems.” 47 According to
Edman’s reporting, the action that first week of January 1919 did not disappoint.
Henderson, Jennings, and Warren let fly the first explosive salvoes: Henderson
found patternedness usually associated with organisms in nonliving things,
Jennings discounted Driesch for violating the determinism observed in the
laboratory, and Warren confirmed that even the seeming exception of conscious
animal behavior could be accounted for by ordinary mechanistic explanations.
So, according to Edman, the three “purely neutral scientific inquirers offering
evidence of an unequivocal nature” could find no redeeming value in the
vitalism of Driesch, Bergson, or anyone else. All “confirm[ed] mechanism with
unequivocal evidences drawn from the distinctly mooted areas” of their
expertise. 48
After the scientists softened the enemy up, the philosopher Walter T. Marvin
moved in for a decisive strike. Before the war, Marvin had joined the “new
realism” movement. It was an attempt to loosen the grip of neo-Kantianism that
the new realists believed had strangled once high-powered philosophy programs
on the US East Coast. 49 Now standing on his profession’s most prominent
platform, Marvin fired at the vitalists. Certainly the scientists had adequately
demonstrated that vitalism was unsupported by any empirical evidence. That
was a good start. But as Marvin expounded upon the history of philosophy for
his audience, he unveiled an even more extensive anti-vitalist argument than the
scientists. Vitalism wasn’t merely bad science or bad philosophy; it was
something much darker. The work of Driesch, Bergson, and others was only the
vanguard of a larger backward march toward primitive animistic thinking. In
fact, according to Marvin’s Manichaean history, there had always been only two
alternatives for human civilization: “the powers of light and the powers of
darkness, in its modern transfiguration, intellectualism versus romanticism.” 50
Romantics—i.e., vitalists—denied progress, civilization, the modern spirit. For
Marvin the new realist, vitalism represented everything that enlightened humans
had been gradually, inexorably suffocating since the days of David Hume, if not
Francis Bacon. Vitalism might appear to be solidly entrenched behind regrowing
sea urchin embryos, harmonious equipotential systems, entelechy, and élan vital,
but as the new instantiation of old animism, it would nonetheless be asphyxiated
by the weapons of mechanistic philosophy. As Marvin pronounced it, and as
Edman framed the talks of the committee at the APA, the case was clear: One
must be for empiricism, mechanism, and thereby believe in scientific progress,
or one must be for animism, vitalism, and stand against science, progress, and
the benefits of modern civilization.
The actual positions endorsed by the committee members called on to settle
the debate between mechanism and vitalism were more nuanced, however.
While it is true that none openly supported vitalism, only Walter Marvin (and
perhaps Irwin Edman, the reporter) celebrated a bellicose either–or, friend–foe
dichotomy. At least one of the headlining scientists, Henderson, had sketched out
his own theoretical compromise for several years before being tapped to
represent physics and chemistry at the APA. (Even that choice was somewhat
strange: By 1919 Henderson worked at Harvard squarely in the biology
department.)
Henderson’s interest in the mechanism-vitalism debate grew from 1909 to
1913, when he published The Fitness of the Environment . 51 This book was his
attempt to glean the best out of the pastiche of approaches then available—from
Claude Bernard and Emil Du Bois-Reymond to C. Lloyd Morgan. 52 His organic
compromise began from the sentiment that the universe is, at bottom, not merely
acceptable for life but positively “biocentric”: The features of the universe we
usually relegate to physics and chemistry mesh together to create a seamless
context for the development of the organic. Though Henderson drew some
inspiration from nineteenth-century German anti-vitalists, and even encouraged
the work of well-known mechanist Jacques Loeb after Loeb’s arrival in the
United States, Henderson retained a deep skepticism toward mechanism. 53
Convinced that the organism’s peculiarly flexible system of respiration and
chemical regulation could not be elucidated by the machine analogy, Henderson
developed what he considered to be a new theory of biological teleology . He
called it “organic homeostasis”—the ability of living things to maintain their
own internal environments in the face of external perturbations. He vigorously
and repeatedly insisted that organic homeostasis and his notion of organic
teleology were completely at odds with vitalism; it may have sounded like
Driesch’s “harmonious equipotentiality,” but Henderson meant something quite
distinct from Driesch. It was a tough balancing act. Henderson admitted that
living beings looked and acted different than nonliving matter, but then he tried
to qualify his bioexceptionalism: It had to be superficial, epistemic alone,
predicated on the relative ignorance of biologists. 54
Henderson must have known his version of organic teleology was not a novel
concept, even among American scientists. The famous cell biologist E. B.
Wilson had advocated something like this for cells quite recently; Immanuel
Kant, for that matter, had sketched something like Henderson’s version of
organic teleology in the eighteenth century. Frankly, there were even closer ties
to concepts being espoused by those on the other side of the Atlantic—J. S.
Haldane in particular. And the resemblance between Henderson’s ideas and
Haldane’s only grew as Henderson shifted his research to focus on the delicate
chemical composition of blood. A few years earlier, as the United States geared
up for its belated entry into the Great War, Henderson published his masterpiece,
The Order of Nature ; Haldane, meanwhile, published Organism and
Environment as Illustrated by the Physiology of Breathing (1917). 55 There were
some differences between Haldane and Henderson, of course. Interestingly, it
was the American, Henderson, who was swayed more by the philosophy of
Herbert Spencer. Like Spencer, Henderson held that the appearance of teleology
is not accidental: Organisms must be purpose driven because the universe itself
is purpose driven. This respect for Spencer could sometimes muddle
Henderson’s biophilosophy. Was teleology part of the universe and the organic
world or was it just a convenient fiction that we used to explain biology to
ourselves? Sometimes Henderson seemed to support the purely pragmatic:
Biologists must continue using purpose-based language in biology because there
was no way around it. Relying simply upon physicochemical explanations would
always prove insufficient because humans were stuck seeing purpose in living
things. On other occasions, Henderson supposed that there was no strong in
principle reason why physicochemical explanations would not suffice. Unlike
Henderson—but like Edmund Montgomery, for instance—J. S. Haldane firmly
believed that the more complex terminology of biology, the dynamic regulation
of systems, for example, captured the actual state of organic life. Those
descriptions, according to Haldane, would eventually apply also to the simplest
physicochemical phenomena—the universe was full of purpose even if we
couldn’t detect it. But this was a minor point of contention; Henderson echoed
Haldane and others advocating the organic philosophy more often than he stood
apart. 56 By the time of the APA meeting, Henderson’s real position—as opposed
to the anti-vitalist simulacrum of Henderson presented by Irwin Edman—was
not so idiosyncratic even in the United States. Edmund Montgomery’s “unity of
the organism” brand of organicism may have failed to attract vocal adherents
during his day. By the end of the Great War, however, a growing number of
Americans, like Henderson, joined the British scientists advocating a non-
vitalistic alternative to mechanism.
The first sign that this group had begun to grow into something more
substantial arrived at the same time, with the publication of a comprehensive
work entitled The Unity of the Organism; or, the Organismal Conception of Life
(1919). Over two volumes, its author, William Emerson Ritter (1856–1944), the
director of the University of California’s Scripps Institution for Biological
Research, summarized the organicist alternative to mechanism as it stood by the
close of the Great War. Interestingly, the conception that Ritter cataloged
repeated what Montgomery had codified decades earlier, only now buttressed by
substantial laboratory work, much of it by Henderson and Haldane. The
distinguishing mark of the “third way” was becoming clearer:

The organism taken alive and whole is as essential to an explanation of its elements as its elements
are to an explanation of the organism . . . all attempts to assign explanatory values to the elements
in their relation to the whole organism, while at the same time denying either expressly or tacitly,
similar values to the entire organism in its relations to the elements, must fail in large degree. 57

Edward Stuart Russell (1887–1954) . Chronologically, Glaswegian ichthyologist

E. S. Russell was the last of the first generation of organicists, yet, in some ways,
his views exhibited the most consistency with Montgomery’s original position.
Russell was versed more explicitly than his peers in the persistent theoretical
questions that had preoccupied biologists going back to Aristotle. Moreover, he
had the most direct impact on the generation of organicists that followed, as he
interacted with members of the Cambridge Theoretical Biology Club of the
1930s. Books such as his Form and Function (1916) continued to resonate with
life scientists for decades after his death.
In 1911, however, Russell easily could have been mistaken for a vitalist
alongside Driesch and Bergson. That year Russell published “Vitalism” (which
made no reference to Haldane’s 1898 essay with the same title) in Scientia . In it,
Russell claimed only two options remained available to the biologist: (1)
Concede that biology deals with unique phenomena irreducible to
physicochemical concepts in some sense, or (2) argue that biology should be
absorbed into physics and chemistry. Russell regarded this dichotomy as the
essence of the vitalism-mechanism debate—and Russell thought it was a crisis.
58 He found the horns of the dilemma particularly sharp, because they hid deep

social ramifications. Should mechanism win, biology would cease to exist as an

autonomous science. Should vitalism win, biology would admit gaps in the
physical laws of the universe—gaps that could be filled in only by appeals to
extranatural forces outside of the scope of science. Either way, biology would
cease to exist in its present form. Russell even went so far as to assert that, prima
facie, compromise proved impossible: “Any attempt to reconcile the vitalistic
theories which admit an intersection of the physical and the psychical with the
prevailing mechanistic conceptions of the physical sciences is doomed to
failure.” 59 Nevertheless, Russell hoped to show that, despite Loeb’s vocal
advocacy for mechanism and Driesch’s equally vocal vitalism, he had found a
way off of the dilemma’s horns.
He objected to Loeb’s mechanism for two reasons. First, biology had always
been a science that appreciated its own concrete empirical facts just as much as it
had been about a general scientific methodology. Biologists could potentially
explain observations by appealing only to the limitless cascade of
physicochemical details. Take the example of an eel migration to the North
Atlantic, a subject Russell studied in detail. One could explain the chemical
machinery necessary to swim a great distance, to consume energy and gasses
with the liquid environment, to reproduce, and so on. Even if such a computation
were possible for a human, however, Russell was convinced that description
would be of little worth. One could not deduce that the intricate motions of the
tail in swimming, cellular transport processes, the adjustments to seawater
salinity and temperature, digestion of prey, and so forth, had as their origin an
impulse to migrate to a particular location in order to spawn. The problem was,
according to Russell, chemists would not even see “migration”; they would see
only “movement”: “To the chemist . . . there is no fact of migration at all, there is
only an intricate enravelment [sic ] of chemical reaction.” 60 Since the days of
Aristotle, biologists looked for explanations of a higher order than mere
chemical or physical transactions in muscle tissue. Biologists’ answers involved
the survival of a class of things—a group of eels, for instance.
Secondly, and more importantly for the trajectory of the “third way,” Russell
argued that unlike inert objects, organisms grew into and then maintained
particular shapes or forms throughout their lives, passing on those forms to their
descendants. Mechanistic explanations could not adequately explain organismic
form. Why this form? —much like Why do these eels migrate to the North
Atlantic to spawn? —was a question with an answer rooted in evolutionary
history and dependent upon the dance of organismal development. Physics and
chemistry might be able to parse with mathematical precision the paths of
materials moving through processes involved in the development of form. But
this sort of an explanation would hardly touch the true mystery of development.
The form of an organism existed in “apparent autonomy and invariability, [in]
great independence of environment,” marveled Russell. Any answer as to why
and how form could be maintained would be “beyond the reach of
physicochemical explanation.” 61 The way out from the dilemma posed by
mechanism and vitalism would have to clear these particular hurdles in the
standard mechanistic story. This “third way” would have to rely upon actual
observations of organisms in the wild, take account of environmental pressures,
understand the evolutionary history of the organism, and deal with the problem
of a constantly changing yet somehow constantly stable form. And it would have
to do so without appealing to extranatural forces beyond the reach of science.
This was Russell’s lasting contribution to the “third way” and the subsequent rise
of epigenetics. For Russell, the twin goals of biology were not merely to find
biochemical pathways and gene expressions, it was to explain the path from egg
to adult—how an organism retained a specific form—and how that form was
inherited from one generation to another. Any theory of evolution had to account
for the persistence of organized form, not merely aggregation.
Russell’s 1911 “Vitalism” did not share quite the same fate as Haldane’s 1898
“Vitalism”—it was not rejected so much as ignored. That reception makes some
sense. Russell graduated from Glasgow University with none of the inherited
pedigree of Haldane nor C. Lloyd Morgan’s connection to Darwin’s favorites T.
H. Huxley and George Romanes. While Lloyd Morgan received recognition for
his work at Bristol, and while Haldane garnered international attention pursuing
the secrets of physiology in coal mines and gas masks, Russell labored away as a
largely anonymous government bureaucrat in the Ministry of Agriculture and
Fisheries (MAF) in London. He quietly served there from 1909 until 1921, when
he was appointed director (the equivalent of chief science officer today). He
retired from the MAF in 1947. 62 During World War I, he became acutely aware
of the overfishing problem while working with fishermen on the southwest
English coast. An adept analyst, Russell became at least as well known for
marshaling statistics in favor of environmental conservation as for his anti-
mechanistic philosophy of the organism. From Russell’s perspective, this
statistical ichthyology went hand in hand with organicism.
While the outcry against mechanism was gaining strength during and just
after the war, Russell quietly published two substantial works: Form and
Function (1916) and The Study of Living Things: Prolegomena to a Functional
Biology (1924). 63 In each he argued for “the validity and independence of
biological laws over against the laws of physics and chemistry.” 64 Biological
laws would have to account for more than those captured by the analysis of heat,
respiration, muscle power, and so on: “The living thing,” insisted Russell, “is not
a machine, for it shows persistent and prospective tendency or striving, and its
responses are adjustable to a wide dance of circumstances.” Vitalism, too, was
wrong for similar reasons: “[The living thing] is not a mechanism actuated by a
psyche, for this formulation simply pushes the mystery further back by ascribing
to an immaterial agent or element of nature the faculties and powers of the
organism. The living thing is not the clay moulded by the potter, nor the harp
played upon by the musician. It is the clay modeling itself.” 65 For this reason,
organisms can be compared metaphorically only to “dynamic works of art.” Just
as there is an irretrievable loss when one analytically dissolves a Monet or a
Kandinsky into the vectors of its individual brushstrokes, the chemical
composition of its pigments, the tensile strength of its canvas, and so on, Russell
claimed that organisms could not “be exhausted of their content by the analytic
and superficial description offered by the physical sciences.” 66 Echoing
Haldane, among others, Russell denied that the objective of biology, even
physiology, should be to “study photoreception and chemoreception,
phototropism and chemotropism, and so on.” (One can hear his dismissal of
Loeb’s work on phototropism here.) Instead, we should always be pursuing
“what the organism is driving at in its dealing with the world with which it is
confronted. . . . [O]ne and the same physical stimulus may mean quite different
things”—thus we should consider not only the organism as unbroken unity, a
dynamic piece of art, but also as an irreducible entity embedded in an complex
environment. 67 In other words, all of biology should orient itself in the opposite
of its usual direction. Instead of looking down toward the most minute parts and
explaining the behavior of individuals and groups by reasoning from the smallest
chemical (or genetic) components, biologists should be looking up from the
individual and reasoning from its response to surroundings, its ecological niche.
This perspective, Russell asserted, was a much older and better established
one than the mechanistic picture or its vitalistic counterpart. In the early years of
the war, Russell immersed himself in the history of biology, including its ancient
Greek roots. It became increasingly clear to him when turning to the works of
Aristotle, Cuvier, Karl von Baer, and many others that the “organic” work he
was pursuing as an ichthyologist was more in line with the far vaster goal of
biology since the time of the ancients than it was with the work of his
contemporaries. From Aristotle onward, biologists had attempted to describe the
world as it appeared with all of its intricacies intact. The drawbacks to such an
approach could be significant, of course. For one, what should be done with
thick descriptions of organisms and the environments to which they were
adapting other than natural history? The Functional Biology proscribed in
Russell’s Prolegomena seemed unduly complicated and full of gaps, especially
where the behavior of the whole organism did not seem obviously functional.
These were places where ignorance would fester.
To Russell, it was telling that modern mechanists usually came from subfields
like physiology, genetics, and biochemistry. Certainly, these fields required a
great deal of memorization and quantification, but they were simple. There were
unexplored areas, but they were able to be explained in the usual ways—find a
new gene or a new biochemical. This relative simplicity of genetics or
biochemistry proved comforting to many because, in these subfields, a
researcher could abstract away from messy processes like development and even
questions of evolution. A physiologist like Sherrington, who opposed Haldane,
or a self-described biochemist like Loeb, who opposed any anti-mechanist, could
work on the individual gears in this process without having to think much about
how it related to the organization of that one. That approach worked, of course.
It worked well : Problems could be solved—they were limited problems, to be
sure, but some headway was possible. But from Russell’s perspective, these
fields were too limited to be models. They merely pulled up one corner of the
vast blanket. To throw back the whole mysterious cloth of biology, maintained
Russell, one needed to discover how individual organisms aligned these
functional processes of digestion, circulation, respiration, reproduction, and so
on, to thrive in particular environments, and then how these organisms persist in
these configurations over generations. The properly oriented biologist must
acquire an appreciation for how multiple processes performing biological
functions link together in a particular formal configuration. In other words, form
had to connect up somehow with function .
Actually, Russell insisted that it was even more complex than merely
identifying current form and function. History mattered. It mattered in two
senses in particular, he argued. In an epistemic sense, perhaps obvious, the
discipline of biology itself has a history. Russell feared that twentieth-century
biologists had forgotten this self-evident fact; they had lost what he called
“historical consciousness.” He hoped Form and Function would demonstrate the
longue durée of morphological thinking, thereby reviving the historical
embeddedness of the biological profession. Perhaps Form and Function could
function as a historical reminder and challenge the growing tendency toward
quick payoff studies that shed little light on the true complexity of the biotic
world. Aristotle, Cuvier, Darwin—these names matter not for any hagiographic
sense, but because they remind us that even those concepts of historical biology
we now reject once were supported empirically, more or less. Perhaps we ought
to know what our scientific forerunners knew, even if it is merely to point out
why they were wrong.
Russell also believed history mattered in a second sense, this one more
strictly biological: Organisms have histories, both individually and as a species.
The core form of any given organism had already been shaped by its
evolutionary trajectory—a trajectory with a function—but new functional
necessities had remolded that form slowly, imperfectly. Russell insisted that the
proper attitude for a biologist was not to focus on the invariant rules of physics
and chemistry that would apply equally well to a suspension bridge or a soap
bubble, but to the commonsense notion of organisms as active and purposeful.
He admitted this would not be easy for a biologist. Russell felt that their training
“drilled out” of biologists any appreciation for the purposiveness of organisms
and the problem of form. 68
By the mid-1920s, Russell’s focus on the apparent purposiveness of living
things had overshadowed most of his other claims. 69 And as time went on, he
increasingly highlighted the notion of “drive,” applying modifiers like
irreducible, sui generis, and so on. That rhetoric made Russell a target even of
his ally J. S. Haldane. Haldane feared that by emphasizing teleology as a way to
discredit mechanism, Russell had strayed into vitalistic territory. 70 Though
Russell would attempt to wriggle out of this charge, it is one that stuck with him.
Even decades later, as we will see below, philosopher Ernest Nagel would
dredge up this aspect of Russell’s thought in order to attack organicism as a
whole.

As the 1920s drew to a close, however, these suspicions hardly seemed to matter.
Anti-mechanism, whether of the vitalistic or the organic sort, had hit its mark.
Pronouncements by philosophers at the APA in 1918 had been forgotten,
apparently. Across Europe, other scientists joined the attack against the once
widely accepted view that organisms could be analyzed as wet machines.
Physiologist Jakob J. Von Uexküll and psychologist Kurt Goldstein were but two
of the most prominent promotors of the view that to explain organisms one
needed to invoke anti-mechanistic final causes; like Russell, they dressed the
ancient Aristotelian notion of teleology in twentieth-century scientific
terminology.
In the 1880s, E. D. Montgomery raised his “solitary voice” to challenge the
conventional wisdom of the dominant group, or what he called the “unanimous
verdict of the most competent judges” who supported the “doctrine” of
mechanism. But he despaired that he would never get across to those who were
committed to viewing the organism as merely “cell-aggregate[s]” and
“coordinative machinery” his “deeper view of life and organisation.”
Nevertheless, Montgomery remained convinced “a more faithful interpretation
of nature” would displace mechanism. 71
 Almost a half century had elapsed. A critical mass of scholars did follow the
trail blazed by Montgomery, though he was all but forgotten by then. From
Whitman to Russell, this first generation of organicists demonstrated the
inadequacy of mechanism in multiple ways. They introduced and elaborated
upon notions of emergence and the internal regulation of the organism. It
seemed, perhaps, that mechanism was on the verge of being replaced by
Montgomery’s “more faithful interpretation of nature.” We might be tempted to
call it a paradigm shift in the making.
Yet it was difficult for them to articulate how exactly their “third way” was
distinct from vitalism. For Russell, especially, the lines blurred between an
organic conception of life and a quasi-vitalistic one enamored with final
purposes. As additional scholars, those not as familiar with professional life
science, joined the growing anti-mechanist movement, they adopted the same
teleological language that Russell had been accused of using. Finalists—those
who emphasized teleology in biology—joined the ranks of older vitalists like
Driesch and Bergson, though they sometimes parroted the language of the first
generation of organicists. Thus, by the end of the 1920s, organic had come to
mean purposeful, perhaps even interchangeable with vitalistic .
 2

NEEDHAM’S REVIVAL OF MECHANISM

T he first generation of “third way” biologists attracted the interest of a broader

audience and had perhaps even instigated a strong challenge to mechanism, but
W. E. Ritter’s vision of a unified organic movement among biologists had not
solidified. Vitalism with an emphasis on teleology remained a tenable position.
Predictably, vitalists gained traction with both popular and scientific
audiences when they attacked studies that treated humans as mere wet machines.
One particularly outspoken vitalist, University of Milan philosopher Eugenio
Rignano, targeted a historically famous mechanistic treatise—Parisian gadfly
Julian de La Mettrie’s enormously influential Man a Machine (1748)—in his
own Man Not a Machine (1926). But the reference was only titular: Rignano was
clearly targeting not La Mettrie in the eighteenth century but modern mechanism
flying under the flag of Jacques Loeb and others in the twentieth. Throughout
Man Not a Machine , Rignano accentuated examples of contemporary scientific
work that purportedly extended mechanistic explanations to embryology and
development—particularly studies of respiration and digestion, heredity and
evolution, and even human social behavior. But then Rignano provided a twist.
Even in these “mechanistic” studies, he detected a residue of purposiveness—of
old-fashioned teleology—that the mechanist couldn’t explain away. In
embryonic development, for instance: “The living substance . . . selects from the
very complex mixture of chemical substances dissolved in the nutritive liquid
exactly those compounds or radicals capable of reconstructing it in the same
specificity as before. And as selection , this process has a marked purposive
aspect.” 1 Obviously, given all of this room for teleology in basic functions like
digestion, Man the Thinker could not be just a machine. Humans, although
Rignano hinted that we should treat the whole of the animal kingdom with the
same care, required other, more vitalistic, explanations.
Young Cambridge biochemist Joseph Needham (1900–1995) would have
none of it. He responded aggressively to Rignano’s Man Not a Machine by
resuscitating La Mettrie’s eighteenth-century title, Man a Machine . Needham’s
Man a Machine even mimicked the style of La Mettrie, thumbing his nose at
Rignano with language at turns playful, sardonic, sympathetic, caustic. Though a
generation younger than Loeb, Needham reanimated the sentiments of that
famous mechanist to upend every argument Rignano had advanced to support
the apparent purposiveness of living substances. His attack on Rignano came on
three fronts: (1) The evident purposiveness of living things is, once properly
examined, not that impressive; (2) the teleology that Rignano and other vitalists
identified exclusively in living things did in fact appear in nonliving things; and
finally (3) Rignano’s evidence for purposiveness or teleology was not
quantitative and therefore could be of no significance for science. 2
To each jab of Rignano’s—each example of some biological activity not
reducible to mere physicochemical mechanism—Needham provided two
counterpunches. He landed his first blow by offering reasonable mechanistic
explanations for biological process similar to, perhaps even better known, than
the ones provided by Rignano. Then Needham jabbed again: Each of Rignano’s
examples were mere red herrings—marginal or recently discovered processes
still awaiting complete biochemical analysis. Given enough time, these examples
would be described mechanistically as well. After several such combinations,
Needham stood back, his opponent on the ropes, and confidently asserted that
future biochemical analysis would bring each process under mechanism’s big
tent. When Rignano insisted that a particular aspect of respiration could not be
reduced to the mere action of chemicals—interestingly, a reinterpretation of J. S.
Haldane’s famous work—Needham wound up for a haymaker: Even if one
aspect of respiration seemed to require vital explanation, the most important one,
the oxygen transport mechanism, was far more typical in the organism than
Rignano’s more esoteric examples. And oxygen transport could be understood
completely mechanistically.
On the ropes, perhaps, but still fighting, Rignano recalled the sea urchin work
of Hans Driesch, now decades old:

What do the ultra-mechanists do in the presence of these distinctly teleological manifestations of
the generative and regenerative processes? They direct all their efforts to an attempt to prove that
given chemical substances exercise a morphogenous action on particular developments, hoping to
conclude triumphantly that the entire series of morphogenous phenomena, constituting the
ontogenetic development, may be explained completely and exclusively by physico-chemical
action. But in this attempt they have mistaken a mere release of morphogenous activities already
potentially in the developing embryo for a genuine morphogenous action. 3

Vitalists, in other words, could always parry attempts by mechanists to attach
some new property to whatever chemicals appeared in development; they could
always assert that the directing phenomena—the form or the purpose perhaps—
preexisted and in fact pre-formed the operations of those chemicals at that
moment in time. Decades after Driesch’s original experiments, the mysteries of
development still supported vitalism, according to Rignano.
Those arguments may have stymied mechanists in the past, but Needham
remained unfazed. He insisted that embryological examples no longer posed an
insurmountable problem to mechanists: When one adopted mechanistic
explanations, one was not putting forth an all-or-nothing, verify-or-falsify
scientific hypothesis. Mechanism was not a conjecture to be refuted. Rather,
mechanism was part of a comprehensive worldview. Future vitalists might
discover this or that biological system for which mechanists could not at present
provide an explanation. But this was a temporary setback; a mechanistic
explanation would be forthcoming. Mechanists could be patient, flexible. Case
in point: Julian Huxley (1887–1975)—grandson of “Darwin’s Bulldog,” T. H.
Huxley and friend of Needham’s—was just then working on thyroxin, a
biochemical that certainly appeared to initiate the “genuine morphogenous
action” Rignano chided the mechanists for missing. “In chains of causes like
[development],” Needham pronounced confidently, “the exact biologists are
working out the complex events in living matter.” He was confident some
fittingly mechanistic explanation would be found.
Then Needham pressed his attack against the core of vitalism. Revisiting a
charge leveled by T. H. Huxley himself two generations earlier, Needham
reminded his audience that the vitalists’ “semi-philosophical arguments will not
deter [scientists sympathetic to mechanism] nor put the clock back by a single
minute. . . . A hypothesis is not regarded as respectable in scientific circles
unless it is a working hypothesis; yet Sig. Rignano never devises any
experiments which could prove or disprove his views.” 4 It was the classic
criticism of vitalism: What actual theory could be tested, what conjecture could
be verified? Wasn’t Rignano writing mere philosophy ? That was the decisive
blow against Rignano’s teleology. It wasn’t science.
Without greater context, Needham’s Man a Machine feels typical—a young
intellectual pugilist trained in one of the most prestigious chemistry labs in
1920s Britain writes in bemused, slightly annoyed terms defending his turf
against an interloper from outside his discipline. Given more historical context,
however, Joseph Needham seems a particularly unlikely candidate to write such
a book. In a short period of time Needham transitioned—one might say
converted—from defender of mechanism to the next generation’s leader of
organicist biology. His story reflects the greater narrative of a new generation on
the hunt for an alternative to mechanism and vitalism.

From birth, Noel Joseph Terence Montgomery Needham was destined to be a

transgressive, an in-betweener, one who straddles disparate realms. The only
child of, as he put it, a “marriage of Fire and Water”—his father an emotionally
reserved physician, his mother an outspoken composer—he spent his entire life
attempting to find compromises between seeming opposites: mechanism and
vitalism, Christianity and natural science, East and West. 5 By all accounts, he
was successful at this pursuit. By his death in 1995, Joseph Needham had
become the only man in British history to be elected a Fellow of the Royal
Society, a Fellow of the British Academy, and a Companion of Honour—an
achievement only possible for someone passionate and talented across the
intellectual spectrum. (When Dorothy Moyle Needham, his first wife, was
elected FRS, they became the first couple to be so honored.) 6
This “bridge building” work remains apparent in the issue Needham is best
known for today: what scholars continue to call the “Needham Question.” The
Needham Question begins with another question. What do the following have in
common: movable wood-block printing, canal locks, the wheelbarrow, chain
suspension bridges, fireworks, kites, and the magnetic compass? The answer is,
they were all invented in China several centuries before they appeared in
Europe, the Near East, or North Africa. Chinese technological inventions and
scientific discoveries antedated many of those associated with famous Europeans
such as Newton, Galileo, Gutenberg, and Da Vinci. As Needham began to
discover the considerable extent to which Europeans borrowed, rather than
invented, the techno-scientific discoveries that comprised the core knowledge of
Western civilization, he was caught by that second-level wonder that is
sometimes expressed by experts who learn that concepts they previously knew as
solid facts were false all along. Thus Needham’s Question became, “Why did
modern science appear first in western Europe, followed by the Industrial
Revolution, when it seems that China was historically the country most likely to
modernize science and revolutionize industry?”
In the twenty-first century, we might take issue with the question phrased that
way. We would, of course, want to ask, “Well, what do you mean by ‘modern
science’?” But that question sidesteps the real issue. The real issue is that
Needham cared to ask the question at all. It was his recognition of relative parity
between Chinese and European culture and the recognition that for centuries
Chinese technology was far superior to European technology that drove the
project that consumed half of Needham’s life: the series Science and Civilization
in China . Thirty-six volumes have been published at present; his passion
continues, now decades after his death, at the Needham Research Institute in
Cambridge. Indeed, it is that passion that author Simon Winchester highlighted
in The Man Who Loved China (2008).
Needham’s path from biochemistry to the history of Chinese science and
technology will become clearer in a later chapter. For now, it is enough to note
something that Winchester deemphasized: Needham’s remarkable ability to cross
cultural lines began long before his acquaintance with Chinese intellectual
culture. In truth, his passion for the history of Chinese science and technology
emerged out of several other foundational factors, including Needham’s blend of
religion, politics, biology, and philosophy. In the religious and political sphere,
for instance, Needham infused the Anglican Christianity of his upbringing with a
radical pursuit of the kind of social and political wholeness he saw both in Jesus
of Nazareth’s Sermon on the Mount and in twentieth-century British socialism. It
would have been natural for a person of his prodigious intellect and charisma
married to those particular political and religious convictions to channel his
drive into leadership in church or state. He did briefly help Julian Huxley birth
UNESCO in the 1940s. But rather than becoming a career-long clerical reformer
or political demagogue, Needham fashioned his religious-political motivation
into a chisel with which he would carve a channel for the organic philosophy.
His teachers at Oundle in Northamptonshire—one of the oldest public
schools in England—found Needham shy and thirsty of mind, and they assumed
he would go on in medicine like his father. 7 After graduating, he departed for
King’s College, London, to do just that, not least because the Great War was at
its bloody peak and medicine seemed a practical pursuit. He received his first
“real world” medical training on school breaks. Needham’s father pulled young
Joseph along with him into the operating theaters at the London military
hospitals. Joseph witnessed firsthand the effects of Fritz Haber’s chemical
weapons deployed on the battlefield, not to mention the ability of high-explosive
shells to maul a body. The experience was harrowing enough to scare him away
from a career as a surgeon.
Fear of blood and gore alone could not dissuade Needham from entering
Gonville and Caius College, Cambridge University, at the close of the war,
despite the fact it had been the home of legendary medical men including
Francis Glisson and William Harvey and continued to feature medicine as a core
subject in the twentieth century. At Caius, his first tutor, Sir William Bate Hardy
—perhaps sensing Needham’s ambivalence toward hands-on medicine—
redirected his serious, almost clerical pupil toward biochemistry. One could
harness biochemistry to heal and still serve society without going near the
surgeon’s knife. Indeed, Needham found biochemistry discipline bending,
refreshing, and a lot less bloody. It was a much better fit. By 1921, he read for
his BA in chemistry and anatomy-physiology. Also that year, he took his first
short research trip to work in a German biochemical lab. 8
These were decisive steps. Upon his return to Cambridge, he latched onto
Cambridge’s legendary biochemist Frederick Gowland Hopkins. Known as
“Hoppy” to his friends and colleagues, Hopkins made a name for himself as the
founder of biochemistry at Cambridge. His work with tryptophan resulted in the
1929 Nobel Prize in Physiology or Medicine. Moreover, Hopkins’s reputation as
a mentor was as legendary as his biochemical prowess. It was widely rumored
that “all of Hoppy’s geese turned into swans.” In that nurturing environment,
Needham flourished for two decades, earning a master’s degree and then
defending a doctorate later in the same year, 1925. When focused on his work,
his output was prodigious, peerless. Hopkins was so impressed he soon made the
young Needham his second-in-command.
Not one for straightforward problems, Hopkins agreed with his protégé that
the most important puzzle to solve would be the labyrinthine biochemical paths
traversed by chicks as they develop in their eggs. Biochemically, development
was a veritable Gordian knot, but Needham made surprising headway through
the 1920s. He happened upon a 1914 German dissertation by H. A. Klein on
changing levels of inositol in developing chickens; it was a key to the chemistry
of ontogenesis. That experimental work resulted in several papers, but they were
merely the birth pangs of a massive study on organismal development he nursed
and sculpted for several years. He published it as a three-volume tome entitled
Chemical Embryology in 1931; it was a monumental achievement, something
that emerges near the end of a distinguished career. 9 Needham, however, was 30
years old. With the cockiness characteristic of talented youth, Needham expected
that Chemical Embryology would initiate a “new branch” of science—which it
promptly failed to do. Its publication did accomplish something else, though. It
brought Needham into contact with two very disparate and very influential
scientific communities: cadres of distinguished German embryologists and
Bolshevik theoretical biologists and physicists. In order to understand why these
two groups were so significant for Joseph Needham’s story—as well as for the
story of the “third way”—we need to understand better the other area in which
Needham became a cross-cultural bridge builder of some renown: his religious
and political affiliations.
Given his pro-mechanistic attack on Rignano’s vitalistic Man Not a Machine ,
it might be difficult to envision Needham as a man who took his religion
seriously. This is understandable. Typically, we regard the mechanistic
philosophy of Jacques Loeb and many others as aggressively atheistic; the
kinder, gentler road of vitalism is supposed to comport better with a religious
outlook. Books like Rignano’s certainly play off this supposition. For Needham,
the reverse was true. Mechanism seemed to place a defensible barricade around
the spiritual realm, leaving a safe space for the flourishing of religious ideals.
(As Needham understood it, this had been Immanuel Kant’s strategy in Critique
of Judgment .)
Yet Needham’s Christianity was anything but traditional from the start. As a
child, his father took him to hear the Master of the Temple in London, E. W.
Barnes, FRS, later known as the radical Bishop of Birmingham. Barnes’s
sermons famously referenced ancient and medieval philosophy far from any
Christian canon. Following these sermons, Joseph would travel home, head
brimming with ideas, and dive straight into his father’s library, which was full of
classical texts. Needham admitted later in life that it was indeed Bishop Barnes’s
philosophy that triggered the pursuit of his own mixture of intellectually and
sociopolitically engaged Christianity. As a graduate student in the early 1920s,
Needham became an Anglican lay brother of the Oratory of the Good Shepherd,
even moving into the Oratory House in Cambridge for a time. Soon he began to
attend the Anglican Church at Thaxted, Essex, led by the controversial Rev. Fr.
Conrad Noel. It was another significant move for Needham that would ripple
through his scientific work.
In the first decade of the twentieth century, Conrad Noel raised the hackles of
churchmen and politicians throughout the UK by displaying at the front of his
sanctuary the banner of the international socialist movement. Both the flag of
Sinn Fein and the traditional flag of St. George scandalously flanked that
unadorned scarlet flag of socialism. Locals nicknamed Noel “the Red Vicar,” an
epithet he bore proudly during the 1911 Social Democratic Federation meeting.
There, he helped start the British Socialist Party. Over the next three decades,
Noel worked ceaselessly to align the aims of the British Socialist Party and the
Church Socialist League (founded in 1906). He tirelessly advocated the ideal
that “the community shall own the land and industrial capital collectively and
use them cooperatively for the good of all.” 10
Noel grounded his publications and even his sermons in the 1889 Fabian
Essays in Socialism . His Christian Fabian socialism deemphasized violent
revolution in favor of political change from within. He crossed traditional ethnic
boundaries by explicitly seeking out partners among leaders in the Roman
Catholic and Presbyterian churches; he invited Irish, Welsh, and Scottish as well
as English to be a part of his congregation and movement. Still more radically,
Noel endeavored to draw manual laborers and their families together under one
steeple with middle-class academics and students. Indeed, Noel’s message
effectively mandated an open, holistic philosophical anthropology. Ideally, his
Christian socialist community would exemplify a cross-cultural unity formed by
mutually interdependent members. His community took practical steps to make
these ideals a reality. Members of the “Thaxted movement,” as the community
that surrounded Noel came to be called, held property and possessions in
common. They saw this practice of holistic mutual dependence, in fact, as the
way to enflesh Christ’s Sermon on the Mount in modern Britain.
That reveals another important feature of the Thaxted movement that
separates it from earlier iterations of Christian socialism: its preoccupation with
primitivism. “Primitivism” can be understood in multiple ways. It was a
motivating factor behind their mutual dependence notion, which harkened back
to the early Christian Church in Jerusalem. But the most obvious expression of
primitivism in the Thaxted movement was the revival of Morris dancing. Morris
is a type of English folk dance dating to the late medieval period. At Thaxted it
was often claimed that the dancing style was pre-Roman, but morris probably is
a derivation of “Moorish.” 11
Joseph Needham pulled fellow biochemist Dorothy Moyle into the Thaxted
movement in the early 1920s. They married in 1924, after Dorothy received her
MA and found a long-term place in Hopkins’s lab. Together the Needhams
invested ever more deeply in both biochemistry and the Thaxted movement.
Joseph even learned Morris dancing, becoming proficient enough to start a
Morris Dancing Ring and make it an established feature of Cambridge
University life by 1934. The new couple agreed that both of them should
continue laboratory work even if that precluded having their own children.
Dorothy remained very private but encouraged Joseph to step into the limelight,
to become a recognized bridge builder even if that also meant he would ruffle
disciplinary feathers. So, with Dorothy’s blessing, Joseph continued publishing
ever more confrontational works exploring the intersection of faith, economics,
politics and science.

Man a Machine , his challenge to Rignano and vitalism, was but one foray
into this region where the boundaries of religion, politics, and philosophy of
science overlapped. Even in the 1920s, it was a kind of no-man’s-land for a
bench scientist like himself. But Cambridge was a special, perhaps even
insulated, place. Needham sharpened his thoughts and debating skills at informal
Cambridge club meetings. There he became known as being so provocative,
erudite, and clear spoken that other groups invited him to give lectures on the
subject to wider audiences. He was made a leading speaker at the General
Conference of the Student Christian Movement held at Swanwick, Derbyshire,
in 1931. Aside from Man a Machine , his popular lectures crystalized into
substantial series of essays, published as “Mechanism” (1926), The Sceptical
Biologist (1930), and The Great Amphibium (1932). 12
Somehow, Joseph managed to write the titanic Chemical Embryology (three
volumes, 1931) while he lectured and scripted these essays about religion,
politics, and the philosophy of science. As he was conducting research for that
project in the late 1920s, he encountered the older debates between mechanists
and vitalists. They intrigued him—so much so that he positioned the debate as a
historical prelude to his biochemical treatise. Soon the intended “prelude”
ballooned into an enormous, almost unwieldy project of its own, long enough to
reprint in 1934 as a stand-alone volume and popular enough to warrant a second
edition in 1959. Needham discovered that, though mechanism-vitalism took
different guises in earlier eras and though it rarely approached the heat generated
during the first decades of the twentieth century, the debate had ebbed and
flowed for centuries. Just the fact of the debate’s persistence bothered Needham:
It suggested that there was no progress in biological theory. Life scientists
continually rehashed the same meta-theoretical debates, without regard to
advances in instrumentation or experimentation. He firmly believed that progress
in theory was possible, even if theory lagged far behind what new instruments
revealed in the laboratory.
In a 1928 article entitled “Recent Developments in the Philosophy of
Biology,” which he published in the high-profile Quarterly Review of Biology ,
Needham intended to map the topography of progress in theoretical biology. He
aimed to convince skeptics—maybe himself above all—that even long-standing
debates in biology did progress toward resolution. 13 Ultimately, Needham hoped
to show, mechanism would win. In “Recent Developments,” he identified three
“schools” that currently stood in the way of that victory. Most notorious were the
“neo-vitalists,” who leaned upon “special entities immanent in living things” to
explain the persistence of bodily form evident, for example, in salamanders’
ability to regenerate limbs. Hans Driesch and Henri Bergson, of course, played
the neo-vitalist villains in Needham’s story. As Needham saw it, any points these
neo-vitalists accrued due to their popularity early in the century had slipped
away by the late 1920s. The reason was simple: Science could not put concepts
of entelechy or élan vital to use. They yielded no analyzable results.
Less vocal, though no less important, the “finalist school” recently
highlighted by Eugenio Rignano’s Man Not a Machine also purported to defang
mechanism. But earlier attacks on mechanism by scientists appealing to
teleology had already lost some of their sting by the mid-1920s. For example,
Rignano, and others like him, deployed thermoregulation as an unsolvable
problem and, therefore, fatal to mechanistic explanations. Needham assured his
readers that this was not, in fact, a problem. Already in 1924, Hartridge and
Roughton had developed a device to measure rapid chemical reactions. 14
Roughton believed that this would help put to rest appeals to teleological
explanations of metabolism, which would certainly hurt the finalists’ arguments.
15 Needham conceded that, in one sense, the teleological camp of Rignano did

useful work: It highlighted just how intricate the biochemical balancing act
organisms had to sustain could be. Yet that complexity didn’t pose an insoluble
problem for mechanism the way the finalist school believed it did.
Thermoregulation would be solved—mechanistically—sooner rather than later.
And Needham exposed a more serious flaw in the finalists’ arguments. Nonliving
substances could do a biochemical regulatory dance of their own, albeit a much
less complex version. For Needham, this fact meant that the apparently
purposeful regulation of their internal states that was done by even very
primitive organisms should not be seen as something spooky, and certainly not
as a reason for life to be regarded differently than nonlife.
Last, Needham turned his critical gaze on “organicists”—he looked squarely
at the first generation of Haldane, Lloyd Morgan, Whitman, Russell, and others
here. Like the neo-vitalists who followed Driesch and Bergson, Needham
insisted the first generation of organicists believed that the organism stood
outside the bounds of ordinary physics and chemistry. Living individuals, even
single-celled amoeba, were just too complex to be explained by the ordinary
mechanistic laws, they said. Unlike neo-vitalists, these early organicists believed
perhaps that living things could be understood by science, as long as scientists
discovered new laws—laws that described the activities of wholes “obstinate and
irreducible.” 16 On the one hand, organicists might not need to appeal to
entelechies, vital fluids, or forces. Needham learned this by sparring privately
with C. Lloyd Morgan. Lloyd Morgan assured Needham that his philosophy

seems to me to be neither mechanism (in that it is impure) nor vitalism (in that it eschews any form
of psychoid business). But I suppose that it is because you think it ought to be called vitalism that
you class me among the neo-vitalists. I’m afraid they won’t own me. I am kicked out from both
camps and get digs in the ribs from both sides. 17

On the other hand, Lloyd Morgan’s philosophy did draw upon the Aristotelian
concept of wholes surpassing the sums of their constituent parts. This view, after
all, was just common sense for a committed empiricist, Lloyd Morgan insisted.
Living organisms really were more complex when in one piece and alive than
they were when chopped up and mounted under the lens of a laboratory
microscope. If certain scientists insisted that biology could be reduced to
physicochemical terms, it was only because those scientists had an obdurate
mechanistic axe to grind. They could not be empiricists; they had not been
baptized in the biology of the whole organism.
This, in any case, was Joseph Needham’s depiction of the older generation’s
organic philosophy in his “Recent Developments.” Surprisingly, Needham found
young, active biologists defending this old deficient organicism alongside the
aging first generation. Here Needham underlined the name of Joseph H.
Woodger. Though he didn’t know it at the time, Needham’s mention of Woodger
began a relationship that would change the future of “third way” biology.
Needham did not intend for “Recent Developments” to be merely a list of
targets; he did hope to offer an alternative to anti-mechanism that he feared had
been gaining popularity. He dubbed the alternative “neo-mechanism.” Needham
explicated neo-mechanism fairly simply: The typical mechanistic explanation for
phenomena would work perfectly well in science, but it had to stay confined
within the territory traditionally occupied by science. Unlike Jacques Loeb’s
version of mechanism, for example, Needham insisted his neo-mechanism could
not be applied to other human pursuits. Art, for instance, deserved its own
territory; so did religion. Needham intended neo-mechanism simply as a way to
define boundaries. Was an investigator working on the problem of organismal
generation and development? They should explain it in the most mechanistic
way possible without fear that they would have to contend with discussions of
unobservable entelechies or élans vital. There could be no science-stopping
boundaries in nature. Anything must be open to mechanistic analysis in
principle. Was an investigator worried about the meaning of life or the ultimate
existence of the universe? Mechanistic science could be of little use there; this
was the domain of theologians, philosophers, and mystics. Needham felt certain
that “Neo-mechanism gives biological science all it wants.” Moreover, neo-
mechanism shielded scientists from criticism by those nonscientists who
regarded the intrusion of mechanistic explanations as the first wave of scientific
imperialism. 18 Neo-mechanism would remain methodological mechanism.
Metaphysics would be left to philosophers.
As he reconsidered it much later, Needham’s attempt to advocate for
methodological mechanism came about because he did not believe the disparate
worlds of politics, religion, and science truly could be synthesized. 19 Needham’s
version of neo-mechanism was Kantian disintegration, a way to keep the worlds
of science and religion apart lest science actually deplete the qualitative richness
from those human endeavors that resist easy quantification like art and religion.
Perhaps not coincidentally given their friendly correspondence many years
later, paleontologist Stephen Jay Gould would echo in the 1990s Needham’s
demarcation between quantitative and qualitative human activities. Gould
described the coexistence of science and religion as one of non-overlapping
magisteria (to which he assigned the catchy acronym “NOMA” in 1997). “The
net of science covers the empirical universe: what is it made of (fact) and why
does it work this way (theory). The net of religion extends over questions of
moral meaning and value. These two magisteria do not overlap, nor do they
encompass all inquiry (consider, for starters, the magisterium of art and the
meaning of beauty). To cite the arch cliches, we get the age of rocks, and
religion retains the rock of ages; we study how the heavens go, and they
determine how to go to heaven.” 20 Dominate your sphere of influence, Gould
stipulated. Dictate the way information is distributed, what counts as
authoritative, and how to call out enemy infiltrators. But stay within your sphere.
Leaky spheres create conflict. Gould came through the ringer of creation science
court battles of the 1980s. Needham’s concerns and motivations were shaped by
the political, social, and economic upheavals a half century earlier. Nevertheless,
Needham and Gould ended up with the same initial proscription—leave religious
questions to religion and scientific questions to science. They recognized that
religion and science approached each other like blocks of wood with grains
running perpendicularly to one another. Rather than simply lenses for
interpreting the same data, competing paradigms, science and religion asked
different questions and heard different answers. As Needham put it, “Science . . .
is abstract, dealing with statistics and avoiding the individual . . . quantitative,
metrical, mathematical and deterministic. It is essentially classificatory,
mechanical, analytical, and orderly, generalising and impersonal. . . . It fights
mystery and teleology. . . . Religion, on the other hand, is concrete and
individual, based on the sense of the holy, with which are connected the
sentiments of reverence and awe. It is qualitative in feeling, opposed to
measurement and analysis, ‘cornucopial’ instead of orderly, personal . . .
essentially irrational and alogical. . . . It naturally insists on ‘free-will’ as against
determinism.” 21

Despite the strength of his endorsement in the 1920s and its utility in separating
spheres of science from religion, Needham largely abandoned neo-mechanism
by the end of the 1930s. His perspective changed as a result of three factors:
wrangling with the “third way” philosophy; his work on Spemann’s organizer
conducted alongside his wife, Dorothy Needham, and embryologist C. H.
Waddington; and the influence of the Soviet delegation to the Second
International Congress of Science and Technology headed by Nikolai I.
Bukharin. All of these events took place between the publication of “Recent
Developments in the Philosophy of Biology” in 1928 and his book Order and
Life in the mid-1930s. The first and most important step in this transformation,
however, was due to the eloquent persistence of J. H. Woodger.
 3

SOCRATES AND THE PRINCIPIA BIOLOGÆ

J . H. “Socrates” Woodger (1894–1981)—the man Needham accused of being a

young representative of discredited “older organicism” in his 1928 “Recent
Developments” article—did not begin his career as an opponent of mechanism.
In truth, as a young man he was much less likely than Joseph Needham to dabble
in these sorts of theoretical questions at all. When he entered University College
London (UCL) in 1911, he cared little for philosophical issues, or even for
experimental biology. Like Needham, he hoped to be a physician; unlike
Needham, blood never bothered him. Medicine proved to be a natural fit.
Woodger earned both the College Prize in Zoology and the Derby Research
Scholarship while a medical student at UCL, showing a natural aptitude for
experimental research. But once the drums of the Great War began beating in
1914, he joined many of his peers and enlisted in the British Army. By 1915, he
would be assigned to the 2nd Battalion, Norfolk Regiment (regulars). Zoological
research would have to wait.
The Great War touched the young Woodger more directly than Joseph
Needham. When the Ottomans joined the Central Powers against France,
Britain, and Russia, the Royal Navy diverted thousands of young men from the
French front to Basra at the southern edge of the Ottoman Empire. At the start of
the Mesopotamian campaign in the autumn of 1914, British command hoped a
show of force would simultaneously knock the Turks back into neutrality and
expand British influence in the oil-rich Persian Gulf. The 2nd Norfolk landed in
Iraq on 15 November 1914 and immediately saw action. 1 Barely a month later,
British forces had secured Basra and the southern Iraqi oil fields. With easy
victories against the Ottomans, British command ordered an advance to the real
prize in the region, Baghdad, over five hundred miles upriver. This time, the
advance was a slog. Roads were few and far between. Disease dogged the
British invaders, eventually stalling the advance.
A second, also costly attempt at Baghdad began two years later in February
1917. By this point, Woodger was there, part of the Royal Army Medical Corps.
During the march north, he fell in alongside Ian Dishart Suttie (1889–1935), the
son of a Scottish physician who was quite capable of “talking shop.” They talked
for days on end; the talking solidified into lasting friendship; the friendship
continued for two decades until Suttie’s untimely death. As Medical Corps, they
experienced more intensely than most the gradual unraveling of the British
campaign: the aftermath of the disastrous Ottoman siege of Kut-al-Amara, Iraq’s
alternating floods and baking hot seasons, and ever-present pestilence that felled
at least as many British soldiers as Ottoman bullets did. 2 In the Mesopotamian
theater of the Great War, disease was as nightmarish as chemical warfare was in
France and Belgium. Waves of cholera, dysentery, and various fevers swept over
the British camps without shedding any light on how best to stop them from
returning. Woodger attempted to put his laboratory training to use combating
disease in the hastily constructed British Central Laboratory in Amara, southeast
of Baghdad. Among other topics, he focused on the spread of plague in rat
populations, the possible insect vectors of amoebic dysentery, and—the bane of
marching infantry—hookworm. 3 Suttie moved into the psychiatric division and
became part of the nascent counseling effort treating mounting numbers of shell-
shocked and permanently disfigured soldiers.
When the 1918 Armistice finally arrived, Woodger, like so many others,
returned home a changed man. He now saw himself as an experimental biologist
far more than as a physician, and he took a position as assistant in Zoology and
Comparative Anatomy under the direction of J. P. Hill back at UCL. There,
Woodger used his war-sharpened microscopical skills in tracing the development
of Golgi bodies during egg formation. 4 Four years later, in 1922, Middlesex
Hospital Medical School in London offered Woodger a Readership in Biology.
He leapt at the chance to teach, but he was soon disappointed to learn that his
students were even more practically minded than he had been before the war.
They had little interest in basic research in the lab, especially as their craft did
not require it. Woodger set out to impress upon medical students the importance
of modern biological knowledge to their field—even if they did not like it.
Within two years he had published a textbook, Elementary Morphology and
Physiology for Medical Students: A Guide for the First Year and a Stepping-
Stone for the Second . Though it dealt with many of the conventional topics one
would expect in a book of this kind, Woodger also explored large philosophical
questions. He revealed his motivations for slipping philosophy into medical texts
to Ian Suttie alone.
Suttie remained Woodger’s wisest, most sympathetic sounding board. With a
medical degree from the University of Glasgow, Suttie rose to prominence
almost immediately upon his return. Only two years after the Armistice, the
Royal Faculty of Physicians and Surgeons made him a fellow. But the
recognition felt hollow. Suttie left working on the body soon after and moved
into the realm of mental health exclusively, receiving appointments at several
hospitals, including the Glasgow Royal Asylum. 5
The two continued their friendship through correspondence and holiday
excursions in Scotland, undoubtedly revisiting war stories. 6 Understandably,
both were excited when, later in the decade, the Tavistock Clinic in London
hired Suttie as a psychiatrist. He remained there after it was reconfigured as the
Tavistock Institute of Medical Psychology and relocated to the Bloomsbury
neighborhood, not too far from Woodger. But the position required so much
more work that, paradoxically, the two seem to have seen less of each other as
the years went by.
It was the excessive workload at Tavistock and his deep attachment to his
patients that killed Suttie, according to his colleagues. Periodically he had
complained of undefined abdominal pains. These had even plagued him during
the war itself. 7 Nevertheless, Suttie had been able to manage both “active
service conditions on the Mesopot[amian] front—1/2 share of 40 lb. tent etc.,”
and subsequent stress from pioneering the field of psychological social work. 8 In
fact, he seemed to need very little relaxation, always keeping up his productive
schedule. By the mid-1930s, however, the pace was proving overwhelming. In
addition to clinical work, Suttie had students to manage: Tavistock began to offer
postgraduate courses and clinical practicum to UCL and London School of
Economics students. Suttie’s bouts of illness were finally diagnosed as a
duodenal ulcer. Occasionally it hemorrhaged, producing severe pain and copious
bleeding. In the fall of 1935, despite the fact that Suttie had attempted repeatedly
to reduce his workload, it erupted catastrophically, perforating his intestine. He
died tragically on 23 October, mere days before the appearance of his
groundbreaking On the Origins of Love and Hate , an attack on interpretations of
Freud and Adler that deemphasized development and interdependent
relationships. 9
Though disconnected from the mainstream psychoanalytical community of
the mid-1930s, Suttie rooted Origins in his conversations with Woodger about
biology and philosophy just after the war. Through the 1920s, they corresponded
often about scientific epistemology and the justification, or lack thereof, for
various claims made ex cathedra by prominent scientists. Suttie, especially,
faulted scientists who made pronouncements in areas outside of their particular
research expertise. Better acquaintance with philosophy of science, he argued,
would serve to discourage this behavior—what he called intellectual “poaching”:

Philosophy then insists in asking questions; science in answering them & concerns itself with
“extending” and “articulating” sense-experience while P[hilosophy] aims merely at “articulation.” .
. . [T]he only thing that remains permanent about [philosophy] is its intolerance of confusions and
antimonies with the insistence that the whole of experience be taken into account. It is indeed easy,
as some “isolated” sciences practice it, [to] confine oneself to a small enough section of experience,
disregarding the rest, to build a conceptual system which will be logically coherent and will
adequately represent the “facts.” But the initial definition of the section of experience which will
constitute the field of any particular inductive science is arbitrary, allowing closely related
phenomena to be represented by distinct and even incompatible systems of knowledge. Worse even
as these systems “succeed” they tend to widen their empirical bases to poach on their neighbors’
territory; so it may come about that the same element of experience may be interpreted or
formulated in two totally different ways and conventions. [ . . . ] The mind whose interests are
“focused” (I won’t even say narrow) does not notice these contradictions and divergences in our
formulation of experience. 10

But only a scientist who recognized the importance of robust philosophy could
lead the reform he was talking about. Suttie implored the philosophically minded
scientist (he definitely pointed his finger at Woodger here) to expose the “felt
need for harmony of interest and systematic continuity of thought” across the
breadth of human experience: “The aim of such must be to bring home to the
individual a sense of his own intellectual confusion: something analogous to the
‘conviction of sin[,]’ which is the first step in religion.” 11
This charge from Suttie sparked something profound in Woodger. Though
drawn to philosophical questions since the war, Woodger’s interest had been
diffuse. Suttie’s twofold admonition—to convict the scientist of his or her own
intellectual confusion, and to resolve the “antimonies” so that scientific fields
could better communicate without “poaching” from one another—sharpened
Woodger’s vision of how philosophy could intersect with his own scientific
work. Articulating that vision would occupy Woodger for the rest of his life. It
would drive him to confront, and eventually convert, Joseph Needham, inspire
embryologist C. H. Waddington, introduce Peter Medawar to the philosophy of
Alfred North Whitehead, wrangle with philosopher Karl Popper, and congregate
the most lasting dedicated “third way” group—the Theoretical Biology Club.
All of these things were in the future, however. First Woodger promoted this
vision in Elementary Morphology and Physiology for Medical Students . In
chapters covering scientific methodology, theoretical biology, even the mind-
body problem, he stressed that different areas within biology rely on different
explanatory frameworks, largely implicit, unrecognized by practitioners. The
imprecision in the language of biology, compared to physics, for instance, meant
that those implicit frameworks ended up influencing areas for which they were
not intended. Case in point, Woodger believed, was the framework adopted by
geneticists:

It is easy to understand why so much attention has been fixed on the chromosomes. The fact of
their careful partitioning between the daughter cells in division has always been suggestive of their
great importance in the normal life of the cell. . . . It cannot be concluded, however, that the
cytoplasm is of no importance in development , whatever may be its importance for genetics. . . .
Mendelism only deals with the distribution of differences between life-histories which are not
dependent upon differences between their zygotes in respect of their parts which are not duplicated
at each cell-division. But such parts are of primary importance in early development because the
establishment of differences between the cells of a life-history must ultimately be traceable to them,
if we assume that the chromosomes are equally divided in mitotic division. This point is in danger
of being overlooked when we try to bring the Mendelian theory into relation with embryological
data, for Mendelism is a genetical and not an embryological, theory. 12

Geneticists were moving into a territory where they did not belong—“poaching,”
as Suttie had put it. But as Woodger undoubtedly understood, most of his
colleagues would not agree that a biological theory could be restricted to a
particular realm of biology and not extended to cover all biological phenomena.
He would have to demonstrate to biologists precisely why genetic theories
should be restricted to genetics, leaving embryologists to pursue their own
concepts. In order to do that, Woodger would need help.
In 1926, Middlesex Medical School granted Woodger leave to visit Vienna’s
Biologische Versuchsanstalt (Biological Research Institute), headed by the
spirited Hans Przibram. Popularly known as the “Vivarium”—a name that
recalled its role as an aquarium and micro-zoo in the nineteenth century—
Vienna’s Biological Research Institute attracted experimentalists from across
Europe. It developed a reputation for promoting mathematical precision in the
biological laboratory and, paradoxically, using the mathematics to undermine
mechanistic interpretations. 13 While working at the Vivarium, Woodger
discussed theoretical concepts with Przibram himself and made contact with the
philosophers in the Ernst Mach Society, one of the legendary Vienna Circles.
Upon his return to Britain, Woodger channeled the conversations with
Viennese biologists and philosophers into a doctoral thesis. He derived
motivation for the project not only from his Vienna trip but also from his
perpetual frustration with “poaching.” Perhaps if he could bring the tacit
assumptions held by his peers in genetics and embryology to light, they could
agree to the boundaries between their subfields. So Woodger attempted to ferret
out the principles of biology: “No one,” Woodger explained, “ . . . had
undertaken a systematic critical study of the fundamental properties and special
requirements of this science in relation to the most advanced metaphysical,
epistemological and logical notions of the day.” 14 He called it only “tentative,”
but the result was an imposing and rigorous piece of analysis—certainly enough
to merit his DSc from UCL. 15 On the way toward publishing the dissertation,
Woodger managed to extract two articles out of his research. 16 It was through
the first of these essays that Joseph Needham learned of Woodger.

In “Recent Developments in the Philosophy of Biology,” Needham had

classified Woodger’s version of organicism as little more than a protracted
retreat in the face of the clearly superior force of mechanism. Probably without
knowing it, Needham repeated the sentiments of Walter T. Marvin at the APA
meeting after the First World War: Like vitalists, organicists were hopelessly
outmoded, standing in the way of scientific progress. After reading Needham’s
essay, Woodger fired back on 28 May 1928, defending the organic philosophy:
“I fully agree that the strong point of ‘mechanism’ is on the heuristic side and
that none of the other alternatives have yet offered anything in that direction
which is fit to hold a candle to it. But I deny that the ‘organic’ view has been
wholly devoid of heuristic merit, and when you consider how few people hold it
compared with the opposite view it is hardly surprising that it has very little to
show for itself in this direction.” 17 Even during the 1920s groundswell of anti-
mechanism there were too few investigators giving organicism a fair hearing,
thought Woodger. And no wonder. If they would face professional tarring and
feathering as Drieschian vitalists or, maybe worse, mere philosophers by those
who, like Needham, retained secure academic positions in laboratories across the
UK, how could they attain the occupational security necessary to defend a “third
way”? Woodger lashed out. There was no in principle reason why biological
work “should not be guided by the organic point of view as well as by any
other.” 18 Needham did not reply to Woodger’s letter, perhaps because at the time
he was publishing Man a Machine and writing Chemical Embryology . So
Woodger continued working on his dissertation without a response from
Needham. When completed, he submitted it to his DSc Committee at UCL and,
simultaneously to K. Paul, Trench, Trubner & Co., who published it as
Biological Principles (1929).
Biological Principles should still be read as the cornerstone theoretical text of
the “third way.” It remains among the most intensive attempts to resolve
theoretical disputes that scientists have passed by for centuries. The fact that it
was written by a research biologist who found practical relevance in these
theoretical issues makes it that much more important. Though Ian Suttie may
already have recognized him as such, Biological Principles signaled to others at
the time that Woodger was a genuine, perhaps even a formidable, biologist and
philosopher. Reviewers reflected that Woodger might be in another league from
other biologists. For these reasons, it deserves a closer analysis here. 19
Woodger began the book with a provocative metaphor: Biology was a crab
that had outgrown its metaphorical shell—its ability to measure and assess had
outgrown its theoretical structure. Drawing from the periodization of Alfred
North Whitehead in Science and the Modern World (1925), Woodger delineated
two previous “shells” in biological thought. The first shell, Aristotelian
commonsense empiricism, dominated for almost two millennia until the advent
of a second shell in the seventeenth century, Rene Descartes’s mechanism, which
Woodger claimed continued through the nineteenth century. By the 1920s,
however, biology had joined the rest of the natural sciences in a “transition
period” instigated by the widely publicized revolutions in physics. 20 In those
first few years of the twentieth century, Mendelians like William Bateson joined
the well-known theoretical physicists—Einstein, Bohr, and Heisenberg chief
among them—in recognizing that the foundations of the science would have to
be torn up and re-laid in order to progress into a new phase. The problem was
that, by the interwar period, biologists lost interest in theory. (American biologist
Raymond Pearl would soon emphasize this point to Woodger, as we will shortly
see.) To Woodger, this meant that the life sciences sorely needed their own
“skeptical biologist” to spur theoretical change—a self-critical mirror of the
same design as Robert Boyle’s Sceptical Chymist (1661). Woodger hoped his
Biological Principles might trigger that prophetic resetting of the metaphysical
and epistemological foundations of his field.
Tellingly, he drew up his attack precisely the way Suttie exposited in their
correspondence a decade earlier. In part 1, Woodger focused attention on the
“sin” of intellectual confusion in the practice and theory creation of the scientist.
Part 2 contained his proposed resolutions to what Suttie had once called
“antimonies” and “incompatible systems.” Woodger selected six dyads that he
believed stood at the core of most debates in theology: mechanism versus
vitalism, structure versus function, organism versus environment, preformation
versus epigenesis, teleology versus causation, and mind versus body. Each
chapter carefully unknotted one of the long-standing dyads.
Woodger opened part 1 by spelling out the philosophy of phenomenalism,
which he saw as the unexamined philosophy most often associated with modern
science; he then proceeded to spotlight the weaknesses of that philosophy.
According to Woodger, Ernst Mach’s Analyse der empfindungen (Analysis of
Sensations ) and Karl Pearson’s Grammar of Science were unquestionably the
most prominent texts of phenomenalism, so he used them to demonstrate why
their philosophy led science in the wrong direction. 21 Woodger proclaimed that
Mach’s and Pearson’s versions of phenomenalistic science were shot through
with self-defeating propositions akin to many reductionistic, mechanistic
philosophies. For one, neither could give an account about how sensations enter
the consciousness without presupposing the sensing processes—that is, without
accepting an account of sensation that could not itself be sensed. For another, the
intellectual gymnastics required to render an anti-metaphysical account of the
world could be undertaken only if, paradoxically, it was supported by a
particular version of metaphysics. Woodger went to great lengths to show how
Machian-Pearsonian phenomenalism actually required the very metaphysical a
priori assumptions that it claimed to discard. In trying to replace dualist
metaphysics with materialistic monism, phenomenalism actually invoked
another metaphysics regarding perception itself. In the second chapter, Woodger
offered his alternative to phenomenalism, which was a kind of local,
circumscribed realism that blended thought with brute sensation and recognized
complexity in the natural world not as illusory and the result of accumulated
simples but as really there in a localized sense—an anti-reductionistic, anti-
mechanistic account.
Woodger opened part 2 of his landmark text by addressing the most imposing
of the tangled messes that had divided life scientists for over a century:
mechanism versus vitalism. Part of the conflict between these two explanations
had to do with the variety of views held by each side. Perhaps it would help
simply to categorize the main approaches.
Did a biologist regard organisms as mechanisms—albeit complex and wet
ones—and nothing more? She would be an ontological or metaphysical
mechanist. Or does our biologist merely require the heuristic of mechanism
without regard to what an organism actually is? In that case, she is a
methodological or provisional mechanist. A biologist from the opposing camp
might regard organisms as special in a sense that cannot be apprehended in any
laboratory; the organism exhibits traits that must ultimately have their
foundation in another process or entity somehow removed from the physical
realm. His vitalism, then, is of the metaphysical sort. Still other biologists avoid
offering any positive assessments of the status of the organism but nonetheless
attack the mechanistic explanation as insufficient. Into this group of diverse
“anti-mechanists,” Woodger placed John Scott Haldane among others. 22
Once he divided possible responses to mechanism and vitalism into these four
categories, Woodger believed the task of finding a resolution between
mechanism and vitalism—a fifth possible category—might prove easier. Only
two of those potential categories would be likely to come to any sort of
compromise: those committed to metaphysical mechanism and metaphysical
vitalism would be exceedingly unlikely to view any possible evidence
introduced by the other side as relevant at all. Those merely committed to a
methodological deployment of mechanism and their anti-mechanistic opponents,
by contrast, might be convinced to abandon their present positions once they had
seen a strict, logical dissection of their view. That is what Woodger intended to
provide.
Woodger delved first into the mechanistic explanations. In fact, most of
Biological Principles dwells on mechanism, since this was undoubtedly the go-
to explanation in the sciences. For an example of a mechanist, Woodger
examined none other than the American scion of cell theory E. B. Wilson.
Wilson openly endorsed mechanism as the only philosophy that “kept us moving
in the right direction.” But according to Woodger, Wilson’s commitment to
mechanism remained merely methodological. For instance, when speaking of the
apparent fine-grained orchestration of cellular movements, Wilson maintained
that the only acceptable means of explanation still “lies in the mechanistic
assumption that somehow the organization of the germ-cell must be traceable to
the physico-chemical properties of its component substances.” 23 Later in the
work, however, Wilson backed off the strength of his statement: “Existing
mechanistic interpretations of vital phenomena, evidently, are inadequate; but it
is equally clear, as someone has said, that they are a ‘necessary fiction.’
Knowledge will be advanced most surely by assuming that the problems of the
cell can be solved by converging upon them all our forces of observation and
experiment.” 24 Including Wilson’s use of this “necessary fiction,” Woodger
identified four distinct definitions that could be classified as methodological
mechanism: (1) interpreting biology in terms of classical mechanics, (2) any
explanation that relies exclusively on concepts in physics and chemistry (this
was likely Needham’s position in the 1920s), (3) the antiquated analogy that
organisms are wet machines (Wilson’s “necessary fiction”), and (4) a weak claim
that living things are “orderly,” i.e., they produce similar effects under similar
conditions. These definitions were all well and good, yet the definitions alone
could not solve the important problem of how far mechanistic explanations
should be pursued. Unfortunately, according to Woodger, “all our forces of
observation and experiment” alone could not decide this question.
This was hardly an endorsement of vitalism. Though later historians would
chide Woodger for failing to bow to the power of mechanism, Woodger’s
critique remained evenhanded throughout. 25 Drieschian entelechies and the like
Woodger dismissed summarily. Vitalists asserted too little about their concepts
of field and force, aside from invoking these “imperceptibles” whenever
mechanistic explanations ran up against some presently insoluble problem. 26
Closely allied, but not really more helpful than vitalists like Bergson and
Driesch, were the psychobiologists, once called hylozoists, who wanted to
extend notions of mind in some sense to all material. E. S. Russell occasionally
seemed to fit in this camp.
According to Woodger, there were three main reasons why vitalists and
associated groups continued to gain an audience within science and the educated
public; each objection to mechanism had its own group of followers, though of
course there was overlap between the groups. One camp criticized mechanistic
approaches for automatically discounting certain concepts—mind, for instance—
without giving solid reasons why these concepts should be ignored. A second
group insisted that secondary qualities of objects must have their ultimate origins
outside of the strictly material and must be supplied from another quarter (the
psychobiologists were of this ilk). The third set of objectors insisted that
mechanists simply gave up too early when it came to biology; they lazily
attempted to explain life in terms appropriate to nonlife but without
acknowledging that there could be a whole new set of physical laws out there
that were realized only in living things.
Woodger agreed with the vitalists that these were indeed difficult issues for
mechanists to address if they had to be faced all at once. Teased apart into three
distinct challenges, each with a backing group, Woodger found them less
problematic for mechanists. The first group—vitalists proper—wanted to expand
the pallet of explanations to include things outside the material universe. While
this might be metaphysically acceptable, Woodger did not see how vitalism
could benefit empirical science; this concern could not then be a scientific issue
resolvable by scientists. The second group, perhaps best called psychobiologists,
traced their intellectual lineage to respectable Enlightenment-era philosophers
like Baruch Spinoza, Johann Gottfried von Herder, and Johann Fichte. But,
again, Woodger could not see how having mind “baked in” to the material
system from the beginning was germane to the scientific practice of biology
even though it might very well be true. Even the third group—we might call
them the “new physical laws” group—failed to satisfy Woodger fully: He did not
discount their approach; he merely found it discouraging. If they were correct,
new answers to biological problems would have to wait until handed down from
someone outside of the discipline—from a physicist, probably.
Because he found each of the three anti-mechanistic positions dissatisfying,
Woodger looked instead for points of agreement between mechanism and
vitalism. And, surprisingly perhaps, he found one point in particular where they
overlapped. Both mechanists and vitalists tacitly assented to a view of the
universe popular most recently among nineteenth-century physicists. For them,
the universe could best be described as built from invisible, indivisible lumps of
stuff knocking about. Physics, the foundational science, described what these
lumps were and what happened as they bumped into one another. Ultimately,
according to Woodger, mechanists were those who insisted that this quantitative,
atomistic view of the universe could in the end account for all other phenomena.
Vitalists, though they came in many shapes and sizes, argued that other forces
from outside of the natural system were needed in order to grant the complexity
that living things exhibited. Very few on either side actually identified and then
rejected this underlying atomism; it was simply that vitalists believed it was
insufficient.
By the late 1920s, insisted Woodger, physics itself was beginning to abandon
this picture of fundamentally disorganized, discrete particles floating in empty
space in favor of organized fields. This point—that physics was no longer
atomistic and therefore biology should not be either—was a theme to which
most advocates of the “third way” would return through the 1930s. Woodger
found it disturbing that biologists—both mechanists and vitalists—had made so
little of this new perspective: “It is a curious fact that organization is being taken
more seriously in the physical sciences than in biology. . . . Biologists, in their
haste to become physicists have been . . . trying to treat the organism not as an
organism but as an aggregate. And in doing so they may have been good
chemists but they have not been good biologists, because they have been
abstracting from what is essential to the biological level.” 27 At the biological
level, Woodger reminded his readers, one never finds free-floating homogenous
chemical masses, as one would expect to encounter in chemistry, for instance. It
is true that biologists might colloquially use terms like “protoplasm,” as if cells
were filled with it as a kind of jelly; this was the term that so enchanted T. H.
Huxley in the nineteenth century. And biochemists might even fashion this sort
of gelatinous stuff in the laboratory. But nothing like a homogenous chemical
mass exists in a living state in nature: “If a large bomb is dropped upon a
populous town we might apply the term ‘town-plasm’ to the debris which
remained, but it would be a little absurd to say that towns were composed of
such town-plasm, and that from a sufficient knowledge of such debris it would
be possible to gain an adequate knowledge of the organization of towns.” 28
Why, then, did biologists act as if cells existed as some sort of aggregate of
protoplasm? Or, worse, why did they act as if cells were some sort of “building
block” of organisms or the proverbial “unit of life”? Woodger’s work on Golgi
bodies after the war and in embryology by the late 1920s militated against these
notions of ever finding any actual heuristic utility in the laboratory, let alone out
in the broader world. Cells were too complex, too differentiated, too variable in
composition and function. And in organized aggregation, as a portion of tissue
or a larger organ system, for example, they operated still differently, taking on
still more complex roles.
Organization seemed to be a basic fact about the biological—organization
extending across both time and space. All organisms have a tendency to alter
their organization serially across slices of time. In other words, organisms are
“serially ordered,” according to Woodger. That change in serial organization
typically goes by the term “development.” Moreover, though we tend to see only
the adult as the thing itself —the entity frog is also the egg and the tadpole—the
expansion of serial ordering continues through all of life. It would not be too
much of a stretch to say that serial ordering must be the base description of Life ;
in fact, the cessation of this process surely meant disintegration and death. Thus
serial organization should be central to the study of biology, something beyond
the purview of those who study merely the physicochemical constituents of
nonliving nonorganisms. Using this concept of an organism as a serially
organized process, Woodger thought he could resolve many of the long-standing
antimonies in biology: structure versus function, organism versus environment,
and teleology versus causation. 29
In his preformation versus epigenesis chapter, Woodger applied his organic
model to both embryology and evolution. In both development and evolution,
we see eerily similar spatiotemporal repetitions of organized parts arranged such
that, according to Woodger, “individual activities are subordinated as part-events
organized in a particular way which is able to issue in biological responses.”
Unlocking the method by which that patternedness occurred embryologically,
that is, how the organic hierarchy itself unfolds over the life of an individual,
might also unlock the process of evolution. As Woodger saw it, evolution should
not be reduced to vague concepts like “adaptedness” or “increasing complexity
of anatomical structure,” as they often have been. Rather, evolutionary progress
had to be tied to the “achievement of increasing independence of contingent
environmental circumstances.” 30 What we see during both ontogeny
(development) and phylogeny (evolution) is increased resilience by individuals
to environmental perturbations. Karl Ernst von Baer made a similar realization in
the mid-nineteenth century, but Woodger was among the first to highlight that
the more sophisticated twentieth-century study of embryology would lead to an
understanding of the greater pattern of evolution. 31 It was a significant
contribution. Yet Woodger was not pleased with this chapter in particular. He
offered only a synopsis of the myriad difficulties in any explanation of evolution
that relied upon preformation or epigenesist exclusively, no real conclusion.
Upon reflection, Woodger felt Biological Principles did largely accomplish
those two challenges Suttie had tasked Woodger with years earlier: to convict the
biologist of his intellectual confusion and to offer resolutions of the “antimonies”
within biological thinking. Beyond that, Woodger believed the book made
several more specific contributions. First, he had described how scientific
epistemology is actually performed in biology as opposed to physics or
chemistry. His hope here was that biologists could now reject “hypothetical
particles” (read: Mendelian genes) that biologists rendered “more real than that
which they have been devised” to explain phenomena despite the haziness of
actual empirical observation. Secondly, Biological Principles stressed the notion
of becoming as a non-atomistic, time-dependent process of relating characters to
parts of whole organisms. These were concepts borrowed from quantum physics
and from Gestalt theory in psychology. So if there was any reduction to
physicochemical theories going on in biology (and Woodger showed that, at
least tacitly, there was), reduction should be to concepts of process, integration,
network, and wholeness rather than particles in aggregations. Finally, in
Biological Principles Woodger believed he had gone beyond William Ritter’s
1919 Unity of the Organism in presenting a realistic depiction of how the
“‘organic view’ of nature” might impact biology. 32 He demonstrated just how
the notion of internal and multiple relationships could be resolved into flexible
hierarchies. This methodology would be crucial for future biology, Woodger
insisted.
He feared, however, that modern biologists might not appreciate his efforts to
clarify biological concepts: “Biologists have taken the celebrated saying of John
Hunter: ‘Don’t think; try’ too literally. This is not the maxim upon which physics
was founded. You must think first and then try. And you must think about the
right things. In biology, we require to think primarily about biological facts, not
about hypothetical billiard balls.” 33 The tyranny of the pragmatic governed
biology; Woodger had experienced it firsthand. In that climate, mechanism was
just easier. It did not require sophisticated theory, though of course tacitly it did
—some biologists had to imagine invisible (or nearly so) particles possessing all
kinds of powers: “genes-for” any identifiable trait, though the genes themselves
were not seen. Woodger believed his post–Great War generation of biologists
would eventually grasp the importance of theoretical thinking and guide biology
back onto the right path toward organic principles. But who could play the role
of that charismatic leader who would inspire biologists of all stripes to buy into
Woodger’s vision of a carefully reasoned nonmechanistic science—a biology
that would “think first and then try”?

Though Joseph Needham had proven himself to be a “think first, then try” sort
of scientist, Woodger was unsure whether Needham was the person to unify
biology behind the philosophy of organism. He was dashing, charismatic, bold,
and a sharp thinker, but Needham was also a known defender of mechanism and
a biochemist—among the most reductionistic of biological subfields.
Nevertheless, enough of their interests overlapped that Woodger wrote Needham
in February 1929, beginning a process that would convert Needham to
organicism and nudge the entire field of biology toward an alternative to
mechanism and vitalism.
In his February 1929 letter, Woodger asked Needham if he had examined
Biological Principles . Reviews had begun to appear and Woodger wondered if
Needham had read those. Or, if Needham had not had the time to read either
Biological Principles or the reviews, perhaps Needham was familiar with the
works of German-speaking philosophers of biology who inspired Woodger to
write the book: Ludwig von Bertalanffy and Adolf Meyer-Abich. (These two
were among those he had become familiar with during his semester at
Przibram’s Vivarium three years earlier. In particular, they impressed upon him
that concept of cell organization that he regarded as the central claim of
Biological Principles —the concept that undermined any easy support for
Needham’s view of mechanism.) The letter was an indirect challenge to
Needham. Would he reply?
Needham did reply to Woodger on 26 February 1929, but his response was
not encouraging. In it, he essentially restated some of the material he used
against Rignano in Man a Machine . For each example of distinction between
living and nonliving, Needham could find counterexamples where the distinction
collapsed. 34 Why should Woodger or anyone else assume there was a line
between the biological and the merely physical and chemical? If we held to
bioexceptionalism, it was only an admission of our ignorance. Eventually
mechanists would fill in those gaps. Woodger did not respond. So here the matter
rested for about half of 1929—another lapse in their communication.
Early in October 1929, the Cambridge Zoological Society invited Woodger to
speak on the topic “Is there any connection between Science and Religion?”
Needham magnanimously agreed for his college, Gonville & Caius, to host
Woodger. Though the content of Woodger’s talk is not recorded, the event went
well enough. Woodger wrote to Needham on 8 October to express his thanks. He
also enclosed the letter to Needham from May 1928—the one Needham never
answered—which contained many of the same challenges to Needham’s neo-
mechanist philosophy that Woodger spelled out in Biological Principles .
Curiously, Ludwig von Bertalanffy wrote to Needham regarding similar issues
the previous day. 35 Perhaps the confluence of these two philosophically erudite
thinkers startled Needham into reconsidering his own neo-mechanism. Or
perhaps it is just a coincidence that he responded very quickly to this letter of
Woodger’s and seemed to digest Woodger’s 1928 objections to “Recent
Developments in the Philosophy of Biology” (this was the letter Needham had
ignored eighteen months earlier). Whatever the reason, in his return letter
Needham thanked Woodger for his congenial, patient tone: “Your letter made me
feel what I always feel about Ludwig Bertalanffy [sic ], namely, that in all
probability we could come to agreement after a good verbal discussion—a
feeling I never have, for instance, about E. S. Russell or W[alter] McDougall.” 36
The friendly tone characterized their subsequent correspondence as well. By the
end of November 1929, Needham asked Woodger’s opinion regarding the status
of organized cell structure. When Woodger proffered the expected organicist
response—“‘Cell’ is not a name for a thing but for a type of organization”—
Needham did not respond with the same disputational examples as he had in the
previous two years of exchanges. 37 It was a small victory for Woodger.
Their correspondence continued and, in fact, the pace between letters
quickened. Needham mailed Woodger the introductory chapter to his
monumental Chemical Embryology , which was at the printer’s proof stage at the
time. Woodger tried not to take offense at Needham’s portrayal of the “third
way”: “I can quite see that an undiscerning reader might regard me as an
‘opponent’ of ‘mechanism,’ whereas all I ask is that mechanism should be used
intelligently rather than blind-enthusiastically. I am even more of a ‘mechanist’
than you are because I believe it has to be taken far more seriously from the
metaphysical standpoint than you appear to do.” The real bone of contention,
revealed Woodger, was the fundamental inconsistency in Needham’s neo-
mechanism—his provisional or methodological mechanism:

 I hope that some day your interest in synthetic [Kantian] metaphysics will wane a little and allow
you more time for analytic metaphysics, logic and epistemology[.]
You say that natural science does not (?even cannot) assert anything about what is
metaphysically the case. Natural science is bound by a certain methodological procedure . . . and
consequently its results and assumptions are bound to assume a certain form[.]
But if this is the case what bearing has the progress of bio-chemical discovery upon vitalism?
As far as I know Driesch has not said that biochemistry should or cannot be pursued. All he has
said is that it is not competent to reveal the “secret of life” (whatever that may mean)[.] But your
own arguments surely lead to that same conclusion???[sic ]
Note that I am not arguing in favor of Vitalism. I am only pointing out what seems to me to be
an inconsistency in your own arguments, and I hope I have made it clear. If you are only going to
give natural scientific propositions a purely “methodological” and not a metaphysical value, you
cannot possibly use them to refute a metaphysical theory. 38

In one sense, Woodger leveled a damning critique, especially showing how neo-
mechanism was not in principle completely distinct from Drieschian vitalism—
the position Needham wished to discredit. Methodological mechanism could
support, or at least coexist with, vitalistic metaphysics, in other words.
Surprisingly, Needham accepted Woodger’s critique without much resistance. 39
In fact, while Woodger penned this attack on Needham’s neo-mechanism,
Needham had finally settled down to read Biological Principles . He was just
then composing a long and largely positive review of it for the journal Mind . 40
Needham’s review conceded that both the vitalism of Driesch and the
mechanism of Loeb contained “kernels of truth.” Because Needham basically
equated “mechanism” with “scientific analysis,” it is unsurprising that he found
truth in it. Yet vitalism had value as well: it reminded researchers that “the living
organism is certainly a harmony of reacting parts [ . . . not] a simple heap of
molecules, as it were, flung together; it is composed of parts which may behave
very differently when in isolation from what they do as parts in the functioning
organism.” 41
Biological Principles mainly dealt with this all-too-acute problem of
organization and, while it offered little for the biologist to do with the
information presented, it certainly chided scientists for their overconfidence; it
convicted the biologist of his intellectual confusion, to use Ian Suttie’s phrase.
Needham took note that Woodger specifically targeted biochemists like himself
for regarding mechanical explanations exclusively. Surprisingly, he largely
agreed with Woodger’s assessment that mechanism was an overly hasty
approach.
So far, so good, but there was more to the otherwise glowing review. Though
he welcomed Woodger’s philosophical rigor, warnings to bench scientists, and
his honest assessment of modern biology, Needham worried that Biological
Principles was too little focused on practical matters. What were biochemists
like himself supposed to do with that conviction of intellectual confusion
Woodger successfully laid upon them?
When he read the published version of Needham’s review, it was Needham’s
insistence on laboratory application in particular that distressed Woodger. “I was
disappointed in your review of my book in MIND,” Woodger wrote to Needham,
“because you seem to have misunderstood my intensions to a greater extent than
I should have expected.” 42 Woodger was hoping to alert philosophers to the vast
tangled web of concepts in biology in need of clarification, to draw in the likes
of G. E. Moore, Bertrand Russell, and Alfred North Whitehead. He feared
Needham scared off philosophers and, in so doing, had killed the philosophy of
biology project in utero, as it were: “As I am particularly anxious to persuade
philosophers of the right type to take more interest in the foundations of biology
(because I think they might help us) [Needham’s review] seemed to me a pity,
and I thought it right to say something to counteract the possible effects of your
review.” 43 The “something” Woodger wrote was a letter directly to the editor, G.
E. Moore; Moore promptly published Woodger’s letter in Mind . 44 Woodger’s
frustration echoes through the letter. Needham had focused on the wrong parts of
the book. Biological Principles had two main missions, neither of which the
review explored. First and foremost, Biological Principles addressed the “well-
known antithetical character of biological thought”—preformation versus
epigenesis, mechanism versus vitalism, and so on—in order to apply “‘the
logical clarification of thoughts’ (Wittgenstein) and the ‘study of the a priori ’
(C. I. Lewis).” So many of the assumptions leading to enduring conflicts in
biology had their roots in the profound messiness of biological concepts.
Needham had neglected to acknowledge this. Perhaps Needham missed this
contribution to cleaning up concepts because, according to C. I. Lewis,
foundational concepts precede scientific inquiry itself—they are assumptions
brought directly to the laboratory bench by the scientist, who is largely unaware
of them. Needham must have been unaware that he, too, brought these concepts
qua scientist. This lack of awareness made Needham miss the second mission of
Biological Principles altogether: to “attempt a rapprochement between the
philosophical and the natural sciences,” to create an incubation space for the
philosophy of biology. Sadly, scientists—and Woodger included Needham here
—seemed to regard philosophy as “either an inferior rival . . . or as a sort of
constable who keeps the peace between science and religion by means of a
cudgel, in the shape of philosophical skepticism, which is flourished from time
to time when science gets in difficulties.” When someone with training and
interest in both philosophy and science takes up the “thankless task” of
investigating biological concepts, attempting to root out the “extravagant and
demonstrably false assertions” made by scientists in defense of their quest for
“unlimited range” for their claims (Suttie had called this “poaching”), those
philosopher-biologists seemed destined to be misunderstood. And that was in the
best case; more likely, they simply would be ignored.
In June 1930, Moore approached Needham to ask for a response to Woodger.
45 A month passed. Eventually, Needham had to apologize: He had mustered a

very inadequate response and, consequently, Moore had let the matter drop. 46
The most prominent early debate setting the table for the philosophy of biology
slipped quietly away without much comment.
The whole episode with Needham indicated to Woodger that, if he wanted a
philosophy of biology friendly to the “third way” that actually impacted
scientific concepts and garnered the interest of skilled philosophers, he would
have to go on the offensive. More than likely, the burden might be his alone to
carry for a while. But he thought his efforts worth it in the end: “I think there
ought to be such a thing as ‘pure biology,’ i.e., thinking about biological data in
biological terms without reference to other sciences. When we have made some
progress in this direction we shall be in a better position to understand how
organization at the biological level is connected with the types of organization
studies in chemistry and physics.” 47 If biology was to be free of the artificial
noose of mechanism, thought Woodger, it would have to be an independent
science, logically—not just in practice—distinct from physics and chemistry.
Organicism—thinking about the nonmechanistic unity of the organism—would
be the best way to ensure biology’s independence.
This encounter with Needham, the third thus far, showed promise. But
ultimately, it may have backfired. Woodger was undeterred. His next move in the
project to extend organicist thinking was to publish a series of three lengthy
articles in the Quarterly Review of Biology . Each essay was philosophically
robust, intimidating even. Yet he built the whole three-part series, entitled “The
‘Concept of Organism’ and the Relation Between Embryology and Genetics,”
upon the relatively simple premise that most scientists possess neither the time
nor the inclination to systematically analyze their own assumptions. As a
philosophically adept biologist, Woodger was stepping into the gap to elucidate
some of those assumptions.
Raymond Pearl of Johns Hopkins University, a leading American biologist as
well as the editor of the Quarterly Review of Biology , sought out Woodger for
his ability to offer this rigorous kind of analysis: “The difficulty is one that you
must have already experienced. Most working biologists, at any rate in America,
do not like to think and look with a very fishy eye on anything which savors of
philosophy. Just now with us anything or anybody not overtly worshipping with
adulation at the shrine of genes and ‘crossing over’ is considered low and wholly
lacking in intelligence.” 48 Woodger may have understood Pearl’s point, but he
forged ahead writing philosophical articles and speaking about the need for
biologists to come to terms with the tacit philosophies they were following. As
Woodger persisted, Pearl wrote again; he felt compelled to prepare Woodger for
further frustration:

The truth is that most biologists (and scientific men in general, but biologists probably more) hate
new, and especially unorthodox, ideas as badly, if not worse, than a parson does. So far as I can see
it has always been so, and probably always will be.
. . . Seriously I think that to get general methodological ideas across it is going to be necessary
to show, by concrete cases thoroughly worked out, that actually such ideas do specifically help us
to get ahead in real comprehension and insight. Simple minds, such as all of us ‘working’ biologists
have need of simple, and above all, concrete pap. 49

Pearl did publish all three segments of Woodger’s “‘Concept of Organism’” over
1930 and 1931. Given the immense amount of publication room he devoted to
the essays in a major journal where page space was always at a premium, Pearl
must have valued the contribution highly.
Each “‘Concept of Organism’” article addressed the perennial conflict of
preformation versus epigenesis to which Woodger had devoted the nearly 100-
page ninth chapter of Biological Principles . He hoped over the series of three
articles to sharpen that analysis—he let on that he was dissatisfied with his work
in the book—through symbolic treatments of the notion of levels and hierarchies
in biology. 50 After all, “third way” thinking must mean that the wholeness of the
organism in some sense influenced the behavior of the organisms’ systems from
the top downward. These in turn influenced the behavior of smaller units: cells,
chromosomes, genes, etc.
Given how he introduced his first essay, it is clear that Woodger took Pearl’s
—and Needham’s and Suttie’s, for that matter—concerns about the need for
concreteness to heart. While in his essays he tried to convince biologists both
that they needed to consider more carefully their underlying concepts and that
philosophy of biology was as important to biology as theoretical physics was to
physics, he did provide definite examples: “[S]uppose we have two rabbits—one
white and one black—born in one litter. Then we say that this difference was
correlated either with a difference between the intrauterine conditions of the two
embryos, or (which would be considered more probably) with some difference
which was present throughout the development of both, i.e., a genetic
difference.” 51 The difference between the baby rabbits, in other words, could be
either strictly the direct result of the immediate context, i.e., the condition of the
mother’s womb, or as a result of the genetic program carried within each rabbit.
Of course, there is the synthetic option—it’s really some of both—which is what
Woodger believed. Indeed, he offered multiple examples along these lines to
point out two otherwise tacit features of biological inquiry: (1) The real locus of
embryological investigation is the differences in relationships between
organisms or systems on multiple occasions, while genetics investigates changes
in intrinsic properties of entities limited in scope, and (2) communication
between biological subfields is so muddled that a new language needs to be
invented to resolve the apparent difficulties.
Primarily, Woodger stressed the same main point through all three essays:
Despite appearances, nearly all biologists really investigate different levels or
compartments of biological content. These levels operate as if they are
balkanized. And the differences between areas are most pronounced when
investigating the concepts of change over time evident at each level. Change
takes three forms in biology, according to Woodger. First, the configuration
between elements of a system can change. Or, secondly, the properties of the
elements themselves can change. Finally, the elements can grow or multiply
(usually necessitating a change in configuration). Woodger represented these
varieties of change with a simple diagram (Figure 3.1 ).

In his figure 2 from “‘Concept of Organism,’” a is his initial organized entity,

b is a change in relationships between the parts, c is a change in the properties of
one of the parts, d is a change in number of parts. 52 By being clear about what
kind of change is operating, believed Woodger, we can clearly delineate whether
we are discussing compatible concepts between levels. For instance, if genetics
treats a realm where the majority of change is “c ” change—change in the
elements themselves—but embryology is primarily “b ” and “d ” change—
change in configuration and numbers of elements in that configuration—then to
describe embryological development in genetic terms would require no small
amount of translation to make explanations commensurable. Simple descriptions
of development in terms of genes would make no sense without this translation
—these two arenas are describing completely different kinds of behavior among
their components.
In “‘Concept of Organism’ I,” Woodger stressed the importance of change not
among entities per se (c in the figure), but change in the relationships,
orientations, or configurations (b in the figure). In fact, for the article he adapted
the terminology he had introduced in a contemporaneous letter to Needham:
“[I]n an organism, the properties of a component depend upon the component of
which it is a component (relational properties) as well as on the parts into which
it is analysable. This I imagine to be what people mean when they speak of
‘organic determination.’” 53 One reason that so few biologists appreciated this
notion of organic determination, thought Woodger, was because biological
communication as a whole was so fragmented. In the particle-centered world of
genetics, there was no language for communicating in terms of relational
properties. Thus Woodger, secondarily, promoted the use of a biological bridge
language, by which he meant specifically a formalized symbolic language. There
were two reasons to use symbolic language in biology, just as it had been used in
physics. Simplified formulae would not only improve clarity in biological
theory, a symbolic bridge language would help provide a path out of the
theoretical entanglements that engendered conflict—especially between fields on
different levels of biology, like genetics and embryology. If geneticists and
embryologists were to speak to, and not merely past, each other then they would
need a clear language by which to do so. Woodger’s real goal in these articles
was to promote easier communication, with fewer complications that led to
division and competition in biology—a unified pursuit of explanations regarding
living things. This emphasis on formalized language in biology was merely
secondary and methodological. Yet it became the aspect of his work for which he
would later become well known and, after his death, derided. 54
He infused such formulae into each part of “‘Concept of Organism’” with
increasing frequency. When explaining experimental results, such as in the rabbit
example previously, we could use what Woodger called a “Postulate of
Analytical Interpretation”: 55

(1) D(A, B).C.D(OA , OB ) or D(EA , EB )
(2) D(A, B).C.D(OA , OB ) and D(EA , EB )

where D is the “‘difference between the following,’” C means there is a causal
correlation, A and B mean two experimental observations of events, O stands for
an organism, and E for the environmental context. So in the first explanatory
formula, we would say that the differences in two observations or experiments,
A and B, were probably due to either changes in the organism or changes in
environmental conditions. That would be true even if “environmental
conditions” simply meant the trials conducted in the laboratory, chemicals
administered, and so forth. The results of many experiments would comport
better with the second explanatory formula: Both the organism and the
conditions changed, leading to whatever change was observed in the experiment.
Most importantly, Woodger added a third explanatory sentence to these two
typical ones:

(3) D(A, B).C.D(RA, RB)

R meaning the relations of the constituent parts. So in this third sentence, the
differences between the relational properties (the “b ” in the above figure) would
account for the differences in the observed phenomena. Woodger saw this as the
most important explanatory sentence in embryological development. In
embryology, “‘totipotent parts’ are transplanted to different situations” and this
transplantation—the rearrangement in the relations itself—makes all the
difference. 56
Woodger’s new generation of organicism becomes clearest here. Properties of
parts that are not intrinsic to the part itself depend upon their relationships within
the organized hierarchy. He contrasted this depiction of biology with what he
called “the machine analogy” or mechanism. In machines there do not appear to
be levels of organization wherein the parts can be arranged homogenously—in
the way that a section of tissue, say, appears to be homogenous—and yet the
parts actually become different things depending upon their relations within the
larger hierarchy. Or, put another way, in a machine, relationships between parts
are external, superimposed, designed; in an organism, the parts are internally
arranged, self-arranged, in fact, and the arrangement matters not only to the
function of the whole but the composition and behavior of the parts themselves.
Still it is not enough to merely study the static relationships, insisted
Woodger. Because these relationships within the organic hierarchy of the adult
come about as a result of development from an egg to larval stages, the proper
arena of biological study is embryology. We can only truly know if we know
developmentally . Only by observing development can we understand how the
hierarchy itself gets formed through time, how cells begin their journey on the
way to becoming a member of a particular level, performing a particular
function in the overall organismal hierarchy.
What about subfields of biology such as genetics or botany? Should
biologists merely study the elaboration of “relational properties” of cells and
ignore the adult version of an organism or the genetic underpinnings of traits?
Woodger did not make such a claim, though more recent scholars have misread
him as doing so. 57 Instead, he asserted that, properly realized, genetics should be
considered master of only a single domain of biology—that of intrinsic
properties. The problem with biology as currently structured was that “lower”
subfields of biology—meaning those that studied smaller entities and processes,
genetics for example—presumed that their level of study was sufficient (not
merely necessary) for understanding higher levels. In other words, fields like
biochemistry and genetics “poached” from embryology.
Primarily this occurred when geneticists held too closely to August
Weismann’s nineteenth-century concept of “chromatin that controls the
properties of cells,” according to Woodger. If geneticists presumed that the
intrinsic properties of genes in the cell nucleus were the controlling mechanics—
Weismann’s “‘primary Anlage,’ ” the formative primordium—they could do so
only by ignoring problems like tissue differentiation: Why do these cells express
one set of properties while those express different properties, given that both
share the same material in the cell nucleus? 58 From Woodger’s perspective,
geneticists were returning to the preformationism of a bygone era. Specificity of
biological properties, he insisted, was not merely a result of cell differentiation
according to the original plan ensconced in the egg: “All this means that there is
another developmental process to be considered, namely the elaboration of
cellular parts, of parts, that is to say, which are analysable into cells in certain
determinate relations, and thus constitute parts belonging to a level above that of
the cells. . . . [I]t is upon these parts and their relations . . . that we have been
driven to place the ‘responsibility’ for the differences in the histological
elaborations of the various non-cellular parts which (in their relations) constitute
them.” 59 Geneticists relied upon a version of preformation: They regarded all of
the instructions for the complete adult organism as preexisting in their entirety
on chromatin in the zygote. In so doing, they actually took explanatory territory
away from embryologists, among others. But all that “genes” provide—all that
genetics should be explaining, according to Woodger—are the directions for
chromosomes to supply the material necessary for “possible histological
elaborations of which a given race is capable” and how these materials persist
and divide within a greater cellular and organismal context. It is up to
embryologists to demonstrate how relationships in the matrix produced by large
numbers of cells have their own rules, as it were. Those rules were not confined
to the nucleus of cells, tied up in chromosomes alone. This was the space for
something at a higher level than genetic explanations.
As Woodger continued with the “‘Concept of Organism’” series into 1931,
his use of symbolic formulae became ever more pervasive. This trend reflected
his reading of Alfred North Whitehead and Bertrand Russell’s three-volume
Principia Mathematica (1910, 1912, 1913). The Whitehead-Russell Principia
analyzed the conceptual structure of mathematics in order to refine loose
concepts that presented problems in quantification itself. To assist their effort,
Whitehead and Russell fashioned an extensive symbolic language. Ironically,
their monumental achievement, invented to add precision and clarity,
inadvertently worked to inhibit communication, broadening the divide that had
begun to separate professional philosophy in the UK from that on the Continent,
with analytical, language-focused philosophy slowly but surely breaking off
from the Continental tradition. For Woodger, the symbolic language that
accompanied the growing Anglo or analytical philosophical movement was
unquestionably the most efficient way to address the persistent dichotomies in
biology that he had already identified in Biological Principles .
By 1931, Woodger was moving toward his own axiomatic masterwork, a true
“Principia Biologae” to place alongside the Principia Mathematica of
Whitehead and Russell. Like Whitehead, Woodger would need an analog of
Bertrand Russell, a young radical willing to challenge orthodoxy. Joseph
Needham would make such a partner. But at the beginning of 1931, Needham
was still in his own neo-mechanist camp.
 4

THE TIPPING POINT

B y the end of 1931, Needham had joined the new generation of organicists.
While Woodger’s logic alone could not convince him to fully accept the “third
way,” a concatenation of unexpected events and figures did finally convince
Needham. These included two unexpected conference speakers; two new
relationships with other British scientists interested in alternatives to mechanism
and vitalism, J. D. Bernal and C. H. Waddington; a more robust encounter with
Alfred North Whitehead’s philosophy; and a tiny experiment conducted by a
young, practically unknown woman in the middle of Germany. By 1932,
Needham had converted from organicism’s most articulate detractor to a
committed proponent. Though a single scholar, his impact, especially as an
organizer and an interdisciplinary bridge builder, would resonate through
biology for years, impacting the rise of epigenetics itself. His story reflects the
larger trend in European science: By the 1930s, the pendulum of scientific
worldviews had clearly swung away from the harder mechanism of Jacques
Loeb. Whether that pendulum would swing toward vitalism or a “third way” was
much less clear.

From 28 June to 3 July 1931, London hosted the Second International Congress
of the History of Science and Technology (SICHST). By this time, Needham had
published his three-volume Chemical Embryology and a curious collection of
essays entitled The Sceptical Biologist , another homage to an Enlightenment
natural philosopher (Boyle rather than La Mettrie). Needham’s ability to mix
science, philosophy, and politics attracted wider notice. On this reputation, he
brought several significant speakers together for the SICHST session “The
Historical and Contemporary Inter-Relationships of the Physical and Biological
Sciences,” including both members of the older generation of organicists (E. S.
Russell and J. S. Haldane) and up-and-coming scientists, including zoologist
Lancelot T. Hogben. Needham invited Woodger to speak as well.
It was here at the SICHST that an unexpected event happened. A cadre of
distinguished scientists and philosophers from the Soviet Union arrived with
little advance warning. Popular Bolshevik Nikolai I. Bukharin headed the
delegation. Needham welcomed Bukharin and his comrades to London, and
then, after the conference, also spearheaded the publication of their English-
language talks as a small book, entitled Science at the Cross Roads (1931). 1
As Needham no doubt knew, V. I. Lenin had once dubbed Bukharin the
“Beloved of the Party.” To those outside of the party, including Needham,
Bukharin appeared a shining beacon of socialism, numbering among the most
important surviving Bolsheviks, once head of the Comintern, a Central
Committee insider, and editor in chief of Pravda . Neither Needham nor the
other British scholars realized the degree to which Bukharin’s position had
suffered of late. Stalin had chastened Bukharin for supporting the late Lenin’s
New Economic Policy—a policy that Stalin had also once supported—by
pushing him out of the Central Committee. Bukharin’s long-term loyalty to the
Communist Party and his role in the revolution ensured that he retained some
role in Soviet leadership, especially once he was reelected to the Central
Committee at the Sixteenth Congress in 1930. Yet his sincerity, popularity, and
connection with Lenin ironically marked Bukharin for destruction. As part of
Stalin’s more general program of eliminating his competition, Bukharin suffered
perhaps the most famous and tragic show trial of 1938. Stalin ordered
Bukharin’s execution that year despite Bukharin’s impassioned appeals to his old
comrade. 2 (His memory lived on: Bukharin was the inspiration behind the
character Rubashov in Arthur Koestler’s heartbreaking novel Darkness at Noon .
Bukharin’s widow had his reputation rehabilitated in 1961.) But these tragedies
were a few years in the future. At SICHST in 1931, Bukharin was a minor
celebrity. His address, “Theory and Practice from the Standpoint of Dialectical
Materialism,” challenged a number of arrangements within the Western
scientific establishment that he regarded as sacred cows—most prominently,
scientific epistemology and the sociology of the laboratory. Bukharin’s address
shocked and appalled some scientists.
Needham, by contrast, was starstruck. 3 He found Bukharin’s critique of the
complicity of science with capitalist exploitation particularly persuasive in
conjunction with Needham’s already strong support of British socialism and
Noel Conrad’s teachings at Thaxted. And Bukharin’s stress on the importance of
the dialectic caused Needham to rethink the framework of the science in which
he had been trained as a biochemist. As he began to reflect on his experience as
a biochemist, he realized that simplistic mechanistic thinking was preached and
practiced exclusively by his colleagues even when that perspective didn’t follow
the phenomena. It wasn’t merely that vitalism was dismissed as unscientific; no
one took any perspective other than mechanism seriously enough to be explored,
even if that exploration would ultimately lead to firmer grounds for rejection.
But perhaps the dialectical view Bukharin trumpeted meant that progress was
possible; the appearance of mechanistic determinism was only a by-product of
the current configuration. One didn’t need vitalism, only a way out of
mechanism, a synthesis, perhaps. It was that aspect of the Soviet contribution to
the SICHST—the possibility for social justice, for progress—that Needham
emphasized in Science at the Cross Roads . Ironically, it was this interest in
social justice and friendliness with the Soviet delegation that would bring
Needham unwelcome attention from British intelligence operatives. That
attention, in turn, dramatically changed Needham’s career during and after the
Second World War.

Crystal physicist John Desmond “Sage” Bernal (1901–1971) also attended the
SICHST and was nearly as impressed by Bukharin as Needham was. While on
the surface, Bernal saw little to dislike in standard mechanism, there was
something more in what Bukharin offered. It made him curious.
In some ways, Bernal proved an ideal counterpart to Needham, and the
relationship between them that started in 1931 would sharpen both their
scholarship and their politics. While equally brilliant, equally pugnacious,
equally inspirational, Bernal had one flaw (or maybe it was an advantage) that
Needham did not share—Bernal was Irish. Born in County Tipperary in the west
of Ireland, Bernal had an acute sense of “otherness,” especially as his world
collided with the English one. In 1926, when Bernal was only twenty-five years
old, he sought to write an autobiography that would capture his experience as
“other.” He titled his unpublished autobiography “Microcosm”:

I have set myself the task of writing down . . . the sum of all the influences that have borne on me,
and what my mind makes of these in its knowledge and action. It would seem in some ways more
suitable to wait till age and experience had taught me my ignorance and futility, such a book may
be without impertinence written at 75, but I do not propose to wait till then. Not that I despise the
wisdom of age or prefer that of youth, but that I think . . . that they are different wisdoms. 4

Certainly he had the same confidence in his own abilities as Needham. And
those abilities did prove to be prodigious.
 Those who have only slight knowledge of Bernal think of him as an
uncomplicated atheist. But Bernal was brought up as a pious Irish Catholic boy
in a nurturing home. He was educated at the diocesan school at Nenagh, and was
good enough at his studies and in his faith to be sent to Hodder, which was the
oldest and most distinguished Jesuit school in the UK, in Lancashire. Bernal
loved the religiosity of Hodder. But Hodder fed into Stonyhurst, about which
Bernal had more mixed feelings. In “Microcosm” he wrote, “James Joyce has
said all that need be said about it. Dark corridors, wandering priests, terrible
sermons of sins that must not be named lest they be practised. . . . But against
that there were forests of flaming candles, golden vestments, and the resonant
chanting that tore out the soul for God.” Like Needham, Bernal would carry this
passionate tension with him for the rest of his life. On the one hand, there was
his contempt for the fear mongering and authoritarian moralizing of the church.
Bernal, unlike Needham, would later state that he had given up his faith—how
could he be a proper Marxist and continue to accept the power of the religious
“opiate”? But on the other hand, Bernal expressed admiration, maybe even
longing, for the systemic order and respect for tradition exhibited by the church.
In fact, like many formerly pious converts to atheism, Bernal expressed that his
conflict was never with Christian ideals, but with the inability of the human
collective to live up to those ideals. “I could not help seeing [the church] as an
active agent of political reaction throughout the world,” he said once. And later
he would admit that, “Seen as the one background of the unexplained of life the
church was impregnable, but seen in its place and time it appears an ideal human
construction, as compelling and as fallible.” Interestingly, some of Bernal’s
distrust of organized religion was born not merely from his experiences at
Stonyhurst, but out of arguments with conservative anti-Republicans during the
1916 Irish rebellion. As Bernal put it in his diary at the time, he was treated
“with absolute loathing” when he tried to argue for the motivations of the Sinn
Fein. 5 Bernal found that Christianity’s adherents quite frequently draped their
nationalism, classism, sexism, and even xenophobia with religious sentiment. It
rankled him. As an undergraduate at Cambridge, it took only a small push by a
classmate to convince Bernal that socialism as practiced by fiery Russian
Bolsheviks better exemplified the Gospel struggle for a brotherhood of all men
regardless of race, class, religion, or scientific discipline than the path preached
by stodgy churchmen. Though the Stalinist bastardization of it would disappoint
him repeatedly, Bernal permanently converted to socialism by his early twenties.
Bernal, like Needham, yearned for the establishment of a utopian society
through the cooperation of technocratic and humanitarian ideals. Though one
remained formally committed to a version of Christianity and the other did not,
both Needham and Bernal articulated the inspiration for their ideals in language
derived from the Gospels. When he was 28, Bernal published his first book.
Though he was already committed to socialism by this time, he entitled the book
The World, the Flesh, and the Devil . He reprinted it forty years later, stating, “I
have a great attachment to [the book] because it contains many of the seeds of
ideas which I have been elaborating throughout my scientific life. It still seems
to me to have validity in its own right.” It articulated a feeling shared by
Needham and Woodger, among others: to pursue knowledge both scientifically
valid and encouraging of human flourishing. Neither mechanism nor vitalism
seemed to offer both.

Woodger, too, listened intently at the SICHST. He had delivered a paper in

Needham’s “Historical and Contemporary Inter-Relationships of the Physical
and Biological Sciences” symposium. He had heard Bukharin and the other
Soviet speakers. But unlike Needham and Bernal, he heard nothing particularly
novel or striking. If anything, Woodger became still more convinced that biology
needed to be placed on new foundations congenial to the “third way,”
foundations that would require more precise language in order to avoid repeating
the same antimonies (e.g., mechanism vs. vitalism) over and over again.
As each scholar at SICHST delivered his talk, Woodger mentally categorized
the speaker into one of three sets, corresponding to their biological worldview.
Interestingly, Woodger defined worldviews according to what each man opposed
rather than the position for which he argued. On the “anti-mechanist” end of the
spectrum, Woodger located the aged but still energetic J. S. Haldane and E. S.
Russell. Unsurprisingly, they “were very emphatic in denouncing the classical
physico_chemical [sic ] methods as applied to biological problems.” 6 Opposed
stood Joseph Needham, John D. Bernal, Victor Cofman, and Lancelot L. Whyte
—all of whom espoused different versions of anti-vitalism. Woodger, of course,
stood for an alternative to both. He was joined there by Boris Zavadovsky from
the Soviet contingent. Woodger and Zavadovsky leveled similar critiques:
Biological concepts were too sloppy to compare to those in physics. Until
biologists were as committed as physicists to the kind of rigor in communication
and in the development of the theoretical structure of their field, conclusions
both for and against mechanism were premature and liable to create more
misunderstandings for future biologists.
During the discussion period after the talks, Woodger pressed his assembled
colleagues. Would they follow through on their talks and take a step toward
clarifying biological language? Could they at minimum commit to an
“operational definition” of life itself? Woodger pointed out that none of them
could competently assess any interrelationship between biology and physics—
the whole point of the symposium—when one could not be sure that the
disparate biological positions presented during the symposium were in fact
compatible with each other. Representatives from both ends of the
mechanist/anti-mechanist spectrum agreed with Woodger, and they gathered
during the meeting to produce and circulate a document entitled “The Concept of
Life: Operational Definition”. Circulated by Cofman, a physical chemist,
“Concept of Life” contained a chart that attempted to identify the properties of
various complex “vital phenomena” and then find analogs among other kinds of
physical entities not usually recognized as “alive.”
When drawing up this table (table 4.1 ), Cofman thought it important to de-
biologize some of the language. Growth, for instance, could be regarded as a
simple “increase in matter and free energy.” Adaptation and evolution might be
merely “maintenance of optimum conditions.” In fact, one striking feature of this
chart is the ease with which it moves between “complex vital phenomena” and
“simpler physical analogies.” Don’t we mean something much stronger when we
use “autonomous movement” rather than “unpredictable movement”? Is a stone
falling down a waterfall really analogous to a salmon leaping up a waterfall? Are
“reproduction” and “heredity” really just “periodicity in time”? By some
appearances this schematic attempt to make clear the possible interrelationships
between physics and biology only begs the question: Don’t we have to assume
that there is some sort of connection here before we draw these specific
analogies? In other words, don’t we already have to agree with the mechanistic
worldview before we can say that what we mean by “memory/volition” has a
deep enough connection to what we mean when we say “dependence upon past
and future events” to make it analogous?
This indeed was Woodger’s critique of Cofman’s table. He disagreed with
how easily the nonbiologists made their analogies: “Growth” couldn’t simply be
“increase in matter and free energy”; “memory/volition” couldn’t be reduced to
“dependence upon past and future events.” Whatever the analogs between vital
and nonvital activities were, these were too simple. J. S. Haldane piled on to
Woodger’s critique. The entire scheme was too austere, too not-empirical for
Haldane. Sounding a complaint belonging to that first generation of organicists,
Haldane stressed that biology needed to go back to Aristotelian induction—
letting the “nature of biological facts themselves” speak. Maybe there weren’t
simple physical analogs.


Needham did not respond to the table itself, but his position at the meeting
was clear enough: “Organization alone is no explanation of anything.” 7
Nevertheless, Woodger’s position intrigued Needham, not least because it
seemed to build from the philosophy of Alfred North Whitehead. “[T]he position
of a part in the whole would be alone responsible for some of the organization
phenomena exhibited by the whole” was how he understood Whitehead and
Woodger’s organic philosophy. Needham had begun to warm to this idea. Add to
it Bukharin’s emphasis on the dialectic—Needham knew there was something
there, even if he couldn’t put his finger on it.

The second transitional moment in 1931 occurred when General The Right
Honourable Jan Christiaan Smuts, former prime minister of South Africa, rose to
speak as the president and keynote speaker of the British Association for the
Advancement of Science annual meeting. Given the intellectual and political
stature of Smuts, his address endorsing the importance of irreducible systems
indicated just how pervasive organicism had become. Within a few years,
support had become so great that one commentator claimed, “Nowadays almost
every self-respecting biologist, philosopher, and psychologist” adhered to it. 8
On paper, there could hardly have been two more antithetical influences on
Joseph Needham and the burgeoning organicist movement than Nikolai
Bukharin and Jan Smuts. One a Marxist revolutionary, the other a stalwart Tory;
one an earnest intellectual who would be imprisoned for standing in the way of
Stalin’s despotism, the other a shrewd political insider who would be honored
and extolled by the British crown even after ordering the slaughter of subjects in
African regions neighboring his own. His former comrade, Josef Stalin, would
execute Bukharin and erase his name as much as possible from the annals of
history; Smuts would be lauded and immortalized as coauthor of the United
Nations’ charter in 1945. 9 Yet when Smuts addressed the British Academy for
the Advancement of Science in September 1931, he tapped into the same
discontent with mechanism that Bukharin had revealed in his SICHST essay the
same year. By 1931, that discontent had crystalized around some of the issues
first addressed by those scientists who just knew that something real
distinguished the living from the nonliving.
After serving as prime minister of the new Union of South Africa from 1919
to 1924, Smuts retreated from public life (temporarily) to research and write his
magnum opus, Holism and Evolution (1926). Though Smuts was not a biologist,
the book accurately captured the growing frustration with mechanism: “At
present our concept of life is so indefinite and vague that, although the kingdom
of life is fully recognized, its government is placed under the rule of physical
force or Mechanism. Life is practically banished from its own domain, and its
throne is occupied by a usurper. Biology thus becomes a subject province of
physical science—the kingdom of Beauty, the free artistic plastic kingdom of the
universe, is inappropriately placed under the iron rule of force.” 10 Modern
biology, thought Smuts, was being robbed of what made it interesting in the first
place. A biologist could not empirically describe the behavior and appearance of
his or her chosen organism and its relationship to other organisms and just leave
it at that. In place of traditional empirical descriptions , biologists needed
explanations . And the only thing that counted as an explanation must be given
using the language of unseen particles—the language of physics, in other words.
Even the one genuinely biological explanation, evolution, had been redefined in
a foreign tongue, according to Smuts. Evolution had once been so wonderfully
captured by Charles Darwin as benevolent, Nature “daily and hourly
scrutinising, throughout the world, the slightest variations; rejecting those that
are bad, preserving and adding up all that are good.” 11 A blind sorting machine,
a Maxwellian demon clothed in the statistics of biometricians and wielding the
atomic units of Mendelism, had replaced this image. 12 Natural selection, the
crowning jewel that set biology apart from all other domains, had been
repurposed to serve the physicochemical mandate.
The reason behind this sweeping reclassification of the biological as
something mechanical, speculated Smuts, was due to the overconfidence of
scientists of the nineteenth and early twentieth centuries. They convinced
themselves and, perhaps especially, Victorian society that they firmly understood
causation itself. They believed that one could not find features in the effect that
were not already there in the cause . Laplace inaugurated this perspective as law
in the late eighteenth century. 13 It was only a short line from that dogma to the
mechanistic reclassification of life and mind. “Life” reduced to matter; so did
“mind.” Using the analogy that Smuts preferred, scientists and philosophers of
the nineteenth and early twentieth centuries saw matter as the musical
instrument; everything else—the living music of existence—was merely the
noise made by the vibrating instrument. Though the music could be quite
beautiful, ultimately this metaphor subjugated its importance to the instrument
itself. It was only the instrument that was lasting, real. The music, which is to
say life itself , was epiphenomenal, lost on the wind once the quivering of the
instrument ceased. Unsurprisingly, this sort of philosophy carried clear practical
ramifications: Any object of study other than physics and engineering became a
preoccupation with “merely transient and embarrassed phantoms on the stage of
existence,” dismissed as so much stamp collecting. 14
Without a doubt, this old physicochemical concept of causation was
understandable, elegant even. But it could not make sense of one glaring
distinction between the living and the nonliving—a difference in type, according
to Smuts, not just in degree: the ability of living things to regenerate and
reproduce. Even if some crystals could tessellate outward in mock reproduction,
this crucial combined ability of regeneration and reproduction existed first,
insisted Smuts, at the cellular level, not at the bare chemical. From the cellular
level onward, repair and reproduction became a trait shared by all organisms, no
matter their size and complexity. This trait did not exist at any other simpler
level: not at the subatomic, atomic, or molecular levels, and not in larger
colloidal substrates—what T. H. Huxley had called “protoplasm”—either. In the
cosmic process of evolution, the combined trait of self-repair and the ability to
pass something of oneself on to the future even though one’s current existence
was coming to an end—this function , in other words—must have evolved out of
preexisting substance. But this substance could not be the nineteenth century’s
version of matter. In order to get self-repair and reproduction, there must have
emerged new properties. Thus through the remainder of Holism and Evolution
Smuts proceeded to spell out nearly the same position that Edmund D.
Montgomery sketched in Texas a full half century earlier and that Lloyd Morgan
traced in detail in his 1923 Gifford Lectures that became Emergent Evolution .
In his 1931 BAAS Presidential Address, an address to celebrate a century of
scientific achievement, Smuts revisited these themes and placed them in a larger
historical context. Nineteenth-century physicists, he suggested—even including
former BAAS presidents Sir J. J. Thomson and Lord Rutherford—reinforced
belief in a rigid, deterministic universe even as their work began to dissolve the
atomic solidity of its building blocks. In the nineteenth century, their physics had
been spectacular in its breadth and precision, but it was after all “a system of
purified, glorified commonsense.” 15 The more that physicists in the twentieth
century peered into the structure of molecules and atoms, the more they found
that the “commonsense” view had little support. Sir J. J. Thomson himself
confirmed Smuts’s assertions in his own address at the same meeting: Only a
few years earlier, “The view was prevalent that all the fundamental principles of
physics had been discovered”; all that was left was to “measure more and more
accurately.” By 1931, Thomson could join Smuts in asserting that view had
proved to be “ludicrous.” 16 According to Smuts, it was first Einstein’s relativity,
then the revelations of Niels Bohr and others into the paradoxical composition of
light and atoms themselves that so rapidly overturned that commonsense
“picture of nature as consisting of fixed material particles mechanically
interacting with each other.” 17
This shift in the worldview of physics would prove especially liberating for
biologists, thought Smuts. Though for years biology had been held in thrall by
“the partial truth of mechanism,” now the “deeper truth of organicity or holism”
would force physicists to turn to biologists to discover “principles at work in
their full maturity which can only be faintly and fitfully recognized in physics.”
18 One of the chief truths, believed Smuts, could be found only by examining the

organism itself:

A living individual is a physiological whole, in which the parts or organs are but differentiations of
this whole for purposes of greater efficiency, and remain in organic continuity throughout. They are
parts of the individual, and not independent or self-contained units which [sic ] compose the
individual. It is only this conception of the individual as a dynamic organic whole which will make
intelligible the extraordinary unity which characterises the multiplicity of functions in an organism,
the mobile, ever-changing balance and interdependence of the numerous regulatory processes in it,
as well as the operation of all the mechanisms by which organic evolution is brought about. 19

Organisms seemed to show top-down organization, in other words. The whole
organism, when properly configured, really was greater than the sum of its parts.
So important was this new structure of causation, thought Smuts, that it
should restructure our view of the universe itself. Whereas the laws of
thermodynamics stipulated that the vast majority of the space-time mechanism is
tending toward entropy, life moves in the opposite direction—from simple to
complex, bacteria to brachiosaurs. On Earth, mere living, breathing, and eating
gave way to sentience, aesthetics, and eventually organized civilization. The
great irony, as this veteran of wars across Europe and Africa knew all too well,
was that the only shining beacon of complex, introspective organization in the
entire known universe—Homo sapiens —spends an inordinate amount of time
trying to subjugate or kill its own kin. Though he had already helped put in place
the League of Nations to halt some of this violence, Smuts recognized this was
only a stopgap measure. 20 Ultimately, he hoped that a widespread appreciation
for the preciousness of humanity as the most complete instantiation of the
holistic evolution of the universe itself would help curb future proclivities
toward murder, war, and other checks against human flourishing.
Smuts’s pronouncements about the place of the organism in modern science
and philosophy were soaring, and his endorsement of emergent evolution was
well received. This is no surprise: By now, the British scientific community had
heard similar, if less lofty, rhetoric from scholars dating back at least fifty years.
And biologists of Smuts’s generation, including especially E. S. Russell, would
continue to argue similar points for another two decades; J. S. Haldane and C.
Lloyd Morgan, too, repeated these general principles until each passed away,
five years later, in 1936. So Smuts introduced little in his BAAS presidential
address, certainly nothing that had not been said repeatedly over the past half
century. Nevertheless, the value of works like Holism and Evolution and Smuts’s
enhancement of it in 1931 was not in originality—he was riding the crest of a
wave in transatlantic scientific thought in 1931. Smuts marshaled his political
clout to synthesize, popularize, highlight.
He did not, however, add much specificity to the debate. How were scientists
supposed to operate in light of Smuts’s charge? Joseph Needham may have been
convicted by Bukharin and inspired by Smuts. But there were more pieces of the
puzzle yet to fall into place.
 5

WADDINGTON AND THE ORGANIZER

D orothy saw it first. While Joseph was putting the final touches on Chemical
Embryology in 1931, Dorothy—an accomplished biochemist in her own right—
alerted Joseph to the recent developments in German biology she read about.
They appeared to be a logical next step, a biochemical experiment for them to
pursue together as a couple. It was called “Spemann’s organizer” after the man
who had discovered it, German embryologist Hans Spemann (1869–1941). They
contacted a few well-connected colleagues and quickly secured a place to
conduct research near the epicenter of all this work: Otto Mangold’s laboratory
in Berlin-Dahlem. Now the Needhams, too, would pursue the identity of this
“organizer.”
The attempt to describe more precisely Spemann’s organizer became
something of a scientific gold rush in early 1930s European embryology and
biochemistry. The Needhams spent most of the 1930s pursuing this question and
assembled the Theoretical Biology Club in part simply because they needed
other young scientists to bounce ideas off of regarding Spemann’s organizer.
In the months leading up to their departure for Germany, they read a great
deal about the discovery. Both Dorothy and Joseph believed that it could put all
of the philosophical musings of Smuts and Whitehead and the organicists on
display. It seemed to hold the secret to the deep morphological structure of the
organism, somehow configuring the most basic spatial properties of the
developing embryo. Whatever the organizer actually was, it was certainly
biochemical. But its impact was associated with the externally related directional
configuration of the embryo itself, which was something the Needhams didn’t
think could be dictated by isolated biochemicals alone. The context of the
biochemicals—the integrated system of chemical expression, rather than the
chemicals themselves, in other words—seemed to be doing the “organizing.” It
was bigger than mere chemistry; they hoped to say that without invoking vital
forces. 1
 They knew a lot. What they did not know in 1931 was that the organizer did
not belong just to Spemann.

Up to this point, Hilde Pröscholdt (later Mangold; 1898–1924) had thrived in the
male-dominated world of German experimental embryology. But now in her first
year of a doctoral program, she was struggling. Her supervisor, Hans Spemann,
had assigned her a particularly difficult experiment for her dissertation. She
hated to admit it, but despite her notable skill in microsurgery, she could not
carry out his original vision. When she approached Spemann, he reluctantly
handed her a replacement project. Though no one envisioned it at the time, her
second project would earn Spemann the Nobel Prize, permanently alter the field
of embryology, and provide the most direct application of organicism in the first
third of the twentieth century.
The German “Entwicklungsmechanik project” had remained strong since
Roux and Driesch. Arguably, it reached its zenith under the guidance of
Spemann in the interwar period. For decades, Spemann patiently studied the
developmental process of multiple species of amphibians and experimented on
the particulars of their ontogenesis. In one notable experiment, he used a hot
needle to lance cells in the part of the developing tadpole that eventually should
have grown into the frog’s retina. As expected, neither the lens nor the rest of the
eye beneath that lens developed. Spemann then revisited the experiment but
killed only some of the protoretinal cells, leaving a few intact. Tadpoles in that
experiment did form the lens of their eye where the untouched protoretinal cells
came in contact with the top layer of skin cells. 2 This result raised more
questions: Did the optic cup directly instruct the ectoderm to form a lens, or was
contact between the protoretinal cells and the ectoderm only a trigger—perhaps
some eye-making potential already existed in the cells, just waiting to be
unlocked. 3 Future experiments to distinguish between these two possibilities—
which he now called “self-differentiation” versus “induction”—failed to clarify
matters.
Along with Spemann, several researchers across Europe and North America
wondered whether self-differentiation or induction was more likely to be the
cause of organ development. Because of this widespread interest in
Entwicklungsmechanik in general, and because of his skill in uncovering the
processes behind organ formation more specifically, Spemann garnered a great
deal of professional notoriety. Just before the outbreak of the Great War, he was
made head of the Entwicklungsmechanik department at the Kaiser Wilhelm
Institute for Biology in Berlin-Dahlem.
During the course of his lens-induction experiments, Spemann removed
portions of the developing eye from one species and implanted them into the
developing embryos of other species. He also surgically implanted those proto-
eye cells into places in the embryo where they didn’t belong—in the developing
abdomen, for instance. In both cases, the newly introduced material conformed
to the cells of the host: When placed into a gut, the eye tissue would become gut
tissue. At least this is what happened most of the time; there were two
exceptions. The first exception, though mysterious, was not unexpected. If the
transplanted tissue was a little bit older, it seemed to lose the ability to adopt the
qualities of its host. That is, if one transplanted slightly older eye tissue onto a
developing leg, for instance, it would maintain some features of being an eye
even though it was attached to a leg. 4
The second exception proved to be more exciting. One portion of the
developing embryo stubbornly refused to conform to its new location. After
transplantation, the tiny portion of the upper blastopore lip that usually became
part of the developing head and neural system not only didn’t become like the
cells in its host location, it actually co-opted the surrounding cells and began to
turn them into a new neural plate. In 1918, when Spemann first conducted these
experiments, he believed this was evidence of self-differentiation in the
transplanted tissue. 5 But over the next three years, he became increasingly
convinced that something more might be happening.
Just after the war, Spemann earned an appointment to head the Zoological
Institute in Freiburg-im-Breisgau. Soon after arriving in Freiburg, he delivered a
lecture at Frankfurt am Main, where Pröscholdt was majoring in chemistry. His
vision of the promise of Entwicklungsmechanik made such an impression on her
that she changed specializations to zoology. After graduation in 1920, she
enrolled as Spemann’s doctoral student, joining an impressive cohort at Freiburg,
including future luminaries in the field Viktor Hamburger (1900–2001) and
Johannes Holtfreter (1901–1992).
Spemann assigned Hamburger to the laboratory bench adjacent to Pröscholdt;
Holtfreter’s stood nearby. Though he was two years her junior, Hamburger and
Pröscholdt spent a great deal of time together. During their first year, they
enrolled in a wide variety of classes together, including a course in philosophy
with the phenomenologist Husserl. They hiked through the Black Forest, testing
each other’s knowledge of local flora and fauna. They drank in that era’s
efflorescence of culture: They read literature copiously, experienced art, listened
to music, shopped at local markets, and debated the meaning of these things late
into the evening. 6 Contemporaries noted their closeness. 7
Along with her intellectual curiosity, Hamburger appreciated Pröscholdt’s
technical skill. Spemann must have noticed as well, because after she completed
her coursework, he asked her to recapitulate one of the most famous experiments
in the history of biology: Abraham Trembley’s work on hydra. Trembley, the
eighteenth-century mathematics-trained Swiss natural-history tutor, had
reportedly turned at least one hydra completely inside out in the 1730s. He used
little more than a Leeuwenhoek-style single-lens microscope and some crude
tools. Curiously, almost no one had been able to replicate Trembley’s results in
two centuries. But Pröscholdt would have the advantage of twentieth-century
scientific instrumentation. Spemann was very interested in her success—the
scientific question she would pursue was crucial to his own interests. When the
ectoderm cells became endoderm, would they take on typical endodermic
properties? In other words, to what degree did spatial context matter in the
determination of parts of the organism?
Despite her close relationship with Hamburger, Pröscholdt married one of
Spemann’s older laboratory assistants, Otto Mangold, on her birthday in October
1921. Mangold (1891–1962) was Spemann’s first doctoral student and one of
Pröscholdt’s instructors. Though they appear to have had very different
temperaments, Hilde recognized Otto’s professional potential. He was poised to
rise quickly through the ranks of German university science and, in fact, did
attain Spemann’s old chair at the Kaiser Wilhelm Institute of Biology in Berlin-
Dahlem in 1924. But when Pröscholdt ran into trouble with her hydra
experiments, Otto Mangold proved unable to help. Frustrated, she approached
Spemann himself. He used a thin glass needle to restrain the polyp after being
flipped in on itself, but not even this master of microsurgery could successfully
coax it into developing in Trembley’s inside-out configuration. The project that
Spemann had assigned Pröscholdt appeared impossible. So in 1921, Hilde
Pröscholdt—now Hilde Mangold—gave up on hydra and went to work on a
project nearer to Spemann’s own work: interspecific, or “heteroplastic,”
transplantation using newt embryos.
In his postwar experiments, Spemann had used Triturus cristatus and T.
taeniatus newts to investigate the process of induction; they had quickly become
his favored experimental organisms. 8 Common throughout central Europe and
adaptable to captivity, they proved to be ideal organisms for Hilde Mangold’s
transplantation experiments as well. 9 The pigmentation difference between these
otherwise closely related species—T. cristatus shows almost no pigmentation at
all while T. taeniatus cells are somewhat pigmented—meant that donor cells
remained visible among the developing host tissue. Otto Mangold demonstrated
that the pigmentation difference lasted long enough to determine with a high
degree of specificity the permeation of individual grafted cells through the
original organism. 10
Hilde Mangold capitalized on Spemann’s and Otto Mangold’s achievements.
Despite her difficulty recapitulating Trembley’s experiment, Spemann’s
confidence in her microsurgical technique was justified. After delicately teasing
a bit off the upper blastopore lip from a T. cristatus , she transplanted it onto the
flank of a T. taeniatus embryo of the same age, just under the ectoderm. Whether
through skill or beginner’s luck, one of her first few transplants began to fold,
then to invaginate. The transplant then formed a complete neural tube—the
beginning of a cristatus head on the belly of a taeniatus . 11 It was a fascinating
result, to say the least. She continued one experiment after another—259
painstaking transplantations in all, using several different species of newts.
Seventy-three of her transplantations (28 percent) survived; twenty-six of the
surviving chimeras developed the beginnings of a neural plate—a result that
compares favorably to transplantation work conducted as recently as a half
century after her own. 12 In the most exciting instances, the host region
containing the other species’ blastopore lip “bulge[d] out into optical vesicles
and add[ed] lenses and auditory vesicles”—beginning eyes and ears, in other
words. 13
She began to write up her results in the autumn of 1922 as her doctoral thesis.
A year later, Spemann selected six of Hilde Mangold’s best examples and
together they published “Über Induktion von Embryonalanlagen durch
Implantation artfremder Organisatoren” (“Induction of Embryonic Primordia by
Implantation of Organizers from a Different Species”). A decade after he helped
her publish “Induction,” Spemann was in Sweden accepting the Nobel Prize in
Physiology or Medicine. It remains one of the few doctoral dissertations to
contribute directly to the award of a Nobel Prize. Notably, Spemann may never
have been a Nobel laureate at all except that he attached his name to her project
—ahead of hers, in fact. Neither Hamburger nor Holtfreter were involuntarily
made second authors on their own dissertations. 14
Presumably Hilde Mangold would have joined Spemann onstage. The initial
ideas were Spemann’s, certainly, but she performed the delicate experiments that
demonstrated the organizer effect. She would have been in her midthirties at the
time; her son, Christian, would have been eleven. In 1924, Otto Mangold
ascended to Spemann’s old post in Berlin-Dahlem; the Mangolds moved outside
of Berlin that summer, excited to begin life on their own. It was at their new
home on 4 September 1924 that the normally careful, dexterous Hilde
accidentally spilled some cooking fuel on herself while attempting to heat baby
food. The fuel caught fire; the flames quickly engulfed Hilde and Christian.
Though Otto extinguished the flames, Hilde and her child soon succumbed to
their wounds. She was twenty-five years old. Her pathbreaking work on the
organizer appeared in print for the first time later that year. 15
Spemann and Mangold’s “Induction” paper represents another milestone in
the path toward the “third way.” The essay capitalized on two decades’ worth of
work by Spemann, his partners, and assistants. The remarkable results of Hilde’s
experiments convinced Spemann that the blastopore lip contained cells that
convinced the embryonic tissue of any particular region into which it was
transplanted to produce, at minimum, a new brain stem. 16 And with better
surgical techniques and antibiotics to keep chimeras alive longer it became
possible to create a large portion of a “piggyback” organism using these few
cells—a “Siamese twin.” 17
Spemann and Hilde Mangold named this small region of cells in the
blastopore lip the “organizer” region. They believed that somehow this region
set the developing embryo into its most basic spatial properties. In “Induction,”
they prudently admitted that the nature of that organizer and the precise process
by which induction of the neural plate happened remained undefined.
Nevertheless, they were confident “that these secondary embryonic primordia
have somehow been induced by the organizer.” 18 Moreover, these early
organizer experiments seemed to be a strike against the self-differentiation
concept initially championed by Roux. In fact, Spemann and Mangold’s essay
concluded with a measured endorsement of Driesch: “But the development of
the [organizer] implant could not be pure self-differentiation; otherwise it could
not have been harmoniously integrated with the secondary embryonic
primordium which is smaller than the primary primordium. Apparently the
inducing part, while in action, was subjected to a counter-action by the induced
part. Such reciprocal interactions may play a large role, in general, in the
development of harmonious equipotential systems.” 19 “Harmonious
equipotential system”—a phrase Spemann invoked numerous times, even in his
1935 Nobel address—belies Spemann’s philosophical predilections. Spemann
was a vitalist; vitalists could do pathbreaking, Nobel-worthy work in the
interwar period.

In 1931, after a relatively fruitful few weeks in Germany, the Needhams returned
to Cambridge to see what could be done about Spemann’s organizer there.
Frankly, there wasn’t much: Cambridge did not have a dedicated chair in
embryology, let alone a whole working group. And as accomplished as Dorothy
and Joseph were as biochemists, the puzzle of the organizer would definitely
require the perspective of an embryologist to move forward. They had just
overlapped with the perfect addition to their prospective team in Mangold’s
Berlin laboratory—although he admitted embryology was a new pursuit for him.
He was British, a bit younger than the Needhams, and he had trained at
Cambridge. In fact, he still worked in Cambridge, at the nearby Strangeway
Laboratory. His friends called him “Wad.”
Conrad H. “Wad” Waddington (1905–1975) was born to a devout Quaker
family from England’s industrial midlands. His father was a tea planter in India
and Waddington spent his first years there. But this was the extent of his
“traditional upbringing”; when he returned to England, he did so without his
parents, living instead with his grandparents and other family members. He
seldom saw his parents. One distantly related uncle, who Waddington called
“Grandpa Doeg,” took the precocious, relatively unsupervised Waddington
under his wing. A black-cloaked kindly man, Doeg was in fact an independent
scientist who labored continuously to explicate a general theory of all sciences,
from electromagnetism to evolutionary biology. It was from Doeg that
Waddington initially learned the rudiments of biochemistry, botany, and his first
love, paleontology. 20 And it was because of Doeg, no doubt, that Waddington
first considered pursuing a broad, transdisciplinary approach to science that
would mark the whole of his career.
His friends called him “Wad” throughout his scientific career. But when he
was younger, attending Clifton College in Bristol, he went by “Hal.” It sounded
more grown up. By the time he matriculated at Sydney Sussex College,
Cambridge University, in 1923, he wasn’t so concerned with putting on airs of
maturity. He didn’t have to: His appearance and demeanor led many of his
acquaintances to guess that he already was older. He wasn’t a large man—much
closer to Woodger’s size than Needham’s tall lankiness—and he remained spry
and nimble until old age. Those who got to watch him work with his hands were
continually impressed by his dexterity. He read voraciously and retained nearly
everything; he only spoke when he was confident of the truth of what he was
about to say, but he knew so much he inadvertently silenced his peers who knew
less. He took up smoking a pipe while an undergraduate and usually puffed away
in the middle of debates. He rarely lost those. On top of this, he began going
bald while still quite young. To those who did not know him that well, he was
always older , at least older than they were. Though he had many acquaintances
from the sciences, architecture, the fine arts, anthropology, even the political
world, he rarely revealed the wide variety of his interests to any but a few close
friends. To those that he allowed to observe his whole self, he became among
their most treasured comrades. To them, he was always passionate and playful,
always younger . 21
Grandpa Doeg had taught him so well that Wad had already been practicing
geology and chemistry as a serious amateur for several years prior to Cambridge;
he had become something of an expert on ammonite shells, in fact. But his
reason for studying geology had little to do with a desire to do basic research
about marine organisms and much more with pursuing his father’s aspirations: to
earn a respectable living as an Englishman abroad, in petroleum rather than tea.
With his background and motivation, he passed the Natural Sciences Tripos in
1926, earning a first-class degree in geology with, if we believe his recollection,
an absolute minimum of preparation. Instead of preparing for his Tripos
examination—and instead of training for a career in petroleum—Waddington
was busy reading philosophy. 22
Smuts’s Holism and Evolution and the challenging philosophy of Alfred
North Whitehead had a profound effect on Waddington, just as it had on
Needham and Woodger. He was likely first introduced to the organic philosophy
by members of the Department of Zoology’s teatime discussion club, which
included Joseph Needham, future Yale University ecologist Evelyn Hutchinson,
and lifelong friend Gregory Bateson (the last surviving son of William Bateson).
They continued their philosophical discussions even after Waddington received
his BS in geology in 1926; Waddington bought a house and moved in with his
new wife, artist Lass Lascelles. The two became well known for their open-door
hospitality; their house was often filled with curious scientists standing cheek-to-
jowl with avant-garde painters. It was Gregory Bateson who lured Waddington
away from geology and toward genetics—at the very moment Bateson was
leaving his father’s genetics to study anthropology—during late-night
conversations while fossil hunting on the chalky Dorsetshire cliffs. 23 Soon
Waddington met Edith R. Saunders, a family friend of the Batesons and longtime
collaborator with William Bateson and R. C. Punnett at the John Innes
Horticultural Institute. 24 As a first-rate geneticist herself, Saunders quickly
recognized Waddington’s potential and put him to work on a genetic question
that had interested her for some time: the problem of seemingly maladaptive
factors in the garden flower Matthiola incana , better known as “stock.”
Waddington went straight at it and conquered the problem quickly, publishing
his results in the prestigious Journal of Genetics . 25
Despite his early professional success in plant genetics and geology,
Waddington longed for a scientific life devoted to those large theoretical issues
he had been discussing with Bateson, among others. 26 Reading Whitehead and
Smuts inspired Waddington to compete for the prestigious Arnold Gerstenberg
Prize in Philosophy in 1929. He titled his winning essay “Philosophy and
Biology”; it revealed the philosophy that would lead Waddington to his
groundbreaking discoveries of “genetic assimilation” and his widely used
metaphor, the “epigenetic landscape,” decades later.

In the essay, he drew explicitly from Whitehead, positing that the

fundamental constituents of the world are “four dimensional events.” 27 This
meant, for Waddington, that what we commonly discriminate as objects are in
reality “possibilities of realisation of certain qualities, which are realized when
the objects have ingression into events.” But as this process of ingression into
events is dependent on perception, it may therefore be “delusive.” Perceived
objects that must be objectified are open to a plurality of interpretations
determined as much by “imagination” as anything else. 28 When applied to
biology, Waddington believed that Whiteheadian emphasis on time, process, and
perception rendered any nontrivial statements about the behavior of the smallest
parts of organisms provisional at best. Then Waddington extended this
skepticism to the growing consensus about genetics and evolution: “Supposing
that at any time we become possessed of a reasonably complete scientific
explanation of the process of evolution, as that it is brought about by natural
selection acting on gene mutations, for instance; we shall not then be justified in
taking this as a complete general explanation of evolution. The scientific
explanation of evolution, as of everything else, is a particular set of relations
embedded in the general fact which it is explaining, whereas a complete account
of the matter must transcend the general fact.” 29 The “natural selection acting on
gene mutations” version of evolution could never be a “complete general
explanation.” 30 According to Waddington’s reading of Whitehead, biological
evolution had to be more interconnected than this.
Skeptical as he was of the consensus position regarding genetics and
evolution, Waddington transformed again, this time into an embryologist.
Through his friend, physicist Sidney Cox, Waddington contacted Dame Honor
Bridget Fell to inquire about his chances of working in her lab. Fell had just
become director of the Strangeways Laboratory in Cambridge in 1928.
Strangeways itself had only recently converted from a hospital to a laboratory
dedicated to solving problems of “normal and abnormal growth,” including the
impact of radiation on cell development. 31 Waddington wanted to study growth
and development as an instantiation of the organic philosophy. As he told Fell,
his interest in development had been sparked by Spemann’s organizer project;
Waddington wondered if Strangeways had the ability to replicate Spemann’s
remarkable results but using endothermic animals like birds and mammals
instead of amphibians. No one in Britain had undertaken such a challenging
experiment. Surprisingly, Fell gave Waddington the go-ahead—even though
Waddington was presented to her as a “paleontologist,” not as an embryologist,
biochemist, or geneticist. She even assisted him in applying for a senior research
grant from the Department of Scientific and Industrial Research (DSIR). 32 By
1930, Waddington found surprising success in this area as well; to the
astonishment of his colleagues, this twenty-five-year-old geologist was able to
publish the results of his experimental developmental work—work that the
Strangeways laboratory founders themselves were unable to successfully
complete—in Nature . 33 Meanwhile, Waddington collaborated with J. B. S.
Haldane, son of J. S. Haldane, on a highly technical problem of linkage in
interbreeding of lines. That coauthored paper appeared in Genetics in 1931. 34
Waddington’s accumulation of successes in areas outside of his formal
Cambridge training, especially in his ability to work with avian embryos,
brought him to the attention of Berlin luminaries Richard Goldschmidt and Otto
Mangold (the latter now a widower). At their encouragement, Waddington
traveled to the Kaiser Wilhelm Institute for Biology in Berlin in 1931 for a six-
month appointment with the Entwicklungsmechanik group. 35 When they later
arrived in Berlin, Joseph and Dorothy Needham became Waddington’s
neighbors. In the fall, all three of them returned to Cambridge and Strangeways
Laboratories flush with ideas. Waddington prepared to take up the
Demonstratorship in Experimental Zoology offered by new chair of Cambridge
Zoology, James Gray, in combination with a concurrent Medical Research
Council fellowship offered by Sir Walter Fletcher. 36 Now he had a substantial
laboratory project to pursue as well—the nature of the organizer.
The collaboration between Joseph and Dorothy, biochemists, and
Waddington, the geneticist/embryologist, on the concept of the organizer proved
to be some of the most fruitful and frustrating work of their careers. The 1930s
became a highlight of their personal and professional lives. Their organizer work
would be a watershed moment in the history of developmental biology, genetics,
and evolution, and another milestone in the history of the “third way.”
 6

THE ORIGINAL THEORETICAL BIOLOGY CLUB

S ecure or insecure, established or peripatetic, from Oxford or Cambridge or

London, they gathered together as transgressors. “The Theoretical Biology
Club” was the name Joseph Needham attached to their meetings. That name may
have sounded impressive in the 1930s, but this was not the group’s formal
designation. Some called it a “biotheoretical gathering”; still others,
“Woodgery,” in recognition of the ostensible captain of the ship. Indeed, the
group typically did meet at the Woodger’s summer cabin on Epsom Downs,
south of London. But the scribbled notes of their meetings belie an egalitarian
working group, a near free-for-all, an intellectual smorgasbord. At Theoretical
Biology Club (TBC) meetings, they shoved disciplinary distinctions,
professional training, and the usual trappings of academic hierarchy—gender,
class, and above all educational pedigree—to the background in favor of
smoothing out the rough edges of the “third way.”
The whole thing was Joseph Needham’s idea, though Woodger put the plan in
motion in April 1932. Now that Needham was taking organic philosophy
seriously, albeit with dialectical materialism mixed in, and now that Spemann’s
organizer was before them—a real interdisciplinary challenge—the time was
ripe to search for an alternative to both mechanism and vitalism again. An
intensive discussion by committed experts seemed like the best way to parse
these much-debated ideas, to dig for solutions, to extend the resultant thinking
into the sociopolitical realm. Waddington was a logical fourth member, after
Dorothy Needham. Woodger suggested they also invite E. S. Russell and Guy C.
Robson, a zoologist at London’s distinguished Natural History Museum who
studied the conundrum presented by speciation in evolution. 1 Needham thought
J. D. Bernal would be a natural fit, along with physicist Lancelot L. “Lana”
Whyte, who had been part of the symposium at the SICHST in 1931 and was
just then attempting to formulate a unified field theory (that he would later
haggle over with Albert Einstein). 2 They bandied other names about, but
Woodger was most excited about a mathematician, Dorothy Maud Wrinch
(1894–1976).
“Dot” Wrinch—who signed her letters in the 1930s with just a Greek delta (δ)
—had already made a name for herself as the first woman to earn a DSc from the
Faculty of Physical Sciences at Oxford University. She was also philosophically
minded, having served as Bertrand Russell’s personal secretary in 1918 while he
was locked away in Brixton Prison as a conscientious objector to the Great War.
She befriended morphologist D’Arcy W. Thompson soon after the 1918 BAAS
—the legendary meeting during which Thompson and John Scott Haldane hotly
debated mechanism. In 1931, perhaps at Woodger’s urging—they knew each
other from Aristotelian Society meetings—Wrinch travelled to Przibram’s
Vivarium in Vienna. There she learned, just as Woodger had a few years earlier,
that mathematical precision in biology did not automatically entail mechanism.
Meetings with Kurt Gödel, author of the famous incompleteness theorem, and
Karl Menger, one of the young Turks of mathematical topology in a different
Vienna circle, enthralled her. 3
Back in England, life was less invigorating. Her husband John Wrinch, also a
mathematician, had just been institutionalized after a “very serious breakdown.”
4 They divorced shortly thereafter. So Wrinch very sincerely was “charmed to be

a member of the team” when Needham and Woodger invited her to join their
Theoretical Biology Club. 5 Given her interdisciplinary interests, she proved an
invaluable core member of the TBC, even though ostensibly her attraction to
biology seemed to run counter to the interests of Woodger and others.
Wrinch wanted to name the entities of life, to define their structures, not
necessarily to sketch the network of interactions, nor to apply Whiteheadian
philosophy of organism to biology. Still, Wrinch had studied with Bertrand
Russell after he and Whitehead had labored at the Principia Mathematica. That
experience left her interested in the relationships between organisms and their
constituent parts. Russell convinced his students, including Wrinch, of the
importance of his “theory of types.” According to this principle, in order for
sentences to be meaningful, they must be arranged hierarchically. At the lowest
level, stipulated Russell, sentences are about individuals. But then above that
level there is another level in the hierarchy, sentences about sets of individuals.
Above that level, there are sets of sets of individuals, then sets of sets of sets ,
and so on. Firmly holding to Russell’s theory of types was the only way to avoid
the dreaded “paradox” he identified, wherein a set that is not a set containing
itself must contain itself.
Russell classically used barbers to illustrate this difficult paradox. Imagine a
town full of barbers who shave only those men who do not shave themselves.
This is their set: barbers-who-shave-only-men-who-do-not-shave-themselves.
Among the barbers in this town, among that set, there is a misfit barber—a
barber who does not shave himself. Paradoxically, given the definition of his set
as barbers-who-shave-only-men-who-do-not-shave-themselves, he must shave
himself. But no barber in the collection, in that set, can shave himself because
they are barbers-who-shave-only-men-who-do-not-shave-themselves. (If so, he
would be a man who does shave men who shave themselves and, therefore,
would not belong in the set to which he already belongs.) It is a brain-twisting
problem.
This paradox is highly significant even when extended beyond barbers. In his
Introduction to Mathematical Philosophy , Russell put it this way:

The comprehensive class we are considering, which is to embrace everything, must embrace itself
as one of its members. In other words, if there is such a thing as “everything,” then, “everything” is
something, and is a member of the class “everything.” But normally a class is not a member of
itself. Mankind, for example, is not a man. Form now the assemblage of all classes which are not
members of themselves. This is a class: is it a member of itself or not? If it is, it is one of those
classes that are not members of themselves, i.e., it is not a member of itself. If it is not, it is not one
of those classes that are not members of themselves, i.e., it is a member of itself. Thus of the two
hypotheses—that it is, and that it is not, a member of itself—each implies its contradictory. 6

Wrinch knew from studying with Russell—and Woodger stressed it repeatedly in
TBC meetings—that biological sentences, i.e., sentences about living things,
must also accord with these principles. Russell’s paradox might be a problem in
logic, but it extends to anything, any scientific explanation that has to do with
classification and with explaining something higher (i.e., sets) by referencing
only lower things (i.e., individuals in sets). By extension, behaviors operating at
a lower level—at the genetic level, for instance—could not by themselves dictate
what happens at a higher level (e.g., at the organism level). They are in different
levels. It was Woodger’s intention to find the logical language for expressing
how these levels might interact, how they could be logically constructed so that
genetic explanations could be properly, rather than sloppily, applied. He hoped
Wrinch’s expertise and new interest in topological mathematics would help him.

For the next six years, the group of five core members—Joseph Needham,
Woodger, Waddington, Bernal, and Wrinch—met over a weekend or longer once
or twice per year. As many as eight other philosophers and scientists joined them
at any one time, occasionally for just one meeting, usually for more than one,
often with spouses or partners or lab assistants in tow. The heart of each meeting
beat to the wide-ranging discussions of exceedingly complex theoretical issues
in physics, chemistry, biology, and mathematics—usually all four areas in
combination with each other. As Joseph Needham’s scribbled notes record,
experts presented their topics semiformally, presuming that attendees would
have a good enough grasp of the subject matter to move forward into a deeper
theoretical problem. 7 Problems , not disciplines , defined them. Woodger
typically led the long discussions that followed each presentation. By this point
in his career, colleagues had dubbed him “Socrates,” given his penchant to
question and re-question issues that many simply had stopped questioning, Peers
found this trait of Woodger’s useful and charming—unless his questioning was
directed at them. At the TBC, his Socratic method meant that few presentations
got by without a thorough cross-examination.
Because of the interests of the attendees of this first meeting in 1932,
geometrical patterning in biology appeared in nearly all of the presentations. For
instance, J. D. Bernal, who was then working in crystallography, proposed a
scale of bodily form that included everything from the level of quantum
structures to metazoa. He suggested that a mathematical formula must exist to
describe these morphological transitions—something like D’Arcy W. Thompson
had described during the Great War in his classic On Growth and Form . 8 That
observation opened directly into the philosophical question: Was there, then, a
hard mechanistic determinism to biological form? The physicochemical limits to
liquid transport in an enclosed envelope of skin cells seemed to stipulate that
only a few basic configurations were possible to any organism. This of course
provided a constraint on evolution; not just any kind of organism was possible.
But Thompson’s On Growth and Form seemed to defend something stronger.
Life was not merely constrained by physical barriers but actually determined by
them. Though we might accept that evolution could not be analogous to
traveling on horseback across an open plain, with any direction as likely to be
taken as any other, could we say that it was like traveling by car over a wide
highway? Or was evolution more like train travel—with deviation from a set
path bringing a swift and terrible end to the journey? Dot Wrinch then spoke of
“geometrical botany” and the mathematical complexity of morphological forms.
Before the meeting in 1932, she had already been in discussions with
Waddington regarding possible topological models of embryological
differentiation rates. 9 Waddington turned out to be one of the most active
participants in the first meeting. Aside from the work he and Wrinch had begun,
he presented once on new findings in Drosophila genetics and again raised the
issue of chemical equilibrium during development. Waddington had begun
thinking about the existence of homeotic mutations in Drosophila , which was
the basis of his important Aristapedia experiments years later. 10
The TBC met regularly through the 1930s. 11 Initially, attendees assembled at
the Woodgers’ cottage in Epsom Downs, but this spot became cramped as the
meetings grew larger. By 1936, attendance by regular members was slated to be
more than a dozen. Partners and other casual attendees would swell that number.
Joseph Needham took it upon himself to find a larger meeting location. He
contacted his friends, poet Frances Crofts (née Darwin) Cornford and professor
of ancient philosophy Francis Macdonald Cornford; the Cornfords owned a
cottage attached to a repurposed windmill near Old Hunstanton, Norfolk, in sight
of the North Sea. They agreed. Thus the 1936 and ’37 meetings took place under
the rough-hewn timber ceiling of the round brick room at the base of the
windmill.
In these relaxed settings, attendees almost immediately chiseled through the
customary walls between their academic and personal lives. Politics found its
way in; religion too—first during their nights camping under the stars in the
Woodgers’ back garden, later over meals and strolls along the windy Hunstanton
sea cliffs. Over the course of years of interactions, something more intimate
developed between them.
In the social network forged by these social, political, and philosophical
outsiders, the organic philosophy finally came into its own. The TBC
conspirators applied themselves to one of the most difficult problems in all of
biology, a problem Aristotle himself noted and that the newly microscope-armed
biologists of the seventeenth, eighteenth, and nineteenth centuries wrangled
over: How can biological complexity be generated out of such simplicity? This
question is the classic conundrum faced by Rudolf Virchow, who gave his partial
answer with the phrase omni cellula e cellulum (all cells from other cells). This
question awed Driesch into vitalism; Bergson, too. TBC members, by contrast,
did not allow themselves to fall back on the explanation of vis viva , entelechy,
or vital fluids. Nor were they satisfied with the consensus explanation that the
prior condition of the fertilized egg and its resultant genetic configuration—the
more mechanistic explanation—provided a definitive answer.

That is not to say the members of the TBC denied the importance of genetics
in development. They fully appreciated, perhaps Waddington more so than the
others, that new combinations of alleles might come to exist in a reconfigured set
of chromosomes after fertilization. But those genes alone did not do more than
limit the set of trajectories or paths for development. The actual construction of
the organism—the way that one relatively undifferentiated cell gives rise to
myriad cells, each with a particular function in a vast matrix of other specialized
cells—requires both more information and more physical material than appears
in the chromosomes of the lone fertilized egg. In other words, biological
development must follow rules that are not derived specifically from the pieces
playing the game themselves. They must obey Russell’s theory of types or risk
being dragged into the paradox. Woodger attempted to formulate a new
definition of biology that would respect this logic:

The science which deals with entities or systems of entities which are capable of repeating
themselves in space. (This covers bacteria, protista, chromosomes, genes, budding of cellular parts
and whole of various orders, etc. etc. At the chem[ical] level this would mean (I suppose) chiefly
polymerization and some sort of [illeg.] or segregation). It is because of this that many-one
relations are so common in biology, and it is because of this that hierarchical (as opposed to merely
serial) order is important in biology.
What are the various types and modes of spatial repetition? What must happen to the parts
(components of various orders) when a complex system repeats itself (e.g., cell division (so
called)). Think this over! 12

Both the material for making the organism and the information for how to
construct that material into an organism must come from outside of the solitary
egg. But how can one use concepts like “outside information” without invoking
references to entities outside of, or at least undetectable within, the
physicochemical system in which science operates? In other words, how can
those serious about the extreme underlying complexity of development possibly
avoid some version of vitalism?

Warren Weaver of the Rockefeller Foundation called these TBC members “queer
fish” after meeting some of them in the mid-1930s. 13 They certainly did seem
like outsiders. All were relatively insecure professionally, perhaps especially
because of their nontraditional social, political, and religious backgrounds.
While Dorothy and Joseph Needham had some security at Cambridge University
working under Sir Hopkins, they were nevertheless also deeply embedded
outsiders as part of the Thaxted Christian socialist movement. Unbeknownst to
them, they were also being continually monitored by British intelligence for
being sympathetic to international Communism. 14 Waddington and Woodger,
too, were Fabian Marxists who retained interest in social forms of nontraditional
Christianity but, unlike the Needhams, neither had supportive supervisors or
prestigious positions from which to challenge the consensus in their fields.
Bernal, too, was in a precarious position in the 1930s, given his ardent support of
socialism; despite his talent and wide influence on other successful British
scientists, he was never able to crack into the more prestigious British academic
research circles. 15 Wrinch was probably even further from the mainstream in
terms of social position and professional security, this despite her DSc from
Oxford and her lengthy publication record. It was perhaps this feature more than
anything—their status as outsiders, a collection of outsiders in fact: the set of
those-who-feel-constantly-pushed-to-the-margins—that made the TBC an
incubator of ideas and an intellectual and sociopolitical touchstone for its first
members and, consequently, for the organic philosophy.
It certainly was a stimulating time for Waddington and the Needhams. In
mid-1933, the three of them made another run at the organizer problem from
Otto Mangold’s Berlin-Dahlem lab. It was a strange summer for them, full of
mixed emotions. They saw Adolf Hitler quickly translate nationalist popular
sentiment into an authoritarian regime and, by the close of their trip to Berlin,
were frightened by the gradual tightening of the noose against any dissent. On
the other hand, their scientific work proved to be both more promising and more
mysterious than ever.
That summer they demonstrated for the first time that the organizer effect,
which Waddington by then labeled “evocation,” could be produced by nonliving
tissue. 16 That convinced the Needhams and Waddington of two things. First,
Spemann’s vitalistic explanations of Hilde Pröscholdt’s experiments—the
experiments that would win Spemann the Nobel Prize two years later—were not
right. 17 Secondly, while Germany in general and Berlin in particular had long
been among the best places in the world for developmental biology since the
Entwicklungsmechanic of Roux and Driesch, it would no longer be a friendly
place for scientists with left-leaning political sentiments like themselves. They
understood as they returned to Britain that a great opportunity now lay before
them. As Germany became more restrictive, they had the chance to organize the
foremost laboratory of developmental biology in Europe. In fact, they now
required such a laboratory because their most recent organizer work presented an
unexpected puzzle: How could nonliving tissue instigate embryonic
development? They agreed to invite participants to the TBC from outside the
initial group to help address what had become a far more complex issue.
By the mid-1930s, the reputation of the TBC had begun to spread, not least
because of the scientific work being done by its core members. The Needhams’
collaboration with Waddington on Spemann’s organizer continued unabated after
their return from Berlin. Now the organizer was among the hottest topics in
European biology, attracting talented young researchers like Jean Brachet, and
Johannes Holtfreter, friend and lab mate of the late Hilde Mangold, away from
Germany and to Cambridge. Woodger’s collaboration with Viennese biologist
Ludwig von Bertalanffy (1901–1971) had transformed into Modern Theories of
Development (1933) and was making waves in theoretical biology on the
Continent. 18 Wrinch sketched a physical model of the chromosome at a 1934
TBC meeting that morphed into a brief letter to Nature and then an important
article, “On the Molecular Structure of Chromosomes.” 19 When untangled and
unspooled, the chromosome, Wrinch speculated, appeared as a cloth mat: The
proteins stretched out in strips, nucleic acid threads woven between, over then
under then over. The proteins, being more complex molecules, would carry all of
the information, the specificity that made traits with the nuclein just holding the
mesh together; nucleic acids were mostly inert, a backbone. 20 The model was
nothing like the one made by Watson and Crick a decade and a half later, but it
anticipated the code metaphor so popular today—the kinds of proteins in the
chromosome aggregate and specify traits in the adult organism. 21 Bernal and
Crowfoot Hodgkin, too, were increasingly bringing x-ray crystallography of
biological chemicals into the scientific spotlight.
Drawn perhaps by the growing reputations of its members, Viennese
philosopher Karl Popper (1902–1994) attended the TBC in May 1935, held at
the Woodgers’. 22 His groundbreaking Logik der Forschung had just been
published and, ostensibly, Popper visited England as a young, already
accomplished philosopher, introducing Britain to sophisticated Viennese logical
empiricism. In truth, he had seen the writing on the wall and was hoping
desperately that his book would provide him with a way out of increasingly anti-
Semitic Austria. 23 The topic during the 1935 TBC meeting was “the nature of
‘organizing relations,’ mechanists and ‘over simplified hypotheses,’ the ‘alogical
core of the world.’” For a Continental intellectual, it was a cornucopia. And the
TBC made a lasting impression on Popper. He left England without securing
employment, but he thoroughly enjoyed the pastiche of mathematics, biology,
philosophy, and the easy way in which these polymaths were able to switch back
and forth between topics. 24 When he returned to Britain in 1936 at the behest of
Susan Stebbing, A. J. Ayer, and Joseph Woodger, Popper insisted on meeting
with the TBC again. As he recalled decades later, this was the first meeting held
at the “‘old windmill at Hunstanton.’” 25


As it turned out, the June 1936 meeting in the Cornfords’ windmill would be
the largest and most important in the history of the original TBC. Core members
Waddington, Joseph Needham (it is unclear whether Dorothy joined him),
Woodger, and Bernal attended. J. B. S. Haldane and mathematician Hyman Levy
joined Popper as special guests. As usual, a great deal of conversation circulated
around mechanism and anti-mechanism in biology—a topic perhaps made more
piquant as they slept under the interlocking gears of the old brick mill. But now,
with darkening skies over Europe and war again springing up in east Asia, the
discussion turned more toward Marxism and the widespread organization of
scientific work, by scientists but for the benefit of the common man. Bernal was
especially vocal: His new partner Margaret Emilia Gardener (1904–2005)
headed the left-wing “for intellectual liberty” group in the Bloomsbury area of
London. Gardener and Bernal had visited the USSR together in 1934, and
together they agitated for leftist causes ever more aggressively. Despite his
earlier interest in socialism, Karl Popper would later recall that he had disagreed
with the entire tenor of the meeting. Though none of the members recalled a
negative reaction from Popper during the meeting itself, his Poverty of
Historicism , published two decades after this meeting, attacked those leftist
political and scientific philosophers he met in 1936 at the old windmill. 26
Popper’s retrospective criticisms notwithstanding, the 1936 meeting at the
windmill represented the apogee of the TBC before the Second World War, not
only because of the number and prominence of the attendees but because of the
way in which the meeting continued to inspire participants in their individual
academic and professional pursuits, their sociopolitical leanings, and even their
personal lives years later. It may have been the moment when the organicist
vision was the clearest, the prospects for the future of science and society the
most promising, the network of scientists interested in theoretical biological
issues the most tightly knit.
Needham attempted to capture something of the spirit of the TBC in those
first few years, and the other relationships that spun off from the meetings
themselves, in his Terry Lectures at Yale University, published as Order and Life
(1936). He dedicated the book to eight members of the group, arranged on the
dedication page as if clustered around an imaginary circular table: the Woodgers
at the twelve o’clock and six o’clock positions, then clockwise: Wrinch, Bernal,
Dorothy Moyle Needham, B. P. Wiesner, Max Black, and Waddington. Just as
they had done during TBC meetings, Needham began the book with a discussion
of the resolution of the mechanism and vitalism debate. A close examination of
Order and Life reveals just how much TBC discussions had changed him
personally. “I myself . . . held opinions which, though very different from the
vitalist ones . . . led to the same conclusions,” he confessed. “I regarded the
nature of biological organization as a purely philosophical question, and
excluded it from scientific biology.” 27 This is true: Prior to 1931, Needham had
sealed off organicism from any actual impact on biology. In 1930, for instance,
he had proclaimed, “The concept of organism, then, is best regarded as a
philosophical concept proper to the domain of the philosophy of science, but in
no sense a scientific hypothesis. . . . The mechanistic principles of practical
research would have continued in use whatever philosophers said, but now there
is no philosophical reason why they should not. The mechanistic schema
formerly covered the non-living world not inadequately, but now without
hesitation is extended to cover the living world as well.” 28 In contrast, by the
mid-1930s Needham was “glad to have an opportunity of cancelling what I then
said.” 29 Paraphrasing Woodger, Needham would now be happy to advocate
“‘legitimate’ organicism.” As he summarized it, that meant identifying three
possibilities in the relationship of an individual biological system to a whole
organism: (1) The system is independent of the organism and, therefore, could be
studied without any fear that something vital—entelechy or élan vital—would be
lost; (2) the system is functionally dependent on the whole organism; (3) the
system is existentially dependent on the whole organism. The hope, according to
Needham—and in this he summarized the ultimate project of the TBC in its pre–
Second World War days—was to concentrate on the first two types of
relationships, analyze them as far as possible without concerns about
overreaching mechanism. And even in those situations where a system was fully,
existentially dependent upon the organism as an irreducible unit—the
phenomena that vitalists would have used to support their claims—they would
continue to analyze. They might look like mechanists in practice, in other words.
Yet their aim was not to explain complex phenomena through reductionistic
appeals to physics and chemistry. According to Needham, they recognized that
the oversimplifications of mechanism were just as damaging to science as the
“dogmatic” or “metaphysical” prohibitions on scientific analysis put forth by the
vitalists.
At its high mark in the 1930s, biology’s “third way”—robust organicism—
could be identified in a number of exemplars: the axiomatic language of
Woodger, the biochemical research of the Needhams, the experimental
embryology of Waddington, the topography of Wrinch, the x-ray crystallography
of Bernal and Crowfoot Hodgkin, and the engineering of Lana Whyte. And it
could be found in the work of other scholars elsewhere in Europe, scholars who
agreed that “the true task of scientific research is not the violent identification of
the biological and the physical, but the discovery of the qualitatively specific
controlling principles which characterize the principle features of every given
phenomenon, and the finding of methods of research appropriate to the
phenomenon of study. . . . Affirming the unity of the universe and the qualitative
multiformity of its expression . . . , it is necessary to renounce both the
simplified reduction of some sciences to others, and the sharp demarcation
between the physical, biological, and socio-historical sciences.” 30 Or, according
to another, “The unity of the universe expresses itself in qualitatively different
forms, the characters proper to which must not be lost sight of.” 31 The organic,
as these “third way” advocates saw it, was not “impenetrable by the human mind
or ruled by unintelligible spiritual entities. Translated into terms of Marxist
philosophy, it is a new dialectical level.” 32
 7

LARGE PLANS VERSUS THE ULTIMATE

LITTLENESS OF THINGS

I n April 1934, W. E. Tisdale of the Rockefeller Foundation (RF) approached

Joseph Needham. Tisdale had heard of the new work being done on Spemann’s
organizer at Cambridge and sensed a great deal of promise in the cooperative,
multidisciplinary attack on a “borderline subject.” Was there a way the
foundation could advance these experiments in “chemical embryology”? 1
Indeed, Needham had a grand vision. He wanted to take the fruitful
collaborations at TBC meetings and give them a permanent academic home with
much more funding and institutional support. It looked as if the RF might be
willing to fund a laboratory wherein the organic philosophy might finally
become instantiated in concrete, mainstream science. Sensing his chance,
Needham needed only two months to submit his plan for a fully staffed and
outfitted Research Institute of General Embryology.
We do not know the true motivations behind Tisdale’s offer to fund the work
of Joseph Needham and, by extension, Dorothy Needham and Waddington.
There are hints that Needham’s Chemical Embryology interested the Rockefeller
Foundation because it suggested a truly mechanistic explanation could be found
for development and morphology. This fit the RF’s preference for funding
projects interested in the “ultimate littleness of things.” 2 And it did accurately, if
only partially, describe Joseph Needham’s own philosophy of neo-mechanism
prior to the influence of Whitehead, Bukharin, Smuts, Woodger, and others in
the early 1930s. But by the time Tisdale approached him, Needham’s tune had
changed. In the end, the negotiations between Needham and the Rockefeller
Foundation would not only fail to secure an institutional home for scientists to
pursue the organic philosophy in the laboratory, they would sharpen into a
wedge that, along with other factors, drove the TBC members in different
directions. Eventually, the acrimony would lead Waddington away from
Cambridge, Needham away from biology altogether, Wrinch to the United
States, and Woodger toward relationships almost exclusively with philosophers
who showed less interest in biology, including Karl Popper, Rudi Carnap, Alfred
Tarski, W. V. O. Quine, and Freddie Ayer. Needham’s grand plan of
institutionalized multidisciplinary organicist biology at Cambridge would never
come to pass.

Needham’s full proposal in 1934 asked for the RF to fund seven research
sections to be added to current Cambridge biological facilities:

(1) Physico-chemical embryology, or biochemistry and biophysics [this is where Needham put the
organizer work];
(2) Experimental or causal embryology or morphology, the Entwicklungsmechanik section;
(3) Genetical embryology or physiological genetics. The study of the action of the genes during
development . . .;
(4) “Explantation” [tissue culture];
(5) “Descriptive morphological embryology” esp. of invertebrates;
(6) Physiological embryology. The later stages of functional differentiation are usually neglected by
embryologists . . . questions such as the physiology of the placenta . . .;
(7) “Psychological” embryology [reflexology];
(8) Theoretical embryology.

Needham then listed possible directors for each section; he nominated all the
main members of the TBC and several leading biologists familiar with the
organic philosophy. For instance, as director of the Theoretical Embryology
section, Needham nominated Woodger, with Nicolas Rashevsky, a recent
Rockefeller Fellow then at the University of Chicago, as a possible backup.
Waddington, he insisted, was “the only active and successful investigator in
Entwicklungsmechanik in the country.” Dame Fell, he suspected, could be
poached from Strangeways to lead the “Explantation” section. Among “Other
Europeans” to consider for inclusion as head of Genetical Embryology,
Needham preferred “Dobjanski.” (By this, he meant Russian geneticist
Theodosius Dobzhansky (1900–1975), who would soon join T. H. Morgan at
Caltech to begin the Drosophila work that would make Dobzhansky a pillar of
the Modern Synthesis). Under no circumstances should the Rockefeller promote
instead of those on his list Needham’s erstwhile comrade Julian Huxley who,
though he had talent as an embryologist, was too often tempted to forsake
laboratory work for “popular presentations and . . . public speaking in general.” 3
It was an ambitious request—too ambitious for the RF. It asked for something a
little more manageable.
Needham’s second proposal in February 1935 requested only five
departments: Chemical Embryology, Experimental Morphology, Crystal Physics
of Biological Compounds, Physico-chemical Cytology, and Theoretical Biology.
He called the whole project “An Institute for Mathematico-Physico-Chemical
[MPC] Morphology” and, just as with his earlier proposal, Needham placed one
of the TBC core members at the helm of each division. 4 This interested the RF
enough that director Warren Weaver traveled to Cambridge to meet Joseph
Needham personally. On 5 May 1935, they discussed prospects for a “large
plan” that, if properly executed, should gradually introduce the law-bound
rigorousness of physics and chemistry to previously unruly fields like
biochemistry and embryology. 5 After that first meeting, Weaver left ready to
throw his weight behind it.
Over the next few days, however, his opinion soured. Though duly impressed
with Needham’s ability to move between biological subdisciplines and with the
work that he, Dorothy, and Waddington had done on Spemann’s organizer with a
modicum of institutional support, Weaver and his other advisors concluded that
the field of chemical embryology was too “young,” Needham’s proposal still in a
“developing” stage. 6 Why the rapid change of assessment? Unbeknownst to
Needham, Warren and Tisdale had spent the next day or two after their initial
meeting perusing the facilities in Cambridge’s biochemistry department and
conversing with other university officials. Their investigations turned up too
little institutional support for Needham and Waddington’s unorthodox attempts
to bridge embryology and biochemistry. 7 Warren and Tisdale also may have
caught wind of the rumor—quite true, as it turns out—that the Needhams and
Bernal, especially, were suspected of Communist sympathies and were being
spied upon by their own government. 8 The RF quietly backed away from the
project.
Nevertheless, there did seem to be enough exciting science going on that the
RF kept up communications with Needham about the possibility of a
“Biochemical Laboratory Extension.” 9 It was not transdisciplinary, but it was
something. Also, Weaver and Tisdale did not hesitate to fund individual research
trips for Needham and Waddington to visit the United States in 1936, and they
continued to show interest in the work of the other TBC members. But Needham
desperately wanted to find an institutional niche for the organizer work. He
pressed Tisdale and the Rockefeller over 1936 and 1937. Wouldn’t they fund
something larger, more permanent, something like the TBC, oriented around
problems instead of disciplines? Unfortunately, the more he pressed, the more
resistance he encountered. Only gradually did Needham realize the ultimate
source of that resistance did not emanate from the RF itself; it was a far more
local problem.
A series of quotidian matters led to the dissolution of the organizer project
and contributed to the breakup of the original TBC. For instance, a small conflict
over Waddington’s salary—a misunderstanding really—grew until it ended up
redirecting the course of Waddington’s scientific career. Beginning in 1933,
Waddington held a research fellowship at Christ’s College that paid £300
annually; £200 more came with the title “Part-Time Lecturer” in Zoology. And,
on account of his philosophical prowess demonstrated by his 1929 essay on the
application of Whitehead’s philosophy to biology, he was awarded the Royal
Commissioners of the Exhibition of 1851 Senior Studentship. The studentship
initially granted him £300 a year, but was reduced to £100 in 1936. While £600
may have been a comfortable salary for a young scientist in 1936 (roughly
£50,000 in 2015), Waddington had a fixed alimony payment of around £300—
leaving him with a mere £250 after taxes. He came to Tisdale more or less hat in
hand looking for some funding to make up the difference. 10 But RF policy
mandated that only departmental personnel at universities could make salary
requests. 11 Waddington was housed in the zoology department headed by James
Gray. Gray did not particularly care for Waddington or his science. 12 When he
learned that Waddington, with Needham’s support, went directly to the
Rockefeller for salary assistance, Gray took offense and went to the Cambridge
Council. The council asked Needham to explain himself. “I understand,”
Needham wrote Tisdale in December, “ . . . a slight verbal slip . . . led the
University to think that the Trustees did not properly appreciate Mr.
Waddington’s position as a University part-time lecturer, but as Mr.
Waddington’s position is the same now as it has been for the last six years and as
we do not desire it to be altered . . . this misunderstanding will be cleared up.” 13
Whatever it was, the Cambridge Council did not look upon Needham’s “verbal
slip” kindly. After some directed consultation, the council and general board
changed their policy to mandate still more direct oversight: “We have reason to
believe that the Rockefeller Foundation approves the suggestion that proposals
for grants to increase the stipends of University Officers should be referred to the
Council and to the General Board.” 14 Though it was touch and go for a while,
Waddington found other sources of income, permitting him to stay in Cambridge
and to continue work at Strangeways Laboratories. 15 But he began to look for
opportunities elsewhere.
Understandably, this episode led to bad feelings between Needham and
Waddington on the one side and Cambridge University officials on the other.
Certainly the hints that Needham and his accomplices were allied with
Communism did little to earn sympathy or financial support from Cambridge
administrators. By 1938, Tisdale revealed to Needham that even Hopkins’s
support was waning. What’s more, with Hopkins nearing retirement, Tisdale
warned Needham that Hopkins’s successor would likely cut off Needham
entirely. 16 Personally disappointed, and faced with an almost total lack of
funding in the immediate future, Needham confronted his longtime mentor.
Hopkins rejected Tisdale’s suggestion as “absurd”: The RF had only consulted
with Hopkins “once or twice,” and he was entirely out of the “loop” on the
Waddington salary dispute. He vowed to approach Tisdale and Weaver directly
and to be more vocal in his support of “your work on the Organiser.” 17 Little
improved, however. By September 1938, with the funding situation ever more
worrisome, Needham approached the Royal Society for funding; Hopkins wrote
in support, though he was “practically sure the R[ockefeller] F[oundation] will
make things right.” 18 In his appeal to the Royal Society, Needham attempted to
convey the precariousness of his and Waddington’s situation at Cambridge. One
can sense his exasperation:

If the grant is not obtained: Nothing further on the nature of the primary organiser will be done in
Cambridge, though Cambridge has at present the lead in this subject. If it is not done in Cambridge,
there is no chance of its being done anywhere else in England. . . . It is conceivable that the work
might then be proceeded with either in Brussels (Brachet), Bern (Lehmann), New York (Barth), or
Amsterdam (Woerdemann). . . . On the whole, it may be said that the subject would be held up for
10 to 20 years. . . . it seems truly shortsighted, especially in a country where belief in pure science
has always been boasted, not to recognise the value of research into the deepest processes of early
embryonic development. 19

This appeal to British pride worked, partially: The Royal Society did see fit to
fund Needham, but not to the sum he had hoped. Permanent damage had been
done. Waddington was forced to transition his research fully to Strangeways
under Dame Fell’s biophysics grant. 20 Cambridge failed to support experimental
embryology consistently after this point.
The relationship with the RF itself also suffered. Needham and Waddington
had been so successful in their pursuit of Spemann’s organiser in the early 1930s
that Weaver felt compelled to invest in the work straddling Hopkins’s
biochemistry lab and the Strangeways Laboratory through the mid-1930s. 21 At
one point the RF funded lab technicians to work with Needham and Waddington
at Cambridge, as the university would not. 22 Yet the succession of failed
attempts at identifying the specific biochemical substance through the 1930s was
not the solution the RF had hoped for, even if it corroborated Waddington and
the Needhams’ suppositions. 23 By 1938, Waddington published his seventh
study “On the Nature of the Amphibian Organization Centre,” subtitled
“Evocation by Some Further Chemical Compounds,” which reported that
synthetic—in some cases toxic—substances nevertheless induced the embryo to
symmetrically develop just as any other “natural” compound. To Waddington,
such a result indicated that the “evocator . . . appears impossible to discover”
given some of the ruling assumptions that “the artificial [laboratory] process
gives a true picture of what happens in nature.” 24 He introduced the speculation
he and Needham shared that, rather than a biochemical substance that induced
change, the evocation principle was distributed throughout a field or gradient in
the developing embryo but suppressed. Once the suppressor was released by
adding the synthetic chemical compound, the embryo began the developmental
process. Johannes Holtfreter—who worked alongside Waddington and Needham
in 1939–40—later confirmed this experimental result, showing that even a
below-normal pH liquid could trigger neural induction. 25 They called this
process the “masked evocator.” Years later, Waddington noted that the masked
evocator concept was “logically almost isomorphic with the non-instructional
theory of enzyme induction in bacteria advanced by Jacob and Monod.” 26 Had
they written in a different era, perhaps the Needhams and Waddington would be
lauded as the discoverers of regulatory genetics, albeit of a much more organic
variety.
While still an exciting and active field of research, experimental embryology
at Cambridge was no longer going to be supported by the RF without requisite
support from the university itself. 27 As Needham explained to Finnish
embryologist Sulo Toivonen many years later, they had not abandoned their
experimental pursuits because their “enthusiasm died.” The funding and, perhaps
more importantly, institutional support simply dried up. “[O]ur research . . . had
to be carried out in the interstices,” Needham recalled. Had he, Dorothy, and
Waddington received “some reasonable encouragement” from Cambridge, the
history of the “third way” may have turned out quite differently. 28

Spemann’s organizer experiments had been a kind of proving grounds for the
organic philosophy. Paradoxically, Waddington’s demonstration that even lethal
substances could initiate neural induction meant that the organic philosophy
passed this preliminary test: No discrete gear was needed to turn on the
mechanism of development. The organizer must be a problem of wholes, a field
distribution, an organism-wide phenomenon, a dialectic between organism and
environment. But this result meant that there were now new questions to ask.
How could something like an organic field be created and distributed? Was it
similar to the concept of field in physics, as Smuts believed, and as TBC-regular
Lana L. Whyte would argue in his lauded book The Unitary Principle in Physics
and Biology (1949)? What did genetics have to do with the development of this
field? And if there was some connection between the field’s development,
genetics, and the physical structure of the molecules involved, as everyone
assumed there was, how could this be expressed while continuing to privilege
relationships and process over particles?
While the TBC would have been a productive environment in which to tackle
these new questions, there were practical matters that precluded their continued
meeting. Most notable, of course, was Cambridge’s failure to support the
organizer work any further. But just as important were the newly divergent
interests of the members themselves—their own idiosyncratic attempts to make a
living while pursing the organic philosophy. The Needhams, J. D. Bernal and
Dorothy Crowfoot Hodgkin, Waddington, Woodger, and Wrinch—each would
peel off in different professional directions by the end of the 1930s.
Woodger did not change professions, but he did move further into analytic
philosophy, toward the logicism of Alfred Tarski and issues of axiomatization.
He spent increasingly large amounts of time with philosophers at the London
School of Economics, the institution to which Friedrich von Hayek lured Karl
Popper. In 1937, Woodger finally published his first attempt at building a system
of biology on the basis of the abstract logical symbolism found in Whitehead
and Russell’s Principia Mathematica. He named it The Axiomatic Method in
Biology , and it acquired a kind of following of its own—but from terribly few
working biologists. 29 Woodger successfully relaunched the TBC after the
Second World War, but with the exception of Popper, ironically, most of the new
members were less acquainted with the “third way” and related topics that
occupied the prewar TBC discussions.
Bernal would forever be a biophysicist and soon accepted a professorship at
Birkbeck College, London, where he would famously pursue greater political
activism and his fascination with crystallization. Rosalind Franklin came to
Bernal’s laboratory at Birkbeck in the mid-1950s and found it a much more
welcoming environment after her negative experiences with Maurice Wilkins,
Jim Watson, and Francis Crick in the early 1950s. 30
Like Bernal, Dorothy Wrinch persisted in her heterodoxy, transgressing the
physics-mathematics-biology boundary. She moved from Oxford to Johns
Hopkins and then to Smith College in the United States, where she would live
for the rest of her life as a molecular biologist. 31 Inspired by the TBC, she
famously debated the molecular structure of protein with Linus Pauling, never
being as sure that the relationship between protein and nuclein was quite as
simple as he and others believed it was. She appealed to large, complex
repeating patterns in crystals that defy easy explanation. Dot lost that battle in
the mid-twentieth century. There are now signs that she was on the right track,
however, and that the pendulum may have swung back in Wrinch’s direction. 32
Though neither Joseph nor Dorothy Needham left England, they had a
profound redirection nonetheless. In 1937, three Chinese scientists—Shen
Shizhang, Wang Yinglai, and Lu Gwei-djen—visited F. G. Hopkins’s
biochemical laboratory. Both Needhams greeted the contingent; Joseph was so
moved by the meeting and by his experience working alongside them that
summer that he immersed himself in the study of Chinese language and culture.
That decision ended up redefining Needham’s entire life. He still considered
himself a biochemist—he completed Biochemical Embryology , possibly his
most important scientific work, in 1940. He was still thoroughly committed to
the organic philosophy and Whiteheadian process. He still attended Noel’s
radical Thaxted church and meetings of the Cambridge Communist Party, and
even delivered impassioned messages at each. But Joseph would also become
the preeminent historian of Chinese science and technology in Britain; this
became his enduring legacy rather than biochemistry or theoretical biology. This
is what made Needham into “the man who loved China.” 33
While the RF pulled back from broader engagement with Needham’s grand
interdisciplinary plans at Cambridge, Tisdale saw fit to offer Waddington a travel
grant to pursue the connection between genetics and embryology elsewhere. In
the summer of 1938, Waddington landed at the Marine Biological Laboratory at
Woods Hole, Massachusetts, United States. But after only a few weeks, he felt
the working environment was too slow and socially stifling. 34 He hoped to
spend time closer to the heart of American genetics, first at Columbia University
in the presence of the original “Fly Room,” and then in Thomas Hunt Morgan’s
new and improved Drosophila lab at Caltech in Pasadena. Over the 1938–39
academic year, he set to work on thorny problems in Drosophila morphology
alongside Morgan’s protégés Alfred Sturtevant and Theodosius Dobzhansky. 35
Waddington gladly incorporated the hands-on genetics instruction into his
background in experimental embryology. Still, his year of work in the United
States reinforced his belief that American geneticists did not know what to do
with the problems of development that embryologists encountered as a matter of
course. They were chauvinistic also—not nearly as willing to learn European
embryology from Waddington as he was willing to learn American genetics. 36
Stimulated by T. H. Morgan’s attempt to synthesize the two main areas
investigating heredity and development in Embryology and Genetics (1934), and
challenged by Dobzhansky’s recently published neo-Darwinian milestone
Genetics and the Origin of Species (1937), Waddington chiseled away at his own
synthesis, Organisers and Genes (1940). 37 The book, the most significant of his
early career, summarized the organizer work and went beyond it. With the
encouragement of Gregory Bateson and Needham, Waddington incorporated his
cutting-edge embryological work into the increasingly powerful and persuasive
American genetic idiom. 38 It was an ambitious and creative project.
Nevertheless, Waddington perceived resistance from the Anglo-American
genetics community all the way through its writing and publication.
Like T. H. Morgan himself, Waddington regarded the science of “genes” as a
means to a greater end—understanding the formal development of complex
living creatures from two haploid zygotes. 39 Genes themselves were heuristic
devices, “thing” names for the invisible networks and processes that actually
produced the traits. Geneticists searching for the isolable gene would invariably
be disappointed because genes as discrete entities could only be properly
understood in this more regulative (rather than constitutive) manner. Genes had a
role to play, a supremely important one in fact, but as complexes rather than
individuals or even sets of individuals. An organ or system inside the organism
was formed by the genotype collectively forming “a sequence of changes” that
Waddington termed “the ‘epigenetic path.’” Those paths tend to be “‘canalized,’
or protected by threshold reactions so that if the development is mildly disturbed
it nevertheless tends to regulate back to the normal end-result.” 40 Genes do not
“code for” a particular trait, according to Waddington. Together, the collective
genome and the developing cellular matrix creates probabilities or potentialities
of trait configurations that become phenotypes. The environment might disturb
normal development. Normal development is buffered against certain kinds of
disturbances; the organism is likely to end up in one of a few configurations. It
was a sophisticated view offered by a talented, philosophically informed
geneticist-embryologist.
But the fact that Waddington could be so circumspect, so non-laudatory, so
hesitant about genes as determiners of development—instead he focused on
epigenetic paths and threshold reactions creating multiple possible phenotypes
from the same genotype—meant Organisers and Genes was placed under the
same shadow as Morgan’s Embryology and Genetics written a few years earlier.
41 No larger rapprochement between developmental biology and genetics would
be forthcoming for nearly the remainder of the twentieth century. 42
 8

AS MANY OPINIONS AS THERE ARE MEN

B y the time Waddington returned to Cambridge from Morgan’s Fly Room in

1938, a war against fascism loomed just over the horizon. Bergson and Walter
Marvin, among others, had framed the First World War as a struggle between the
philosophies of mechanism and vitalism. Would a similar interpretation be
drawn from this new conflict? The First World War postponed or reshaped the
plans of many researchers in the UK—Woodger, for instance. If another world
war broke out, would the young scientists of the 1930s have any control over the
fate of their research, or would they find themselves marching, huddling in
trenches, sampling intestinal bacteria in dun-colored tents?
With core members moving in different directions, and with the organizer
collaboration between the Needhams and Waddington on hold, possibly
permanently, the original TBC did not reconvene. 1 Bernal, Joseph Needham,
Lana Whyte, and Waddington joined another group, this one specifically focused
on sociopolitical issues, especially the scientific ordering of Britain to face the
coming conflict. It was dubbed “Tots & Quots”—a phrase borrowed from the
second-century-BC Roman playwright Terance’s “Quot homines tot sententiae”
(“As many opinions/statements as there are men”). The group ended up being
unexpectedly influential for the future of British science and even the outcome
of the Second World War. Historians would dub them Churchill’s “tame
magicians of the War.” 2
Indeed the war pushed even these champions of organicism away from their
previous preoccupation with theoretical biology. Tots & Quots focused neither
on theoretical issues nor specifically on biology; it encouraged activism and
public planning. Concerned that Japanese brutality would hamper Chinese
science, Needham went to China as part of the first Sino-British scientific
outreach. Waddington joined Patrick Blackett in Operations Research (OR) work
with the RAF; his OR group was instrumental in the Battle of the Atlantic.
Unable to pursue laboratory biology, he published widely read and debated
publications connecting evolution and ethics. Both Needham and Waddington
joined other Tots & Quots members (also former TBC members) to write the
anonymously published Science in War , which lobbied both for specific changes
to scientific and agricultural policy and for the British scientific and political
elite to join in support of long-term national projects.
To be sure, important theoretical biology work continued for a time. Woodger
introduced the German notion Bauplan while addressing the problem of form
and organization in his essay for D’Arcy W. Thompson’s Festschrift . 3
Waddington published two crucial articles and a groundbreaking book between
1938 and the beginning of his OR work. Joseph and Dorothy Needham, Bernal,
and Wrinch, by contrast, contributed to theoretical biology much less during and
after the war. Personal issues—the dissolution of the collaborative network in
particular—contributed to the marginalization of organicism and epigenetics in
the mid-twentieth century in combination with the theoretical issues discussed in
later chapters. Understandably, even supporters were more concerned with the
war than with debates in theoretical biology. During their relative absence,
however, other philosophical narratives less congenial to the complexities of
“third way” thinking would emerge, especially in the United States.

Oxford primatologist Sir Solly Zuckerman (1904–1993) created a short-lived

invitation-only Tots & Quots in the early 1930s, around the same time Needham
and Woodger convened the TBC. 4 In 1939, Zuckerman restarted Tots & Quots
with a new goal: to collect “people with different interests to discuss common
problems” and to “influence” those attendees through “non sterile” debate. 5 He
brought with him Julian Huxley, now a radio host of The Brains Trust and
secretary of the London Zoo. Huxley and Zuckerman had just produced a
documentary film on the behavior of apes. Zuckerman and Huxley successfully
recruited other scientific luminaries with political connections, including
controversial cell biologist Cyril Darlington, mathematician Hyman Levy, soil
chemist William K. Slater, marine biologist and neurologist John Z. Young, and
science journalist James C. Crowther. (Several of these Tots & Quots comrades
had attended TBC meetings over the 1930s.) The ostensible goal of Tots &
Quots may have been like any other drink-dine-and-discuss club, but
Zuckerman’s real purpose was to connect British scientists with political figures
who could put their science to work for the nation and the world. Careful
minutes of their meetings list a litany of British, American, Canadian, French,
and even Soviet political, military, scientific, and literary notables, including H.
G. Wells, N. M. Victor Rothschild, Royal Society chair A. V. Hill, Warren
Weaver, Harvard president James Conant, US ambassador John Gilbert Winant,
and Admiral Harold R. Stark, Commander of US Naval Forces in Europe.
Bernal set the tone of the group in the first meeting on 23 November 1939.
Most of the Tots & Quots had attended the fateful 1931 SICHST meeting and
had listened intently to the Soviet contingent as they exposed the deficiencies of
capitalism. While Needham found Bukharin the most inspirational, Bernal was
moved most by Soviet State Planning Commission (GOSPLAN) economist
Rubenstein’s “Relations of Science, Technology, and Economics Under
Capitalism”; he based his lecture on Rubenstein’s implication of the waste,
corruption, and inefficiency generated by the capitalist system. Rubenstein’s
point was that waste—in his case, the unemployment of talented scientific minds
—was the intentional by-product of the plutocrats to keep demand at a level
where it benefited the few who controlled the means of production rather than
the many who actually labored. If a system existed whereby this waste could be
reversed, where people could be employed according to their talents, then both
higher production and greater contentment among the workers would result.
Bernal lamented in 1939 that the British government had not even attempted to
analyze this situation in the intervening eight years, let alone do anything about
it. If they could actually know what was needed, maybe they could plan for how
best to distribute the existing know-how, to get the most out of the technical and
intellectual talent available without exploiting that talent.
Bernal repeated Rubenstein’s other criticism: His audience of scientists spent
too much of their time in the “metaphysical spaces above the clouds.” 6
Rubenstein had challenged the British scientists gathered at the SICHST to
renounce the traditional division between “pure” and “applied” science, to work
toward innovations that would harness the natural world in service of humans,
not merely to understand the composition of that world. Some no doubt balked at
Rubenstein’s suggestion that civil engineering should stand at the same level
with quantum physics or agricultural production with biochemistry. A
preoccupation with commodities and practical solutions was an obsession of
American academics, not British ones. But Bernal had taken the Soviet critique
at the SICHST to heart. Though many of those assembled excelled at the “pure”
discovery aspect of science—the TBC was a group created to debate exactly this
—Bernal hoped that their future aim should be to serve the people through
innovations, thereby connecting techno-science to the working class.
Following Bernal’s charge at the first meeting, one of the main objectives of
Tots & Quots scientists became the creation of a “scientific general council” to
advise Prime Minister Winston Churchill’s cabinet. In the meantime, they
authored pamphlets for broad distribution that would educate the public and
government officials through “factual statement and with suggestions from the
scientific point of view.” 7 In this way, the Tots & Quots scholars could render
even their most densely philosophical work more practical, less “metaphysical.”
There was also the added benefit of contributing to the centralization of
scientific planning for the betterment of all classes, not just the plutocracy.
Three publications emerged from these initial discussions on the eve of the
Second World War. Levy, the mathematician, issued an essay entitled Science:
Curse or Blessing? (1940), the most direct reinterpretation of Bukharin and
Rubenstein. 8 Zuckerman and Needham followed Levy’s effort by compiling
eight essays from core Tots & Quots members into the anonymously authored
Science in War (1940), a call for broader ordering of research according to the
needs of the people rather than the whims of funding sources. 9 Waddington
started with his anonymous Science in War submission but then went beyond it
in a third publication, The Scientific Attitude (1941). 10
He formulated Scientific Attitude in dialogue with Robert Thouless, a lecturer
in education trained in psychology and occasional attendee of the Cambridge
Moral Science Club in 1940 and ’41. Thouless had a practice of walking and
talking philosophy with Ludwig Wittgenstein after dinner in the Roundabout
Garden at Trinity College. Waddington frequently joined them in the summer
evenings early in the war. 11 Their conversations sometimes devolved into
monologues as Wittgenstein attempted in his characteristically halting way to
define distinctions between private and public languages, grammatical versus
factual statements, and the practical impossibility of translating thoughts into
universally transferrable axiomatic formalisms. Despite the fact that he deeply
desired biology to be more self-consciously philosophical, Waddington had
trouble envisioning the larger relevance of Wittgenstein’s kind of analytical
philosophy—especially during the Blitz. Instead, these evening chats actually
helped crystalize Waddington’s motivation for incorporating theory into his
science: Even philosophical excursions should end up addressing social
problems rather than playing what he took to be “word games.” 12 The Scientific
Attitude , then, may have been Waddington’s admittedly somewhat hasty attempt
to articulate a scientist’s philosophy in the face of the Wittgensteinian trend
growing at Cambridge.
As it turned out, Scientific Attitude was only the first of Waddington’s
publications to deal with his lifelong interest in what might be called the
anthropology of ethics. His more thoughtful Science and Ethics would appear in
1942, capturing a controversy that engaged philosophers, scientists, clergy, and
journalists across Britain. That controversy began with a brief statement
Waddington originally published in Nature . 13

We must accept the direction of evolution as good simply because it is good according to any realist
definition of that concept. We defined ethical principles as actual psychological compulsions
derived from the experience of the nature of society; we stated that the nature of society is such
that, in general, it develops in a certain direction; then the ethical principles which mediate the
motion in that direction are in fact those adopted by that society. Of course the good is, as the
anthropologists pointed out, different in different societies, and particular cultures which regress
may be actuated by principles at variance with the cosmic process. But in the world as a whole, the
real good cannot be other than that which has been effective, namely that which is exemplified in
the course of evolution. 14

By the “good,” Waddington meant the apparent increase in complexity of
individuals, groups, and the diversity evident in whole ecosystems that seemed
to be the overall trajectory of evolution on a global, if not universal, scale.
Science could judge a culture as ethical or progressive if the individuals in that
culture comported to the increase in complexity and, therefore, diversity,
unethical or regressive if they advocated those things that lead to
oversimplification and the breakdown of diversity.
Karin Stephen, a psychoanalyst who had written the well-received Misuse of
Mind —a nuanced defense of Bergson—challenged Waddington on his
assumption of an evolutionary “good.” 15 Waddington had identified a holistic
superego as the basis of ethics. Stephen pointed out that Waddington must be
mistaken, since the superego of Freudian psychoanalysis could only be guilt
inducing, regressive, even evil. Instead, she suggested that there existed some
unnamed “spontaneous tendency” toward maturity, mental and moral health—in
short, goodness. 16 This “tendency” she characterized as a kind of élan vital
running counter to both the superego and to evolution, which she defined as a
brutal life-destroying process, along the lines of T. H. Huxley’s depiction of it in
his famous lecture Evolution and Ethics (1893).
Waddington responded in several ways. First, he appealed to the labors of his
friends Margaret Mead and Gregory Bateson in support of the concept that
“good” in Stephen’s sense was not good in an evolutionary sense. Her use of
good as “mature” was merely culturally relative: “[M]ature and well-balanced
personalities are in point of fact dependent on the culture in which they live. The
anthropologists Ruth Benedict, in her Patterns of Culture , Margaret Mead in Sex
and Temperament and Bateson in Naven have all pointed out that the type of
personality who feels well-adjusted and happy is not the same in different
societies.” 17 Mead’s and Bateson’s fieldwork suggested that radically different
kinds of personalities could be “mature and well balanced” in their cultures. This
culturally dependent idea of good could not be what Waddington meant by
“good.” So, secondly, Waddington took on Stephen’s suggestive question about
this very issue. Did Waddington mean that evolution as a process could be
working toward our “mature personalities” in a teleological sense? “I rather
dislike speaking of evolution having a goal. I should wish to say that we observe
an evolutionary change of which we can specify certain characteristics (to take a
random selection—an increase in size, in complexity of structure, of subtlety and
precision of movement, of capacity to react to the relations between objects). A
tendency for change in this sense has then in point of fact been operative in the
organic world.” 18 Because of this built-in increase in complexity in the
evolutionary process, Waddington dismissed Stephen’s appeal to Bergsonian
vitalism as unnecessary both in the biological and in the psychological realms. In
phrases like “change in this sense has . . . been operative in the organic world,”
Waddington was merely summarizing a non-vitalistic “third way.”
Waddington encountered an entirely different challenge from biologist
Miriam Rothschild, sister of Victor, 3rd Baron Rothschild. While Stephen’s
psychoanalysis represented the Scylla of vitalism, Rothschild’s biology
personified the Charybdis of mechanism. She concluded her objection to both
Waddington’s organicism and Stephen’s vitalism with a call to acknowledge the
“hereditary element in our ethical codes.” Drawing from examples about the
behavior of birds observed by ethologists including Niko Tinbergen, Rothschild
wondered, “Can science cope with the fact that, mentally, man has raced ahead
of his genes?” Somehow humans had managed to come up with altruistic
“golden rules.” These would be self-defeating because, she asserted, human
ethical standards were due entirely to “hereditary determination,” which was
more or less self-centered, given natural selection. 19 Though self-identifying as
a geneticist by this time, Waddington remained unconvinced by Rothschild’s
appeal “to the importance of the hereditary determination of our ethical
standards.” “We actually know very little about the causal mechanisms of
behaviour in birds, and the mere invocation of a genetic basis whose expression
cannot be modified by the environment seems somewhat too easy. . . . The
reference to Black-headed Gulls [Rothschild’s example, borrowing from
Tinbergen] is in fact not only a long range and dubious analogy, but is an
analogy with something which we understand even less fully than the human
behavior which is supposed to be illuminated by it.” 20 Still, Waddington agreed
that, references to birds aside, genes certainly shape (not determine) behavior.
The question was to what degree could genes in particular be thought to
specify discrete traits ? According to Waddington’s “theory,” which he did not
spell out in much detail here, genes were not the proper agents. “But on my
theory one is bound to admit that the genetic nature of mankind sets limits to the
range of human ethical performance, just as does that of chimpanzee to his. But
that does not in the slightest imply that differences within the range of normal
human behavior are also genetically determined. That would only follow if we
supposed that the environmental differences met with among men have only
negligible effects on the development of the different genotypes.” 21 In other
words, Waddington supported the concept that human culture and behavior
flowed like a river between wide, steep banks. According to this metaphor,
genetics determined the landscape. But the actual flow of culture and behavior
could not be specified by genetics. Waddington’s claim here was bold: We could
assign genetic constituents for particular behaviors if we assumed environmental
differences contributed little to the “development of the different genotypes,” but
because we suppose that the environment has greater than “only negligible
effects” on development, it must follow that the particulars of human culture and
behavior are outside of direct genetic control and require a fairly large
environmental contribution.
The view of human culture that Waddington defended against Stephen’s
vitalism and Rothschild’s mechanism emerged quite directly from two sources:
his familiarity with the anthropological work of his friends Gregory Bateson and
Margaret Mead and from the TBC discussions. Largely because of these
influences, Waddington would gradually infuse his version of the “third way”
with a sense of proper behavior, an ethic of living organically. But the full
realization of what this meant would be delayed for decades by the war and a
host of other life circumstances.

Though Science and Ethics earned Waddington the limelight, his most important
contributions to theoretical biology in the early 1940s came in the form of two
brief articles, one in Nature and the other in the popular scientific journal
Endeavour. After the MPC morphology confusion, the salary debacle, and the
disappointing conclusion to the organizer experiments, Waddington had been
quietly chipping away at one of the problems at the core of evolutionary biology.
In his Nature article, “Canalization of Development and the Inheritance of
Acquired Characters,” he tackled a question that dated back to Aristotle: How do
organisms change from one form to another while retaining efficient
coordination with their environmental niches? 22 He stressed that neither of the
options then considered by evolutionary biologists—neo-Lamarckism and neo-
Darwinism—offered satisfactory accounts. But why, then, were both accounts so
attractive?
Naturalists and systematists—those biologists who dealt with whole
organisms and their lineages—relied upon neo-Lamarckism well into the
twentieth century. That wasn’t because of intellectual laziness; Waddington
thought there was a much more likely reason. The version of Lamarck’s theory
formulated by American paleontologists in the late nineteenth century accounted
for the observed phenomena: As environments change, organisms respond,
occasionally dramatically, and their progeny seem to exhibit the effects of that
response. The strict Darwinian interpretation insisted that change would be
random with respect to the environment and gradual, almost imperceptible. As
paleontologists insisted in the early twentieth century, the responses of
organisms seemed neither random nor slow but tightly coordinated to
environmental pressures and rapid, at least geologically. Nevertheless, as
Mendelian breeding experiments and new cytological observations emerged,
those biologists who worked in laboratories insisted that the mechanics of
inheritance were not Lamarckian. No matter what the paleontological record and
the studies of field naturalists might suggest, the environmental context of one
generation could not directly impact future generations.
Waddington disagreed that the modern theory that synthesized Mendelism
and Darwinism solved the problem that the neo-Lamarckians identified. By the
1940s, the modern, or neo-Darwinian, synthesis was the dominant interpretation
of evolution in part because its pioneers, R. A. Fisher and J. B. S. Haldane,
incorporated mathematical models in biology. This made biology feel a good
deal more scientific, like physics. But Waddington pointed out that, despite its
popularity, it relied too heavily on imagined stochastic processes. Neo-
Darwinians, in other words, postulated a “random walk” of genes and selection
—genes change any which way and are expressed in organisms under the
assumption that selection eliminates unsuitable individuals and, therefore, the
number of aberrant genes in the population. However, Waddington asserted,
when they applied their quantitative models to real organisms, not even the
“most statistically minded geneticists” seemed wholly convinced by this system
of mutate-and-filter. Take, for example, callouses on the underside of ostriches.
(Waddington borrowed this example from Robson and Richards’s The Variation
of Animals in Nature [1936]—a source also cited by both Theodosius
Dobzhansky and Sewall Wright in their famous neo-Darwinian accounts.)
Ostrich callosities seem to be intragenerational adaptive characters. This means
that, though callouses are caused by friction on the underside of the ostrich as it
kneels onto hot sand and rocks, callouses mysteriously develop before birth, that
is, before any stimulus that would cause the skin to thicken. Field naturalists in
the early twentieth century were tempted to see those callouses as examples of
the inheritance of characteristics acquired during the adult ostriches’ lifetimes—
support for Lamarckism. But by the 1930s, most laboratory biologists regarded
this explanation as a nineteenth-century fiction. Robson and Richards then
entertained the neo-Darwinian explanation: A randomly mutating gene must
have caused these callouses. But how could a randomly mutating gene be
selected for in utero, as proponents of the modern synthesis had to assert? 23
Waddington attempted to cut through the dilemma: “[S]uppose that in earlier
members of the evolutionary chain, the callosities were formed as responses to
external friction, but that during the course of evolution the environmental
stimulus has been superseded by an internal genetic factor.” 24 But how could
this be true? Wouldn’t one have to believe that somehow the environment caused
genetic change—a Lamarckian hypothesis? In order to cut the knot, Waddington
made two smaller claims. First, he asserted that the possibility of response in the
organism had to be under genetic control. This was not a controversial idea from
a neo-Darwinian perspective. To this Waddington added a slightly more
unconventional postulate: When actually “submitted to natural selection,”
organisms did not develop haphazardly. In fact, there were only a very small
range of phenotypic variations for any given species, and this was true even in
variable environmental conditions. Developmental responses—meaning the
changes that an organism could undergo from fertilization through the end of its
growth phases—tended to persist in a particular configuration despite the
environment. Development is, in Waddington’s nomenclature, “canalized,” that
is, restricted, nonrandom, following one or another pre-generated path. 25
Canalization is not a difficult concept. By 1942, it was not even new. Many
biologists, even after Darwin’s On the Origin of Species , observed the relative
imperturbability of organismal development and form and assumed that taxa
were essentially static categories. Sure, small variations might lead to new
species , but why should we assume the differences between genera, families,
orders, and so on are all acquired the same way? Evolutionists of whatever stripe
thought that, given the right environmental context, change would come either
through the direct pressure of the environment or through the expression of some
gene that a new environmental context allowed to flourish and spread. But the
process of development belied such easy assurances. A canid develops in a dog-
like fashion in just about any environment. Butterflies morph through four
distinct developmental stages but almost always end up butterflies. Escherichia
coli and Drosophila can be made to develop in just about any conceivable
configuration in the laboratory; in an ordinary environment, however, they tend
to follow predictable developmental patterns unique to their genera and even
species. By “canalization,” Waddington merely asked his readers to take that
appearance of equilibrium seriously—rather than assuming it to be illusory, a
figment of our brief life spans, as gradualistic evolutionists since Lamarck have
tended to do. As Waddington saw it, “constancy” in the phenotypes of
nonlaboratory creatures “must be taken as evidence of the buffering of the
genotype against minor variations not only in the environment in which the
animals developed but also in its genetic make-up. That is to say, the genotype
can, as it were, absorb a certain amount of its own variation without exhibiting
any alteration in development. . . . [I]t is not difficult to see its advantages, since
it ensures the production of the normal, that is, optimal, type in the face of the
unavoidable hazards of existence.” 26
For Waddington, canalization also accounted for the immediate, or
intragenerational, fitness that so lured an older generation to neo-Lamarckism.
Look again at the strange case of ostrich sternal and alar callosities, which arise
long before the animal is subjected to environmental stimulus. Long ago, these
callosities arose only with the appropriate environmental stimuli, only after
repeated contact with harsh surfaces. But as time went on, the threshold of skin
reactivity lowered considerably to the extent that almost no environmental
stimulus would be needed to produce the callosities. Over time, ostriches that
developed callouses faster under ever-slighter stimulus would outreproduce their
kin, until, one day, a new pathway appeared during the course of development.
After this point, the ostriches would develop callosities in the appropriate places
without any environmental stimulus. Even if reared in a soft nest—even if it was
born into an area without rough sand and rocks—an ostrich chick would
nevertheless emerge from its egg already possessing sternal and alar callosities.
Crucially, however, the environmental stimulus–biological response system
did not close at that point, becoming forever fixed in the phenotype by virtue of
a change in the genotype. Canalization meant that things could change again,
could return to the old process of development or even to a new one not before
seen. This is where Waddington parted ways with the neo-Darwinism of his
peers. Subjected to “extreme and specialized” environmental conditions at
precise moments in its growth from egg to adult, the organism might suddenly
switch all at once from one canalized developmental path to another. On that
new path, assuming it provided selective advantages and the intense and unusual
stimulus continued for several generations, the organism would settle into a
different standardized developmental pathway, producing an adult—the new
“normal”—buffered against change by the usual whims of the environment.
Only then, suggested Waddington—only after an environmentally buffered, new
normal developmental pathway was established—would the genome catch up.
By then, of course, the favorable traits would be produced even without the
original environmental stimulus. 27 Thus, according to Waddington’s account, a
once-acquired characteristic would assimilate seamlessly into the development
of a new phenotype, largely through hereditary mechanics familiar to
evolutionary biologists—and certainly without appeals to the direct action of
either environmental forces or internal orthogenetic drives.
It garnered little attention during the war. Nevertheless, “Canalization of
Development and the Inheritance of Acquired Characters” deftly blended
genetics derived from T. H. Morgan’s Drosophila lab, embryology from Otto
Mangold’s Berlin lab, and the philosophy of Whitehead as channeled through the
TBC. The term “canalize” itself Waddington appropriated directly from
Whitehead. And the notion that genetics works in potentialities while
development works in actualities came straight from his organizer experiments
with the Needhams. Moreover, the article briefly highlighted for a more
specialized audience the powerful metaphor he had introduced in Organisers and
Genes two years earlier: development as a landscape, a river delta, a possibility
space.
On the frontispiece of Organisers and Genes , Waddington had tasked his
friend John Piper (1903–1992), a modern artist who traveled in the same circles
as Waddington’s new wife, architect Margaret Justin Blanco White (1911–2001),
to depict this process of canalization. He had spoken of the process in the book
as a railyard; the forms of an organism were only able to change developmental
paths if switched, shunted to a new rail line. But Waddington did not like this
metaphor. It was too obviously mechanical, too restrictive. So he asked Piper for
something more organic, natural, free-flowing. Obviously, a river delta came to
mind, fingers forking out from the central channel. Water consolidated in the
central channel for a while, but then any given drop might end up in one or
another of the paths at the end, before dumping into the sea. Development,
Waddington thought, must be a lot like that. The metaphor, and the image Piper
produced to express it, stuck.
“The Epigenotype” was Waddington’s second major contribution in the early
1940s. Today scientists often hold up the article as the foundational work of
epigenetics. 28 At the time, however, it attracted even less attention than
“Canalization of Development and the Inheritance of Acquired Characters.” The
essay was Waddington’s attempt to capture the late 1930s work leading up to
Organisers and Genes in the language of nonspecialists. By the “epigenotype,”
Waddington meant the process by which the genotype becomes the phenotype—
development, in other words. Only Waddington did not mean that development
was merely following a “program” scripted out by individual genes, as later
molecular biologists would. He meant that biologists attuned to both embryology
and genetics should examine, for instance, “concatenations of processes linked
together in a network, so that a disturbance at an early stage may gradually cause
more and more far reaching abnormalities.” 29 He called out for an appreciation
of complexity and multicausality—a pursuit that should engage biologists
trained in multiple subfields. Perhaps now “the analysis of the effects of genes”
could be blended with experimental embryology: “The two methods of analysis
whose rapprochement has for so long been no more than a pious hope can now
actually and in practice come together.” 30
But Waddington underestimated the division between geneticists and
embryologists. With the Second World War raging, basic research into the
complex processes of epigenetics would have to wait. In fact, by the time “The
Epigenotype” appeared in print, Waddington was already directly involved in
war work, far from the biological laboratory.

Unlike J. S. Haldane and the other first-generation organicists in the First World
War, the organicists of the 1930s largely spent the Second World War far away
from the front lines. Combat still had its face-to-face horrors, but in the absence
of large-scale trench warfare, dueling chemists and physiologists would give
way to physicists and mathematicians. Death could be delivered almost without
warning by airplane, battleship, or submarine beyond sight or sound. All sides
understood from the first shot that the best-organized, most efficient side would
win. As much as it was a war of men, guns, and machines, the Second World
War was a conflict written in the quantitative script of logistics.
In their collection Science in War , Bernal, Waddington, Zuckerman, and
twenty other Tots & Quots scientists confidently asserted that men like them
would be more successful with the organization of soldiers and technologically
sophisticated weaponry than the business-trained class of experts usually
enrolled in such work. Scientists educated without the motivation to increase
fiduciary profit should directly influence wartime policy. 31 But through the first
year or so of the war, Waddington remained largely capable of concentrating
exclusively on his scientific work. That changed by 1941. Whether through
pressure by Zuckerman and the other Tots & Quots or because of encouragement
from Gregory Bateson—who left his very pregnant wife, Margaret Mead, in
America to offer his services as an anthropologist to his homeland, attending a
Tots & Quots meeting in the process—Waddington volunteered with the Royal
Air Force. 32 His analytical skill made him perfectly suited for the RAF OR team
assigned to snuff out the U-boat threat to Allied shipping. 33 By all accounts,
Waddington was good at being one of “Churchill’s Scientists.” Yet he could not
anticipate that OR work during the war would have a reciprocal effect on his
scientific life.
Waddington’s scholarly output had been prodigious in the 1930s and early
’40s, but it more or less halted during the war. Perhaps suspicions that he had
never been a serious developmental biologist, just an interloper with a degree in
geology and a scholarship in philosophy, began to reemerge. 34 For whatever
reason, once the end of the war was in sight, Waddington feared that he would be
returning to unfavorable circumstances after already being on unsure financial
footing before the war. His fears turned out to be more or less accurate.
Cambridge zoology had not softened its stance toward experimental embryology.
Though his work in the 1930s had been exemplary, he found the path back into
Cambridge and Strangeways obstructed, the door barred by the head of the
zoology department, James Gray. 35 Waddington nearly left academic biology
altogether. But at last, through the unexpected intervention of American
agribusinessmen in the last months before the end of the war, a door opened for
him. He would work for the Agricultural Research Council (ARC), applying the
administrative and statistical methods he acquired doing OR work and his
knowledge of genetics gleaned from his experience at Caltech to issues like
national food production and animal hybridization. Indeed, these were the same
topics he explored for his anonymous contribution to Science in War.
Then, in May 1945, he received still better news. Unexpectedly, scion of
Scottish genetics F. A. E. Crew recommended that Waddington replace him as
the director of the Institute of Animal Genetics at the University of Edinburgh.
This professorship might allow Waddington to continue theoretical biology
work. At first, he was undecided: The ARC work was rumored to be going to
Oxford. Though he did not record his feelings about the move at that moment,
the choice to leave the Oxbridge-London triangle for Scotland seemed
unappealing: personally, because the Waddingtons lived in London not far from
the Woodgers, Bernal, and a host of Justin’s artist friends; professionally,
because he certainly recognized the potential cost to his career to escape the
orbit of Britain’s most distinguished universities. Still, as he reflected many
years later, the war had left its mark on Waddington. He was prepared to turn his
back on the possibility, however fleeting, of life as an Oxbridge don and take up
a more secure position putting his extensive scientific knowledge to practical
use. When, through a series of negotiations between the ARC, the University of
Edinburgh, and R. G. White, professor of agriculture at Bangor University in
Wales, the two positions were combined, Waddington was indeed named
Director of Animal Genetics at the University of Edinburgh. He joined a group
that had once hosted such luminaries as future Nobelist H. J. Muller. 36 Happily,
he would later look back on his Edinburgh experience as a good fit; after the
science-for-the-people spirit imbibed at Tots & Quots meetings had transformed
into hunting U-boats with the RAF’s OR team, Waddington no longer felt like an
“ivory tower(?) Cambridge experimentalist” anyway. 37
 9

“OFF IN ALL DIRECTIONS LIKE AN

EXPANDING UNIVERSE”

S ome wars end. Germany surrendered; Japan soon followed. Joseph Needham
cared a great deal about how Japan’s withdrawal from China would be handled.
Over the eight years since he had been introduced to Lu Gwei-Djen and her
biochemist companions, his love and admiration for all things Chinese
intensified. By 1942, less than five years after his initial introduction to the
language, Needham had learned enough Chinese to be invited to head a formal
British scientific mission to China. 1 For the remainder of the war, he supplied
Anglo-American instruments and demonstrated the latest techniques in
biochemistry and embryology to Chinese researchers while touring their
scientific facilities, preserving anything that seemed imperiled.
On the one hand, his experiences in China and the relationships he made
there would change Needham’s life. Though Dorothy continued, Joseph
relinquished his role as a dominant figure in British biochemistry. Nearly all of
his time would be spent in pursuit of the history of Chinese science and
technology. By the 1950s, Waddington was the only one of all the core TBC
members still pursuing theoretical biology with the same rigorous eye on organic
philosophy. On the other hand, the more he learned about Chinese philosophy
and science, the deeper Needham’s already existing commitment to the “third
way” became. Perhaps Eastern philosophers had long understood a stance
toward the science of life that Western biologists were only beginning to
appreciate in the twentieth century.
In one of his final publications before leaving for China, Needham reflected
on the impact of the organic philosophy on his own work. According to
Needham, another war, the “war of ideas” that first engaged him—mechanism
versus vitalism—had been over for a while; the turning point in the conflict had
come in the early 1930s. (He only hinted at how his own conversion at that time
reflected those cultural and philosophical contours.) Despite Bergson’s
overwhelming popularity in America in the first quarter of the twentieth century,
vitalism never really stood a chance, Needham insisted. Vitalists had fought the
battle for good reasons: the attempt to keep alive the spirit of “‘religion, poetry,
morality’” against the “over-simplified explanations of biological processes” by
mechanists such as Loeb, who wished to reduce all phenomena to “‘use,
pleasure, self-preservation.’” It was in fact for the “‘recognition of this
mysterious and unfathomable gulf that all the higher pleasures of life depend.’” 2
At the same time, mechanists promised progress, real economic gain, and the
death of superstition. This was a problem for vitalism. If vitalists insisted on
saying the mysteries of organisms were “inscrutable and axiomatic, rather than a
subject for investigation,” then they also appeared to be stuck in the past,
antiprogress and antiscience. 3 Vitalism was unacceptable, according to
Needham, for its tendency to say “this far and no further” to scientific
knowledge, while mechanism was unacceptable for its tendency to oversimplify
and devalue. Thankfully, Needham summarized how organicists had found a
way out of the dilemma. That war, too, had ended, Needham thought; both
mechanists and vitalists had surrendered to organicism.
That confidence was misplaced, as Needham surely knew. He had studied the
long history of the conflict between mechanists and vitalists while writing
Chemical Embryology (1931). He bought into Whitehead’s diagnosis of the
problem, as well. So Needham must have sensed at some deeper level how
intractable the conflict once seemed to be. Since Descartes, the goal of physics
had been to deduce the location in space and time of every kind of object in the
universe. When a new phenomenon was discovered, it had sufficed to describe
its motion and its location. Even once mysterious phenomena such as electricity,
heat, and magnetism had been quantified and categorized by simple location. As
Whitehead put it, “To say that a bit of matter has simple location means . . . it is
adequate to state that it is where it is . . . apart from any essential reference of the
relations of that bit of matter to other regions of space and to other durations of
time. . . . This idea is the very foundation of the 17th century scheme of Nature.”
4 With the concepts of space and matter as foundational principles in the

seventeenth century, it was no wonder that simple location could be sufficient for
understanding a thing. And that governing worldview was contagious, according
to Needham. By the mid-nineteenth century, biology had become physics-
envious and, therefore, also sought to describe biological phenomena in terms of
simple location; only when described this way, in fact, could a biological fact be
counted as explained.
But, as his reading of Whitehead reminded Needham, simple location could
never be enough. It was only an abstraction, a choice not to see certain
connections as relevant to the matter of real interest. To be sure, that way of
thinking made vague, difficult problems clearer. But one must always keep in
mind that clarity was the product of abstraction. To go beyond this, to say that
this observation or that one is the concrete fact—to forget that one is abstracting
away from the interconnectedness of this phenomenon with others—is what
Whitehead called the “Fallacy of Misplaced Concreteness.” And from
Needham’s perspective, biology before the 1930s was shot through with it.
Thankfully, the philosophy of organism that Whitehead shepherded through
the 1920s and that the TBC refined and applied in the 1930s freed the biologist
from having to bow before the altar of mechanism. Needham offered the
organizer experiments as examples of how that organic philosophy played out in
the scientific laboratory. By the time of publication of his massive Chemical
Embryology in 1931, Needham had encyclopedic knowledge of how
biochemical substances act during development. What he learned during the
collaboration with Waddington and Dorothy Needham and the conversations
with Woodger and others at the TBC was that identifying and describing simple
locations of particles in mechanistic fashion—the bedrock of the natural sciences
since Descartes—offered little help in the face of embryonic development:
“[C]hemical substances (the Evocators and Organsers) do not act at random, but
faithfully in accordance with that plan of the body which is decreed by the
characters of the species . . . a plan the field properties of which have earned for
it the name of Individuation Field. Hence the fate of a given monad, protein
molecule, atomic group, or what have you, in the original egg, is a function of its
position in the whole.” 5 Merely recognizing that the function of the part depends
upon its position in the whole, however, does not capture why a “third way” is
better for biology than mechanism or vitalism. After all, removing a gear from
one part of a machine and moving it to another part of that machine would
produce the same observation: same part + different location = different
function. What attracted Needham to the organic philosophy was that embryos
persistently formed adult organisms even under adverse circumstances, even
with different parts, even with some parts removed. The process of growth and
determination was maintained even if the initial components were altered
somewhat.
Vitalists like Driesch and Bergson noticed this, of course—trumpeted it as
vindication of their views, in fact. The reason that organicism was not vitalism,
according to Needham in 1941, was that it eschewed the old dichotomy between
form and matter altogether. Vitalists recognized matter as mute, inert, lifeless.
For them, living things could only be described as “‘an X in addition to carbon,
hydrogen, oxygen, nitrogen, etc., plus organizing relations.’” 6 Form—whether
named élan vital, entelechia , or nisus formativus —had to be added to inert
matter to make it alive. Mechanists held matter to be inert as well but, if they
were atheists, regarded it as capable of producing living organisms on its own
once aggregated sufficiently. Form was merely matter configured. (Mechanists
who believed in the supernatural would need to believe their God was a
watchmaker, imposing order on material chaos from beyond the material
universe, as it were.) Needham, and organicists like him, rejected form/matter in
favor of organization/energy. Organisms were instantiations of time-dependent
nodes in fields or networks rather than collections of particles. And the
organization needed to produce the living being was not the simplistic
combinatorial depiction of the mechanists, full of unopened “black boxes,” but a
vastly more complex, synchronized, coordinated, and fluid process of becoming.
Any statements comparing biological systems to mechanical ones—comparing
the heart to a pump, a brain to a computer, a chromosome to a book written in
the code of genetics—could only be held tentatively. A body was not merely
animated meat. An organism was always a set of minute systems within a larger
set of systems, itself set within ever larger systems. We might take a bit of one
system apart to examine it—how else could science be done? Yet the removal
process itself, as the first generation of organicists feared, might result in the loss
of some piece of information. We were always plagued by epistemic
indeterminism. Every time one killed an organism to analyze it, or even isolated
some biochemicals to understand them better, it was like opening Schrödinger’s
box—one would always find a dead cat, whether or not the cat was dead a
moment before opening it. That, of course, couldn’t be helped: Scientists would
always open the box. It was the only way forward, and vitalists were wrong for
suggesting otherwise.
Notably for Needham, organicism could not and should not remain a
philosophy of biology per se, locked within the biological laboratory. As a way
of living in the world—a weltanschauung—Needham saw in the organic
philosophy the same principles he saw in dialectical materialism, the philosophy
of Karl Marx, Friedrich Engels, and V. I. Lenin. Organicism worked because it
acknowledged the nestedness of concepts in tension, of dialectics, the synthesis
of which would emerge into a higher level. From the thesis of decay and entropy
and the antithesis of growth and aggregation came orderly, constrained processes
of development; from the individual, self-sufficient cell and the destructive
environment came multicellularity, tissues, organs, and organisms. Out of death,
richer life; out of richer life, greater death.
Viewing human society dialectically, organically, meant that Needham
envisioned the tension between individuals eventually leading to tribes, factions,
communities, states, and, hopefully, the universal brotherhood of humanity. For
Needham, this was no utopian vision, however impractical it seemed in the
1940s. The resolution of all human tensions and wars, the hammering out of
selfishness and avarice into general peace and understanding for mutual benefit,
seemed to be promised not only by the words of Jesus of Nazareth and Karl
Marx, but written into the unfolding script of the universe itself.

In 1943, Needham officially began his term as an emissary of the British

delegation to China. Stationed in Chonquing, he made eleven different
expeditions out into the Chinese hinterland over the next two years. He visited
“Cave 15” on the remote western edge of the Gobi Desert, wherein was hidden
the oldest printed book known, the 1100-year-old “Diamond Sutra.” Dorothy
came along for parts of the trip, to the consternation of some of the funding
parties. Just as troubling for some, Joseph successfully hired Lu Gwei-Djen to
accompany him as a “nutritionist.” 7
China and the Chinese entranced Needham. But he was not too hypnotized by
beauty to miss a crucial observation. While visiting historical landmarks and a
panoply of research institutions and universities, he began to notice a trend:
Chinese scholars and tinkerers had hit upon most of the major discoveries made
in Europe from the thirteenth through eighteenth centuries much, much earlier. If
the Chinese were so obviously intelligent, and if they had developed so many of
the major technologies of the so-called scientific revolution years, even
centuries, earlier, why do we associate the origins of modern science with the
observations of Copernicus, Galileo, Kepler, Newton, Gassendi, Hooke, Boyle,
Huygens, and so on? Why did the Industrial Revolution first erupt and take root
in Manchester, Glasgow, Liège, and Düsseldorf and not Chengdu, Kaifeng,
Liaoyang, or Hangzhou? His determination to solve this riddle soon overtook
many of his former passions. He collected notebooks full of observations that
would enable him to attack this question.
In 1946, he was temporarily sidetracked from this new quest. Erstwhile
comrade Julian Huxley had written to him from out of the blue. Would Needham
join Huxley in setting up a particular wing of the United Nations dedicated to
cultural understanding? It would be entitled the “United Nations Educational and
Cultural Organization” (UNECO). Needham agreed, flying home to Britain
almost immediately, Lu Gwei-Djen in tow (Dorothy had already returned). He
joined Huxley in their makeshift offices near Victoria Station, with frequent trips
to the Majestic Hotel in Paris, their headquarters. (It had been spared the horrors
of war by virtue of its cavernous beauty, which proved useful to the occupying
German forces.) Fatefully, in those first few months of 1946, Needham insisted
on inserting “Science” into the mission and the title of the UN department—we
have him to thank for UNES CO.
Needham’s reputation as a formidable scientist, philosopher, authority on
China, and left-winged rabble-rouser began to spread within UN circles. British
intelligence, which had been tailing him since 1931, began feeding their
suspicions about Needham’s politics to the Americans. 8 American intelligence,
in turn, jumped to the conclusion that UNESCO’s leadership was using the
ostensible goal of cultural and scientific exchange between all nations as cover
for spreading support for the new bête noire, Comrade Stalin’s USSR. In a
communiqué to President Harry Truman, CIA director Hoyt Vandenberg singled
out Needham: He was unquestionably a Communist (despite the fact that British
intelligence confessed to having no record of Needham’s membership in the
Communist Party) and was using his scientific stature to negotiate an agreement
between UNESCO and the World Federation of Scientific Workers (WFSW), a
group the CIA had good reason to believe was a Soviet front. Truman acted
swiftly to deny UNESCO funding and political support. 9 In truth, the WFSW
had been established by a group of scientists interested in promoting the use of
science for peaceful rather than militaristic purposes. The group was never fully
accepted by the USSR, despite the fact that many British and French members
were Communists, or at least sympathizers. J. D. Bernal not only helped
organize the WFSW, but served as its first vice president; the connection with
Needham and Huxley at UNESCO was one of longtime friendship rather than
nefarious “negotiation.” 10
Disgusted by the American overreach (he never let on if indeed he had
suspicions his own government had been spying on him for over a decade),
Needham resigned from UNESCO in 1948. He planned on returning to
Hopkins’s lab to resume biochemical work—he had originally told both
Cambridge and Julian Huxley that he would take only two years’ leave—but Sir
Hopkins died in 1947; Needham was left in limbo. He certainly recalled the
veiled threats made by Rockefeller agents in the 1930s that his work would find
little support at Cambridge were it not for Hopkins.
At first he returned to the biochemistry he knew, albeit halfheartedly. By
1950, he had revised and republished Biochemical Embryology as Biochemistry
and Morphogenesis. Though the book was largely a backward glance at the work
he had done with Dorothy, Waddington, Brachet, Holtfreter, and others a decade
earlier, he demonstrated extensive knowledge of the biology that had taken place
since the war. The tome nevertheless led to some head-scratching by the new
generation of biochemists that had grown up in Hopkins’s dotage: Needham
prefaced detailed treatments of the minutiae of laboratory biochemistry with a
discussion of Aristotle’s ancient division of forma and materia , detailed
refutations of Drieschian vitalism—by then a ghost story from a bygone age told
to frighten young geneticists—and wandering paths into Lotke’s topographical
mathematics. 11
On one photographic plate in the middle of the thick book, Needham
displayed a clunky, awkward clay or plaster model of a hill with grooves
scooped into it. It looked like something a bored child might have made out of a
large pile of mashed potatoes and a spoon during his parents’ dinner party.
Needham captioned it a “fate map” in 1950—an attempt to depict the potential
trajectory of cells and tissues, even the whole developing organism. 12 The figure
was a reprint of the same model published in Needham’s Order and Life more
than a decade earlier. In actuality, it was a three-dimensional attempt at casting
an epigenetic landscape. This notion of an epigenetic landscape had only
appeared in pictorial form twice before: this image in Order and Life and in
1940, in the guise of John Piper’s painting of a river valley, as the frontispiece of
Organisers and Genes. It would reemerge as a classic line drawing in
Waddington’s The Strategy of the Genes , seven years in the future, at which
time it would become iconic.
One might believe that the metaphor would have caught on in 1950. After all,
Waddington coined epigenetics in the previous decade. As figure 44 in
Biochemistry and Morphogenesis , however, this strange-looking grooved hill
was just another bit of trivia, obscured because—despite the fact that he had
been elected Fellow of the Royal Society in 1941 for his work in biochemistry
alongside Dorothy (the first couple so honored)—Needham was no longer
regarded as a proper biochemist. By the late 1940s, he did not work in a
laboratory at all; he was, in fact, employed as a librarian upon his return from
UNESCO.
Surprisingly, given his world-class research record in biochemistry, this new
position suited him fine. He had largely turned his back on original biochemical
research to commence a project so immense that the three-volume Chemical
Embryology —a project that Dorothy confessed nearly triggered his mental
breakdown—seemed a mere pamphlet by comparison. Science and Civilisation
in China was Needham’s true life’s work, and he pursued it not from a
laboratory but from his own rooms in Gonville and Caius College, Cambridge
University.

Though it seems ridiculous in retrospect, especially given his inability to

corral earlier projects, Needham guessed that he could fit Science and
Civilisation in China into a single volume. He soon expanded that to seven
planned volumes. But once he had decided, roughly, what would lodge within
those seven, he realized there was far more to say than conventional volumes
could hold. Indeed, he would be saying it to the end of his long life and beyond.
Needham limited his 352-page first volume, published in 1954, to his
introductory thoughts. Volume 2 (1956) captured the history of Chinese natural
philosophy from antiquity to Europe’s medieval period. Volume 3 (1959)
traversed Chinese arithmetic and geometry. By the beginning of the 1960s,
Needham was ensconced in Chinese physics and engineering and here
encountered such a staggering amount of information—mechanical inventions
and concepts that predated by centuries even the most majestic achievements of
Renaissance and Enlightenment Europe—that he was forced to break the
“volume” up into “parts.” Volume 4 thus was bound as three separate books,
each one quite large. But that was just a taste of what awaited him when he
finally turned to China’s innovations in chemistry, Needham’s area of expertise,
in volume 5. As of 2008, thirteen parts of volume 5 have been published. Now,
decades after his passing, Cambridge’s Needham Research Institute continues
his labor of love through the two remaining volumes on life and the social
sciences, the surfaces of which Joseph himself had barely scratched by the time
of his death at the age of 95. 13
The Science and Civilisation in China publication record suggests feverish
work for years on end. Indeed, he only took breaks to visit with friends and to
head Sino-British relations committees and Gonville and Caius College itself in
1966. 14 He also stayed heavily involved in political activism.
Activism got him into trouble even after his resignation from UNESCO. That
negative experience, among others, made Needham suspicious of Western
capitalist governments and especially the United States. The continued invasion
of North Korea by American troops after halting the North’s initial invasion of
the South incensed him. As a member of the International Scientific Commission
to China, Needham heard allegations raised in 1952 that the US military had
engaged in banned biochemical warfare by releasing artificially infected animal
vectors on Korean and Chinese populations. 15 The British government and press
offered Needham multiple chances to recant. In one rancorous episode in
September 1952, journalists cornered him at a multi-hour press conference. His
skill at answering every charge soon quieted them. Needham was convinced that
America was at least conducting something large, “a full-dress rehearsal,” he
thought, in preparation for a larger biological attack. 16 It may have been a
baseless conspiracy theory, but the American government nevertheless
overreacted to Needham’s deft handling of the situation, blacklisting him; he
would be unable to obtain a visa to visit the United States until the 1970s. 17
Needham never answered the question that bore his name. He remained
unable to decipher why the scientific revolution, and the industrial one that
followed it, occurred in England, Scotland, France, Italy, and the Belgian, Dutch
and German states, or why sixteenth-century Portuguese sailors “discovered”
China rather than the reverse. On the one hand, it hardly mattered who was first:
One could imagine similar inventions and discoveries in Europe and Asia
coming about independently just as easily as the alternative—centuries of
knowledge exchange through tangled trade routes that simply failed to attract the
attention of historians. On the other hand, the more that Needham learned, the
more astonished he became. Nearly every “modern” scientific achievement had
a predecessor in China. How could the British be the Victorian-era conquerors of
the globe? Why did so much of the world speak versions of English, French, and
Spanish and not Mandarin or Cantonese? Guns, germs, and steel alone could not
be the answer: The Chinese had them, too—often first.
Perhaps the answer could be located not in resources or material differences
as much as in a fundamental difference in worldview. Here Needham turned yet
again to the organic philosophy to which he had been converted decades earlier:

I spoke of the desire of the Taoist thinkers to understand the causes in Nature, but this cannot be
interpreted in quite the same sense as would suit the thought of the naturalists of ancient Greece.
The key-word [sic ] in Chinese thought is Order and above all Pattern (and if I may whisper it for
the first time, Organism ). . . . Things behaved in particular ways not necessarily because of prior
actions or impulsions of other things, but because their position in the ever-moving cyclical
universe was such that they were endowed with intrinsic natures which made that behavior
inevitable for them. 18

Needham never quit advocating for the “third way.” In fact, the more he
studied Chinese thought, the more he found organicism everywhere. He saw the
Chinese organic influences in Gottfried von Leibniz’s metaphysics. 19 And,
though perhaps they had skipped the revolution in European thought caused by
the triumph of Democritean-Cartesian mechanism in the seventeenth through
nineteenth centuries, there was every reason to believe that the organic Chinese
way of thought would be the scientifically endorsed worldview of the future. It
seemed fair to say that Europeans were late even in articulating the “third way.”
Only in the nineteenth century had a first generation of Europeans begun to see
the outlines of an organic alternative to mechanism and vitalism. European
scientists were recent explorers of a territory that the Chinese—like they had so
many times before—had already mapped: “[O]ur proper conclusion seems to me
to be that the conceptual framework of Chinese associative or coordinative
thinking was essentially something different from that of European causal and
‘legal’ or nomothetic thinking. That it did not give rise to 17th century
theoretical science is no justification for calling it primitive. What remains to be
seen is whether it was not related to a view of the world which modern science is
now being obliged to incorporate into its own structure, namely, the philosophy
of organism.” 20

Waddington had a new position at the University of Edinburgh. In fact, he was

now the head of his laboratory. Given the precariousness of his situation
throughout the 1930s, it was a bit of an irony: Waddington was the only one of
the TBC core who came out of the war squarely a biologist. In 1931, Woodger
hoped Joseph Needham might become the next champion of the “third way.” By
the end of the 1940s, Waddington had supplanted not only Needham but also
Woodger as the most active and best-known adherent of the organic philosophy.
It took some retooling, of course. Even before fascination with DNA dominated
titles in both Nature and the Times , Waddington saw that the language of
modern biology would be written in genetics. Whereas he had learned
embryology in Germany when it was the world’s scientific powerhouse, he went
to America to study genetics. In a little over ten years, he had moved from
petroleum geology to the paleontology of ancient marine animals, to embryology
of amphibians and birds, and now to Drosophila genetics. In the meantime, he
had become an expert in modern art criticism, taught himself enough of
Whitehead’s process philosophy and the symbolic notion in Principia
Mathematica to earn a prestigious scholarship in philosophy, helped Joseph
Needham bring medieval Morris dancing to Cambridge University, maintained
the network of British socialist scientists, learned how to identify and sink
German U-boats, and authored books on ethics, embryology, and genetics. To
call Waddington a polymath would be an understatement.
None of this, however, prepared him for the process of setting up the new
Institute of Animal Genetics. In the atmosphere of strict rationing after the war,
scientists willing to give up positions elsewhere and move to the Scottish capital
were usually younger, less socially settled. Edinburgh, as Waddington already
knew, was distant from manufacturing and scientific centers in the south of
England; equipment gathering took more time and effort than he expected.
Delays and disappointments accumulated. 21 In Cambridge before the war and in
London during it, Wad and Justin had run a house known for freewheeling
conversations with many sorts of interesting people, another sort of Bloomsbury
Circle, blending artists and scientists. In an attempt to replicate the intense and
productive relationships he had experienced then, Waddington convinced the
university to rent Mortonhall, a country manor a full three miles away from the
Institute of Animal Genetics. With that much space, they could accommodate up
to ten families, a few single researchers, and the house staff. In Waddington’s
mind, perhaps, the scientists would work together in the institute and then return
home together to discuss grander theoretical issues. It would be a kind of
scientific commune. Initially, there was some promise. But he missed the
Needhams and Woodgers; they had been the charismatic and hospitable bedrock
of the TBC. While the Mortonhall experiment survived for a good six years,
through some of the worst of the postwar privations, eventually sheer distance
from the institute and lack of reliable transportation for the staff, combined with
the inevitable conflicts arising from the combination of nonrelated families
under the same roof, ended hopes that Edinburgh could have its own
Waddington-led Theoretical Biology Club. 22
The Mortonhall experiment was only the most obvious attempt by
Waddington to foster the connections between laboratory dynamics and
philosophical reflection that he had experienced in the 1930s at Cambridge.
Though he kept in touch with former TBC members, mainly through his own
travels south, since few made the journey to Edinburgh, the situation had
changed perceptibly since before the war, and not just because Needham was
permanently infatuated with China. Wrinch left to the American side of the
Atlantic for good. Something had changed in Woodger’s circles as well. Postwar
theoretical biology seemed not to be following the same anti-mechanistic
trajectory that it had in the 1930s.

Compared to the Needhams, Waddington, and Wrinch, Woodger’s trajectory

stayed more or less consistent after the war. He restarted Theoretical Biology
Club meetings, though now those meetings shifted to London or Oxford
University’s Magdalen College instead of his home in Epsom Downs. 23 His
position at Middlesex Hospital remained secure despite his increasing
involvement with prominent analytic philosophers far from biological or medical
circles including Carnap, Tarski, and Quine. As much as Waddington, the
Needhams, and the other TBC members needed their “Socrates,” he found that
he needed them. Woodger’s new iteration of the TBC had a feel decidedly unlike
that of the prewar years.
Karl Popper became a regular interlocutor and, for a time, Woodger’s closest
contact. That connection was only natural since it was Woodger that shepherded
Popper’s publications through the press while Popper was in New Zealand
attempting to flee from Nazi persecution. That did not mean, however, that they
held political, religious, economic, ethical, aesthetic, or social interests in
common—quite the opposite, as Carnap noted. 24 Though their philosophy
dovetailed, in future years they had a falling out so pronounced that Popper
complained that he felt the usually amiable Woodger would never forgive him
(though he did not actually apologize for any offense, either). 25 When Popper
wrote Woodger’s obituary in 1981, he felt compelled to point out his criticism of
Woodger’s Axiomatic Method in Biology (1937) and highlight his own role in
introducing Woodger to Alfred Tarski. As prickly as he could be at times,
however, Popper largely supported Woodger’s attempt to refine the language of
biology to highlight the explanatory power of organicism. By far the most steady
opposition to Woodger’s project within the postwar TBC came not from Popper
but from a more surprising person: a former student of neurophysiologist John
Zachary Young (1907–1997). Young himself was another TBC and Tots & Quots
regular. Peter Brian Medawar was the name of Young’s student. Popper recalled
Medawar as the person “who became most important for me.” 26
Immunologist Sir P. B. Medawar (1915–1987) would win the Nobel Prize in
1960 and the Order of Merit in 1981. He came to Magdalen, Oxford, in 1932 to
read zoology with Young. The work on squid axons and synapses they did
together inspired Medawar’s work in modern organ transplantation and led to the
Nobel award. Embryologist Gavin De Beer—who was himself Julian Huxley’s
student—had tutored Young. Earlier, De Beer, in turn, had worked alongside
Woodger. (The circles of Britain’s elite intellectuals overlapped extensively:
Medawar would tutor Huxley’s son, Francis.) Young introduced Medawar to
prominent philosophers, including Oxford’s rambunctious Kant scholar, T. D.
“Harry” Weldon. Medawar, like so many other Magdalen men, both loved and
feared Weldon’s acerbic wit. Whether it was through the guidance of Weldon, or
of Young—who by the mid-1930s was deeply involved in the TBC—or perhaps
through his own curiosity, by 1935 Medawar had journeyed to the base camp of
the philosophical peak known as Whitehead and Russell’s Principia
Mathematica. Few at Oxford had ascended that mountain, so Young introduced
Medawar to his hoary-bearded man on that philosophical mountain, none other
than Woodger. Though Woodger was in London and Medawar in Oxford—now
a recent graduate in zoology just awarded a Magdalen senior demyship—they
worked out a system that would allow Medawar to continue scaling the
precipices. As he later admitted, the pursuit “was more a matter of vanity than
anything else: I wanted to be able to say of myself—as a character in one of
Aldous Huxley’s novels said . . . there was nothing any man had written or
thought that I could not master if I chose.” 27
He came down from the mountain of the Principia Mathematica later in the
1930s to pursue his “proper business” in biology. But he never wanted to be a
straightforward biochemist or embryologist. D’Arcy W. Thompson’s On Growth
and Form found its way to Medawar while he got reestablished in Oxford’s
zoology department, now as a junior instructor; he also came across the
celebrated work of Russian biologist Ivan Schmalhausen. Combining the
insights of these two celebrated thinkers, Medawar studied the relationship
between growth, form, and time—attempted to formalize it axiomatically as
Woodger did—on tissue cultures, treating the tissue as a substitute for whole
organism transformation as described by Thompson. He wrote up these findings
as “The Shape of the Human Being as a Function of Time.”
Given this evident ability to traverse philosophy, mathematics, biology, and
chemistry, Young recommended Medawar write a thesis to take the DPhil at
Oxford. Two examiners were appointed: Oxford biochemist Rudolph Peters,
who would soon be called upon to find an antidote to mustard gas in the lead-up
to hostilities in 1939, and a biochemist who was more familiar with
development, morphology, and Whitehead—Cambridge University’s Joseph
Needham. Peters rejected Medawar’s thesis as overly speculative; Needham, no
doubt already familiar with Young’s budding theoretical biologist, advocated for
Medawar. Medawar was ultimately granted the ability to “supplicate” for his
DPhil based on Needham’s attestation. Curiously, Medawar turned down the
honor, regarding it as too expensive—the £25 fee was almost the cost of
Medawar’s first two automobiles combined, he later claimed—and unnecessary
in the world of 1930s British biochemistry, besides. 28
Now a genuine biologist and an Oxford man, Medawar began corresponding
with TBC members with whom he shared so many interests. He turned to
Needham and Woodger in particular when he had a difficult time getting his
philosophically flavored developmental morphology work accepted by his
Oxford peers and superiors. Needham attempted to get Medawar’s “Shape of the
Human Being” article published in the Proceedings of the Royal Society in 1942.
After several revisions, and continued pressure on the editors by Needham (who
was in China by then), it saw the light of day in 1944. 29 That experience, among
others, embittered Medawar toward what he saw as the elitism and conservatism
entrenched in Oxbridge science. He measured the impact of that perceived
snobbishness on Woodger, for instance: “Provided a thing is done vaguely,
generally, and badly (as D’Arcy Thompson did, while paying due respect to the
brilliance of the idea) people will accept it and call it brilliant; but if (cf. your
Axiomatic Method ) you make some attempt to be rigorous and precise, then the
whole thing will be dismissed as being unbiological, or as being of minor
technical interest at the most.” 30 It was, in Medawar’s estimation, Thompson’s
obvious indebtedness to the mysticism of the classics—to Plato and Pythagoras
above all—that made him a darling of Oxbridge dons, “acclaimed and
universally quoted, although, from a formal point of view, it is unprecise [sic ]
and incomplete; . . . faulty in many ways biologically.” 31

During the war, Medawar began to shift away from the “EntwickMech
[Entwicklungsmechanic]” tradition at Oxford—established, he thought, by
Julian Huxley and Gavin De Beer—and toward matters of “surgical
importance.” 32 During the Battle of Britain, Medawar heard about a badly
burned crew of an aircraft that had crashed just to the south. He wondered if later
physical pain and social distress of burn victims like these could be avoided if
there was some way to graft new skin onto their burns. Unfortunately in the
early 1940s, surgeons were only vaguely aware of the relationship between
recipients’ immune systems and the chance that the graft would be rejected.
Understanding the behavior of immune systems in the context of transplantation
became a project that consumed Medawar. Tissue grafting work was cutting
edge, exciting stuff in the mid-1940s, a field where a young ambitious man could
make his mark. Medawar turned thirty in 1945, just before Germany
surrendered, and by all accounts was just such a man—ambitious and quite
interested in leaving a scientific legacy.
Medawar felt flattered when Woodger invited him to join the second iteration
of the TBC in the fall of 1945. 33 (Late in the war, many of his most memorable
theoretical discussions came with Oxford giants C. S. Lewis and J. R. R.
Tolkien. Medawar dined next to Lewis at High Table; he and Tolkien shared
duties as air-raid wardens in Oxfordshire. But there were few opportunities for
him to discuss the philosophy of biology specifically. 34 ) The group began
meeting as soon as the petrol ration was lifted early in 1946. Karl Popper and
Young joined Woodger, Medawar, and W. F. Floyd, a colleague of Woodger.
Popper—who, thanks partly to Woodger, was now at the London School of
Economics—invited another Austrian expat, physicist Hans Motz. In future
meetings, Medawar would invite two of his own especially promising students,
Francis Huxley and N. Avrion Mitchison, the grandson of John Scott Haldane,
the nephew of J. B. S., and the son of Naomi Mary Margaret Mitchison, one of
Scotland’s best-known novelists and a lifelong activist for feminist and socialist
causes. 35
Between Popper—who was just beginning to become internationally known,
largely for his attack on authoritarianism, The Open Society and Its Enemies —
and Medawar, Woodger found his reboot of the TBC much less congenial. 36
Woodger continued sculpting his axiomatic method in biology, still hoping to
complete a Principia Biologae. Medawar had previously been supportive of this
approach. But Mitchison—who later became a renowned immunologist in his
own right—recalled how the atmosphere had begun to change after the war:
“Wittgenstein filtered through Freddie Ayer” convinced many young biologists
that there were a great number of formerly important theoretical questions that
they no longer needed to worry about. Medawar, perhaps prompted by Popper’s
own skepticism, argued that “Zoology based on Darwinism led nowhere,” and
that induction as a scientific method—in other words, “collecting facts without a
hypothesis”—was bankrupt. 37 Thus Woodger’s attempts to get clearer and
clearer about the content of observations in order to build theorems from them
seemed a bit of a waste of time to Medawar. 38
Interactions between them became more strained. Medawar turned on a target
close to Woodger: Waddington and the Needhams’ organizer work. “I do object
to a very large part of the embryology of the ’30s; its technical methods, its
sloppy experimental design, its reckless generalizations, and its ghastly
efflorescence of ‘evocation,’ ‘competence,’ ‘individuation,’ ‘field’ jargon—and
much more besides!” 39 Nevertheless, he regarded TBC meetings and Woodger’s
assistance in particular with fondness: “As we spend so much time at
Th[eoretical] Biol[ogy] meetings merely wrangling with each other, you
probably don’t realize as clearly as I should like how deeply I know myself to be
indebted to you, right from the days of my first reading of the Axiomatic Method.
I owe you, and latterly Popper too, anything I may have picked up about lucidity
and coherence of thought.” 40 As years passed and Medawar received acclaim for
his immunosuppression work, such recollections became still more muted,
equivocal.
At the end of the war, Tots & Quots stalwart Solly Zuckerman moved from
the RAF, where he was scientific director of the British Bombing Survey Unit
and one of Waddington’s superiors, to a professorship of Anatomy at
Birmingham University. Mutual acquaintances, including Young, Needham, and
Bernal, introduced him to Medawar in the mid-1940s. Zuckerman took an
immediate liking to the young, outspoken zoologist. After enlisting fellow TBC
and Tots & Quots member Lancelot Hogben to become chair of the medical
school, Zuckerman appointed thirty-two-year-old Medawar to fill Hogben’s
position as professor of Zoology. Only three years later, Medawar was elected
both Fellow of the Royal Society and dean of the faculty at Birmingham
University.
Now the busiest and most sought after that he had ever been, Medawar found
it difficult to think at all about theoretical issues or even to reply to Woodger’s
letters addressing topics in the philosophy of biology. When they did correspond,
it was clear that Medawar no longer cared to engage the concepts that Woodger
still held dear. His increasingly “impatient and, I fear, rude, briskness” with
Woodger eventually led to a long-term falling out between them in the early
1950s when Medawar, among other things, explicitly dismissed Woodger’s focus
on organizing relations in the organism—the heart of organicism. 41 He later
critiqued Woodger’s axiomatization in its entirety: “Woodger creates the
impression that genetics is in a great big muddle, with everyone talking at cross
purposes. . . . Luckily, this is simply not true . . . the patient is not nearly ill
enough for Dr. Woodger’s physic. I am therefore out of sympathy with his
scheme of treatment.” 42 Whereas once Medawar had largely agreed with
Woodger that biologists needed more precision in their language and appreciated
Woodger’s axiomatization—like Suttie, he saw it as a way to prevent poaching
and to resolve antimonies between subfields—he now found the cure worse than
the disease. Now that he had become part of the inner circle, the dominant
group, analyzing the core concepts of biologists seemed like a waste of time. 43
Medawar continued to have a negative impact on the organic philosophy after
leaving the second TBC. As his fame grew through the 1950s and ’60s as
director of the National Institute for Medical Research, he found more
opportunities to publicly pronounce on issues of scientific and social importance.
He was elected president of the British Association for the Advancement of
Science in 1969, the position once held by General Jan C. Smuts. But his
presidential address depicted a very different vision of biology than Smuts’s
exposition in 1931. Both implicitly and explicitly, he had undercut Woodger’s
axiomatization and mocked organicism. Among his favorite targets were Hans
Spemann and the organizer work of Joseph Needham and C. H. Waddington in
the 1930s. And he had powerful support from other authoritative realms: For
now, in the wake of the Second World War, physics-inspired Anglo-American
philosophy of science had taken aim at the “third way.”
 10

MECHANISM REDUCED TO MOLECULES

M edawar’s skepticism regarding Woodger’s insistence on organization—a

small personal disagreement on the surface—reflected much larger trends that
would turn the tide against organicism for decades. That global shift was so
profound that even in the middle of it historians of biology felt that looking only
a few decades back was like staring into a lost world: “But look back at the
situation in the 1930s and compare this present state of affairs with the state of
affairs then. In the 1930s, embryology was riding high on the crest of the wave.”
1 Embryologists had been some of the chief promotors of “third way” thinking;

the wave broke after the Second World War with the rise of a new philosophy of
science, the birth of molecular biology with almost exclusive focus on the
molecules of the cell nucleus, and the concomitant resurgence of a popular
mechanistic scientism.
The next few chapters outline a few of these larger trends. The present
chapter offers a “flyover” view of the postwar scientific, philosophical, and
cultural landscape. Those that follow this one provide “on-the-ground”
assessment, primarily of Waddington’s interactions with molecular and
evolutionary biologists in the 1950s. This includes smaller episodes, such as the
attempts by Waddington and Joseph Needham to preserve the legacy of the
organizer project and the memory of the original TBC, and their efforts to
position epigenetics and organicism as a counterbalance to implicit genetic
reductionism in the neo-Darwinian synthesis.
One could pick any number of representatives to be the subjects of this
“historical context” chapter. Two in particular illustrate just how broad were the
transformations in theoretical biology after the Second World War: analytic
philosopher Ernest Nagel and biochemist-turned-science-popularizer Isaac
Asimov. Nagel was among the most well-connected expositors of the
philosophical rejection of organicism. He seems to have turned his critical lens
on the “third way” following the publication of Ludwig von Bertalanffy’s
“General Systems Theory.” (Bertalanffy originally developed his anti-
mechanistic theories in conjunction with Woodger in the 1930s.) Asimov’s case
demonstrates that, by the 1950s, a new DNA-centered mechanism had spilled
over from the laboratory to become part of popular consciousness.

Ernst Cassirer, Polish philosopher of physics and friend of Albert Einstein, died
in 1945. But in 1950, the translation of his Problem of Knowledge: Philosophy,
Science, and History Since Hegel reintroduced a younger generation of
American and British philosophers to the mechanism-vitalism debate. As so
many had done before, Cassirer faulted vitalists for statements regarding
scientific concepts that allowed for no empirical test—Driesch’s entelechy,
Bergson’s élan vital, even Jakob von Uexküll’s umwelt concept. Yet he refused
to side with the mechanists. Many mechanists, Jacques Loeb, for instance,
promised control over life itself, including the creation of artificial organisms
from ordinary chemicals. Obviously no such thing had happened. And there
were other deeper and more disturbing problems with both the logical structure
and the implications of mechanism. “[A]t present all is still in flux,” Cassirer
reflected, “and there can be no mention as yet of any conclusive and generally
accepted outcome.” 2 Both sides had convincing arguments; both could point to
evidence they counted as definitely in favor of their position. Thus, despite the
fact that Needham had declared this debate closed in the 1930s, Cassierer’s
stature as a philosopher and public figure signaled that perhaps the mechanism-
vitalism debate was not closed after all. The Problem of Knowledge cast doubt
that a fully satisfactory solution could be found—as long as mechanism and
vitalism were the only two possibilities.
It was in this new moment of uncertainty introduced by Cassirer that
Woodger’s old Viennese friend from the 1920s and ’30s, Ludwig von
Bertalanffy (1901–1972), reappeared to open a new attack on mechanism. This
time, he gave it a sophisticated moniker: “general systems theory” (GST). But he
began his “Outline of General Systems Theory” essentially the same way he had
Modern Theories of Development , his collaboration with Woodger, two decades
earlier:

Corresponding to the procedure in physics, the attempt has been made in biology to resolve the
phenomena of life into parts and processes which could be investigated in isolation. This procedure
is essentially the same in the various branches of biology. The organism is considered to be an
aggregate of cells as elementary life-units, its activities are resolved into functions of isolated
organs and finally physico-chemical processes, its behaviour into reflexes, the material substratum
 of heredity into genes, acting independently of each other, phylogenetic evolution into single
fortuitous mutations, and so on. As opposed to the analytical, summative and machine theoretical
viewpoints, organismic conceptions have evolved in all branches of modern biology which assert
the necessity of investigating not only parts but also relations of organisation resulting from a
dynamic interaction and manifesting themselves by the difference in behaviour of parts in isolation
and in the whole organism. 3

General systems theory, then, was another example of the by now familiar
attempt to find a “third way” around the mechanism-vitalism debate. Though it
was well received in the early 1950s, GST offered little that had not already been
expressed in Woodger’s “‘Concept of Organism’” (1930–31), Needham’s Order
and Life (1936), or, frankly, even in Edmund Montgomery’s “The Unity of the
Organic Individual” (1880). What was new was the way Bertalanffy dressed up
GST. Just as Whitehead and Smuts had done in the 1920s, he liberally deployed
the terminology of physics, especially thermodynamics and new discoveries at
the subatomic level—reflections of the Manhattan Project and the Copenhagen
interpretation of quantum physics. He also included a new factor, already
becoming an early Space Age buzzword: information.
The physicists and engineers of the Second World War invented not only new
ways of killing but also new methods of communication—or at least new jargon
to apply to theories about communication. Central to this new way of speaking
about speaking was the concept that some difference, some data, in a system
would drive the outcome one direction instead of another. According to the
concept as spelled out by Claude Shannon and Warren Weaver in their influential
The Mathematical Theory of Communication (1949), a product of the early
1940s Macy Cybernetics meetings, this “difference maker” would be the
information in a system; entropy was the name given to the predictability, or lack
thereof, of any particular outcome being reached or any desired message being
conveyed. This was the mathematized, cybernetics-infused language Bertalanffy
adopted for his GST—his retooled version of first generation organicism.
Whereas once J. S. Haldane had been fascinated by regulation systems within
organisms, in Bertalanffy’s GST, biological regulation would become
“homeostasis” and “feedback loops”; L. J. Henderson’s notions Bertalanffy
called “dynamic equilibria.” Organisms themselves he typified as “open
systems” and sources of increasing order, the antithesis of the concept of entropy
—“negentropy.” Living beings, in other words, are capable of taking from the
disorderly universe and reordering it in service of their own self-maintenance,
repair, and growth. Nonliving systems might also be capable of this negentropy,
making order out of disorder. The formation of snowflakes, thermostats
controlling the climate of buildings, and so on all seemed to be examples of
negentropic systems. But Bertalanffy added that organisms are also defined by
“equifinality,” the ability to reach their final states given a broad spectrum of
initial conditions and using multiple pathways—something like the old
Whiteheadian notion of canalization filtered through Waddington.
Bertalanffy insisted that GST was not a new version of vitalism. But it was
definitely anti-mechanism:

The mechanistic world-view found its ideal in the Laplacean spirit, i.e. in the conception that all
phenomena are ultimately aggregates of fortuitous actions of elementary physical units.
Theoretically, this conception did not lead to exact sciences outside the field of physics. . . .
Practically, its consequences have been fatal for our civilisation. The attitude that considers
physical phenomena as the sole standard-measure of reality, has led to the mechanisation of
mankind and to the devaluation of higher values. The unregulated domination of physical
technology finally ushered the world into the catastrophical crises of our time. After having
overthrown the mechanistic view, we are careful not to slide into “biologism,” that is, into
considering mental, sociological and cultural phenomena from a merely biological standpoint. As
physicalism considered the living organism as a strange combination of physico-chemical events or
machines, biologism considers man as a curious zoological species, human society as a bee-hive or
a stud-farm. . . . The organismic conception does not mean a unilateral dominance of biological
conceptions. When emphasising general structural isomorphies of different levels, it asserts, at the
same time, their autonomy and possession of specific laws. 4

Augmenting at least two generations of anti-mechanistic rhetoric, Bertalanffy
stressed that continued reduction of social concepts to biological ones and
biological concepts to physics and chemistry would ultimately lead to an
impoverished view of humanity and what he called “the catastrophical crises”
humans would face in the Cold War, including the looming threat of
thermonuclear armageddon.
His goal—the upshot of his redeployment of the organic philosophy using the
new idiom of cybernetics, in other words—seemed somewhat schizoid, at least
on the surface. On the one hand, Bertalanffy claimed GST would contribute to
the unification of science effort championed by his and Woodger’s mutual
friend, philosopher of science Rudi Carnap. On the other hand, Carnap’s
unification efforts had been physics-centric, and Bertalanffy clearly disagreed
with those he deemed mechanists for doing what Carnap seemed to be doing:
regarding biology and psychology as more or less soft, animistic pursuits just
waiting to be reduced to physics. Bertalanffy believed it was possible to have it
both ways. The general in general systems theory meant that he believed all
phenomena from quanta to organisms to governments and economies could be
analyzed as systems or complex networks. But he also regarded each of these
levels—subatomic, molecular, cellular, organismic, social, and so on, as discrete.
Each level had its own laws, some possibly unique to that level. It was Russell’s
“theory of types” reborn, or perhaps the notion of levels from the works of
Woodger, Needham, and Waddington in the 1930s. Reality is nested , such that
there exists a “correspondence or isomorphy of laws and conceptual schemes in
different fields.” Science could be unified because different levels of reality have
some structural elements in common, not because psychology was just biology
dressed up or biochemistry anthropomorphized. Bertalanffy’s “third way” was
cybernetic organicism—not the reduction of organisms to machines, per se, but
the recognition that the universe was far more complex than any simple machine
theory would allow.

GST did not prevent philosopher Ernest Nagel (1901–1985), a colleague of

Carnap’s at Columbia University, from taking “organismic biology” to task in
1951. In order to do that, however, he would have to avoid directly addressing
GST and even Cassirer’s widely read Problem of Knowledge. Instead, Nagel
aimed at E. S. Russell’s Interpretation of Development and Heredity . 5 In
retrospect, the use of Russell’s book was a regrettable choice. To be sure, Russell
fairly represented first-generation organicism. Had Nagel centered his crosshairs
on Bertalanffy and Woodger’s Modern Theories of Development or Needham’s
Order and Life instead, Nagel’s critique may have turned out differently. Perhaps
the damage done to the organic philosophy may have been less pronounced;
perhaps the philosophy of biology would have attracted more sympathetic
spokespeople—as Woodger hoped back in the early 1930s.
As it was, Nagel effectively looked past the second generation of organicists
to stab at an ichthyologist’s work written two decades earlier. 6 He focused on a
small number of Russell’s quotes, including: “[T]he activities of the organism as
a whole are to be regarded as of a different order from physico-chemical
reactions, both in themselves and for the purposes of our understanding.” 7 This
statement, Nagel said, was nearly an admission to vitalism in that it suggested
organisms were outside of the physicochemical order “in themselves,” that is,
ontologically. Russell’s position, according to Nagel, denied that mechanistic
explanations could ever suitably describe the organism and that organisms could
not be accounted for by those explanations found in chemistry and physics, even
in principle. Ultimately, Russell claimed, according to Nagel, that biological
laws stood apart from laws in physics and chemistry. He took this to be the
central claim of all organic philosophy, though he referred merely to the works
of first-generation organicists.
After picking apart claims about the nonanalyzability of organisms, Nagel
turned to the claim that organisms are wholes that share certain relations in
common that are not reflected in nonbiological realms. He offered two
counterexamples. First imagine a clock: We possess physical laws that describe
the forces at work in springs, cogwheels, and pendulums. Therefore we have a
way to describe the behavior of a clock, a complex whole, by the sum of its
parts. Then consider a historical example: In the nineteenth century, physicists
understood the thermal properties of a gas; they assumed the thermal properties
of solids could be explained in the same fashion. Repeated observations,
however, proved them wrong: The Newtonian kinematic theory did not work
precisely when applied to solid masses of a certain size. As it turns out, when it
comes to heat at least, solids act differently in aggregate. They are literally
greater than the sum of their parts. Once quantum mechanics came about,
however, the problems with the thermal behavior of solids were more or less
solved. In the one example, a complex whole could be understood completely by
the analysis of its parts using known physical laws and, in the other, a previously
problematic whole could be successfully explained once new physical concepts
were discovered. For Nagel, these examples at least hinted that organicists who
insisted that the organism was a whole irreducible to the sum of its parts were
making much too strong of a claim:

To say, therefore, that the behavior of an organism is not the sum of the behavior of its parts, and
that its total behavior cannot be understood adequately in physico-chemical terms even though the
behavior of each of its parts is explicable mechanistically, can only mean that no body of general
theory is now available from which statements about the total behavior of the organism are
derivable. The assertion, even if true, does not mean that it is in principle impossible to explain
such total behavior mechanistically, and it supplies no competent evidence for such a claim. 8

The most the organicists could maintain, argued Nagel, was that we had not yet
reduced biology to physics and chemistry.
All told, Nagel made a historically uninformed claim. Early in the twentieth
century, Jacques Loeb had insisted that he would provide said mechanistic
explanations for all manner of organic behaviors and systems. Now Nagel could
only hold to the claim that it was still possible—in principle, at least—that such
mechanistic explanations would be forthcoming one day. He did not argue for
the reduction of biology to physics. He did not even discount the attempt of
organicists from Morgan, Henderson, Haldane, Russell, and beyond to resolve
the debate between mechanists and vitalists: Organicists, he admitted, made a
“heuristically valuable” stand against the imperialism of physics. 9 Nagel merely
pointed out that, as Russell formulated it, there was always the possibility that
organic processes could one day be explained solely through chemistry and
physics. But few of the more recent generation of organicists had claimed to
establish the in-principle impossibility of reducing biological laws to physical
and chemical ones—imagining, of course, that what counted as a physical law
expanded. This was precisely what Bertalanffy advocated with his GST: a
broader concept of physical laws that would include the priority of systems.
Even Alfred North Whitehead in the 1920s had maintained that physics itself
was organic, just in a way that we did not or could not yet measure. Physical
laws were not supplemented by vital entities; they just should not be interpreted
as Democritean-Cartesian, according to Whitehead. Nagel also ignored the fact
that proponents of a “third way”—including Woodger, Needham, and
Waddington—had for two decades attempted to move the discussion away from
ontology into the territory of scientific epistemology, methodology, and
language.
Ironically, Russell himself, on the page immediately following the one quoted
by Nagel, stated that his entire project—his contribution to that first generation
of organicism—merely captured the “conceptions which are tacitly followed by
most biologists in the practical conduct of their researches, and, accepting the
broad facts of observation at their face value,” Russell would “generalize these
without paying much attention to ultimate questions of philosophy.” 10 In other
words, even Nagel’s main target made different claims than those he critiqued.
Even if he was not the most up-to-date example of a consistent “third way”
theorist, Russell was actually arguing that most of the time biologists think and
speak organically; they investigate specimens as whole organisms; and whether
or not biological laws are reducible to physical ones, biology should remain an
autonomous science with its own ways of seeing.
Nagel’s partial critique made an oversized impression. And it proved to have
two lasting, perhaps unintentional, effects on the history of the “third way.” The
first was in the fast-growing field of philosophy of science. Nagel suggested in
his critique that organicism was merely a species of vitalism. In doing so, he
reforged chains between these concepts that two generations of biologists
interested in the organic philosophy had labored to sunder. After Nagel, “vitalist”
and “organicist” would often be used interchangeably, and the discrediting of
vitalism could easily be seen as a concomitant discrediting of “third way”
philosophy. 11
Secondly, Nagel’s essay signaled a definitive change in the terms of the
debate from mechanism to reductionism in both biology and philosophy. Both
vitalists and organicists had objected to the uncritical application of mechanical
concepts and metaphors to living things—this is what Whitehead termed
Democritean-Cartesianism. Organicists recognized the complex patterns of
relationships exhibited within and between organisms. Unlike vitalists,
organicists insisted that these patterns and organizing relations were analyzable
using the tools of empirical science, not ephemeral entelechies or vital spirits.
Nagel’s critique of organicism highlighted a subtle but significant shift in
language. With the advent of molecular biology in the postwar period, the
question was no longer focused on whether concepts of heat and electrical
charge should play a crucial role in discussing animal behavior. Should a master
molecule be discovered, a molecule coding for the secrets of life, it would no
longer matter whether the regulation of CO2 had an analogical example in the
mechanical world, as it had to Haldane. Instead, biological processes would be
describable in terms of the behavior of that biological molecule. That molecule
might be unique to living things, but it would still be a molecule like any other.
As a molecule, composition and behavior would be describable entirely in
physicochemical terms. And as a master molecule, it would determine what cells
do, what organisms do, what whole populations do. Thus Nagel’s “Mechanistic
Explanation and Organismic Biology” really initiated a debate about the
pervasiveness of reductionism: Are all the things that matter about an organism
really just features of its master molecules? Is the rest of the organism a mere
marionette, with these molecules ultimately pulling the strings?

Nagel was only a few years early. Biologists had known about the importance of
deoxyribonucleic acid (DNA) for a while, of course. But when Dorothy Wrinch
toyed with a model of how DNA might be bound up in the chromosomes in the
1930s, she saw it as structural support for proteins. Proteins obviously exhibited
more complexity and, consequently, more specificity than DNA. If there was
anything to the notion of chromosomal genes making phenotypic traits, those
genes would have to be complex and specific—written in the language of
proteins, in other words.
The first solid model of how DNA might work came early in 1953. Wrinch’s
rival Linus Pauling, the discoverer of the single-helix formation of protein,
devised a far more complex triple-helix model for DNA. But he possessed fairly
poor imaging data and it led to a major error in judgment. It was Jim Watson and
Francis Crick who deduced the double-helical structure a few weeks after
surreptitiously examining Rosalind Franklin’s “B-form” x-ray diffraction images
—the crispest, most precise images available. Soon the world heard about the
iconic image of two spirals dancing around each other with pyrimidines and
purines hovering between like rungs in a twisted ladder. Watson believed all
along that DNA was the master molecule, the secret of life itself. After Watson
built a cardboard model, Crick, too, was convinced, proclaiming to “everyone
within hearing distance that we had found the secret of life.” 12 In truth it did
seem, following the Cold War developments in nuclear physics, that molecules
in general—and elegant ones like DNA in particular—held secret powers just
waiting to be exploited.
This was certainly the suggestion made by prominent physicist Erwin
Schrödinger in the middle of the Second World War. In his What Is Life? (1944),
Schrödinger wrestled with the same large theoretical questions that members of
the TBC contemplated a decade earlier: How does physics impact biology?
What is going on in heredity? Is life based on the same mechanical laws that
govern everything else? What can we say about the size and complexity of
genes? What are genes, anyway? Though evenhanded in his approach, he
definitely sided with the notion that scientists should focus on the behavior of
molecules and even submolecular units if they want to understand life itself:
“We find [life] controlled by a supremely well-ordered group of atoms, which
represent only a very small fraction of the sum total in every cell. Moreover,
from the view we have formed of the mechanism of mutation we conclude that
the dislocation of just a few atoms within the group of ‘governing atoms’ of the
germ cell suffices to bring about a well-defined change in the large-scale
hereditary characteristics of the organism.” 13 It was true that Schrödinger felt the
discovery of new laws of physics might be necessary in order to deal with the
complexity of organisms. This indeed was the perspective he received by
reading the “Three Man Paper” (3MP) of physicist Max Delbrück, geneticist
Nikolai V. Timofeev-Resovskii, and physicist Karl Zimmer. 14 But whereas
Delbrück’s interpretation was not a typical mechanistic reading of genes-to-
traits, Schrödinger read him in a more mechanistic light than Delbrück intended.
15

In the early Cold War, fascination with the molecular spread beyond scientists
and philosophers, thanks to science popularizers who actively promoted the
notion that the behavior of molecules was the key to understanding all manner of
formerly difficult puzzles. As word spread that simple molecules held the key to
life, “third way” sentiments that an understanding of the organism remained
crucial seemed immaterial, a quaint relic of a less sophisticated era. The
popularizers trumpeted that story.
 Among the most effective of them was a young man of Russian descent who
grew up in Brooklyn, New York, and taught biochemistry at Boston University.
His name was Isaac Asimov (1919–1992). Understanding Asimov allows us to
envision how powerful and persuasive gene-centric mechanism became even
outside of scientific circles.
Asimov began his prolific writing career even before the war, and had
composed the opening of his famous science fiction trilogy Foundation by 1942.
He would complete the trilogy by the time Watson and Crick published their
short letter to Nature describing the structure of DNA. That coincidence would
resonate through Asimov’s writing when he turned from science fiction to
science popularizing.
The Intelligent Man’s Guide to Science , his attempt at a synthesis of
midcentury scientific knowledge, became a best seller and went through several
editions. 16 The book ran to two volumes. The second volume tipped Asimov’s
hand. He began it with “The Molecule” (the first volume, on “The Physical
Sciences,” apparently had been too concerned with “The Universe,” “The
Machine,” and “The Reactor” to include molecules there). It featured triumphal
passages including the following: “Modern science has all but wiped out the
borderline between life and non-life. Nowadays the question ‘What is life?’ is
asked by physicists as often as by biologists. In fact, biology and physics are
merged in a new branch of science called biophysics—the study of the physical
forces and phenomena involved in living processes. . . . And it is to biochemistry
(‘life chemistry’) that biologists today are looking for basic answers to the
secrets of reproduction, heredity, evolution, birth, growth, disease, aging, and
death.” 17 Asimov did not hide his justification for making such a bold statement:
“Once we get down to the nucleic-acid molecules, we are as close to the basis of
life as we can get. Here, surely, is the prime substance of life itself. Without
DNA, living organisms could not reproduce, and life as we know it could not
have started. All the substances of living matter . . . depend in the last analysis
on DNA.” 18 As a competent reporter of the recent history of biology, Asimov
could find nothing to compete with the overwhelming perspicuity of the DNA-
as-molecule-of-life story. Given that DNA was just a molecule, biochemists like
himself—and, ultimately, physicists—would in the very near future offer all
relevant information about that molecule, about life itself.
Asimov’s DNA-centric vision, that life itself depended upon this molecule,
this “prime substance,” flowed effortlessly out of the philosophy implicit in his
Foundation trilogy. The first episode opens on a galaxy-spanning human empire
that has controlled thousands of inhabited worlds for millennia. A handful of
brilliant scholars study psychohistory : the behavior of peoples—even the
residents of entire solar systems—based on the simple philosophical premise
(and some arithmetical hand waving) that humans operate like molecules of gas
in a closed container. Those molecules behave according to the clockwork of
Newtonian kinematics: “[Psycho-history] was the science of mobs. . . . It could
forecast reactions to stimuli with something of the accuracy that a lesser science
could bring to the forecast of a rebound of a billiard ball.” 19 Psychohistory was
Asimov’s projection of James Maxwell’s demon applied to living things in the
same way that it could be applied to gasses in closed containers—a Laplacian
vision of a closed, deterministic, and therefore predictable universe.
The story of DNA as molecule-of-life dovetailed nicely with Asimov’s larger
deterministic vision. No one would doubt that the behavior of molecules, even
ones as complex as Schrödinger’s aperiodic crystal, could be determined by the
unchanging laws of physics. Since DNA, that prime substance, obviously
controlled biological development and at least preconditioned the behavior of all
living things, it was an easy logical jump to regard the control of living things as
a wetter version of engineering.
The takeover of organismal biology by reductionistic mechanists was already
evident by the middle of the twentieth century, according to Barry Commoner,
Washington University in St. Louis botanical physiologist. He expressed this
fear in his review of Asimov’s Intelligent Man’s Guide. One need only examine
the various laboratories belonging to university biology departments around the
United States, thought Commoner. One kind of laboratory featured expensive
electromechanical devices, well-paid postdoctoral fellows, and scores of
students. The other kind displayed fewer people, less expensive devices. This
obvious material distinction had an interesting historical cause: “a widening gap
between the more traditional areas of biology,” by which Commoner meant
those areas associated with evolution and development versus those “closely
related to modern chemistry and physics.” 20 He saw this division reflected even
in the other reviews of Asimov’s Intelligent Man’s Guide. One reviewer, a
historian of astronomy and physics from Princeton University, extolled Asimov’s
reductionistic mechanism: “For [Asimov] . . . biology is a system that proceeds
from biochemistry to the associated subjects of neurophysiology and genetics.
All else, as they used to say . . . is stamp collecting. . . . I happen to agree firmly
with Asimov about what is central in science and what is not and I will defend
him to the death against traditionalists who might deplore his . . . giving short
shrift to ‘Natural History.’” But for Commoner, this long-term trend looked
frightening. Not because his profession was in jeopardy—plant physiologists
would likely be needed for a long while yet—but because the investigation of
the actually observed biological phenomena of the world, whole organisms and
their ecosystems, in other words, could not be done solely by physicochemical
means. Commoner would become ever more emphatic in this critique, as we will
see later.

By any measure, a new and improved mechanism had emerged out of the
Second World War and had shrugged off the challenges leveled at mechanism by
the organicists of the 1920s and ’30s. One could witness this trend in small,
personal cases—Medawar’s ambivalence toward organicism and falling out with
Woodger in the midst of his rising success in less theoretical, more practical
matters. In addition, broader scientific and even sociocultural changes were
visible. Nagel’s critique and the work of his followers became ensconced as
disciplinary wisdom in a midcentury analytical philosophy of science
preoccupied with physics rather than biology. Asimov’s promotion of DNA-
centrism certainly struck a chord with his American audiences. Intelligent Man’s
Guides surpassed mere best-seller status to become a classic example of
twentieth-century science writing.
Upon reflection, the mid-1950s situation in biology echoed trends seen a half
century earlier. In that era, it was Wilhelm Roux who attempted to extend
mechanism over what was thought to be the last mystery of biology, embryonic
development, through his Entwicklungsmechanik program. Back then, sea
urchin cells nudged apart by Roux’s follower, Hans Driesch, had slowed the
locomotive of mechanism for a time. In the 1950s, just as molecular biology was
extending a new and improved reductionistic mechanism focused on the
behavior of DNA in the nuclei of cells, Waddington was conducting experiments
on Drosophila that would again highlight the promise of epigenetics as the new
“third way” biology of the future.
 11

THE LYSENKO MORALITY TALE AND THE

EPIGENETIC LANDSCAPE

E arly in the Second World War, Waddington wrote to Joseph Needham that he
was “thinking of popularising the word ‘epigenetics’ instead of the cumbersome
phrase ‘experimental embryology.’” Needham knew the history behind this term
“epigenetic” or “epigenesis”; his stand-alone History of Embryology covered it
in detail. 1 Woodger, too, had discussed the concept in Biological Principles. It
was, in fact, a focal point of TBC meetings. Waddington could see a number of
advantages to popularizing the old term: Epigenetics had a substantial scientific
pedigree, “having meant since Aristotle’s day the causal aspects of
development.” He also thought the Aristotelian term captured the spirit of
modern biology better than Roux’s nineteenth-century term
“Entwicklungsmechanik” because epigenetics seemed to indicate that modern
“genetics and [embryology] are sister sciences which are rapidly growing closer
together.” 2 In Waddington’s mind, twentieth-century epigeneticists would
investigate the causes of development without either privileging genes-only
explanations or ignoring genetics altogether. Epigenetics was still a “third way
idea,” even though it was tied to modern genetics: It required conceptualizing of
the organism as a system of interactions between the lowest levels (i.e., genes),
the developing systems of the organism, and the highest levels (i.e.,
environment). Epigenetics indicated a constant dialogue between the whole
organism, its context, and its parts—no simplistic genetic “program” to unroll.
Waddington’s “The Epigenotype” (1942) spelled out the questions
practitioners of this new subfield might pursue, but it was much easier to say
“epigenetics” or “epigenome” than it was to rally researchers into the science.
Though today’s epigenetics researchers point to “The Epigenotype” as a
landmark paper, Waddington did not see it as such. In fact, a full decade elapsed
between this re-coining and his first major publication that really demonstrated
what he meant by epigenetics in the laboratory. In 1952, Waddington re-unveiled
epigenetics at the Society for Experimental Biology’s Symposia on Evolution
with the talk “Epigenetics and Evolution.” 3 By that point, he had been at the
University of Edinburgh for almost five years and—when he wasn’t dealing with
incessant challenges running the lab and maintaining Mortonhall—had buried
himself in Drosophila work. He published eight papers dealing with
development, genetics, and evolution from 1945 to ’52; that pace increased
significantly after “Epigenetics and Evolution.” While Watson and Crick
tinkered with the model of DNA that would eventually earn them the Nobel
Prize, Waddington published a small flood of studies attempting to combine
genetics and embryology.
Before the war, scientists celebrated such synthetic work, at least in Europe.
So, one might think that the modern subfield of epigenetics possessed all the
features necessary for it to emerge just after the Second World War. Needham
and Waddington had already introduced the concepts through works like Order
and Life and Organisers and Genes , and Waddington’s “Epigenotype” had
already provided the terminology. Scientists at the time were developing the
analytical tools to observe and trace the tiniest features of the living organism—
the tools that would lead to the milestone discoveries in molecular biology. The
pieces were all in place, in other words. Yet epigenetics nearly disappeared until
finally emerging at the very end of the twentieth century. Why?
That answer has something to do with the general context that we have
already reviewed: changes in Western philosophy exemplified by Ernest Nagel’s
dismissal of organicism and widespread support for DNA-centric mechanism
evident through Isaac Asimov’s popular works, to name two such factors. Just as
important was the marginalization of organicism by the very network of
scientists with whom Waddington had close affiliations—geneticist H. J. Muller
and ornithologist Ernst Mayr in particular. That marginalization process is the
subject of this chapter.
Muller was the chief narrator of the “morality tale” woven in the wake of the
tragic events surrounding Soviet geneticist Nikolai Ivanovich Vavilov and
agronomist Trofim Denisovich Lysenko, now known as the “Lysenko affair.”
The lessons Anglo-American biologists drew from the Lysenko affair made
epigenetics unpalatable and “third way” thinking suspect more generally.
Waddington’s epigenetics also raised concerns regarding the interpretation of
evolution extolled in the “modern” neo-Darwinian synthesis of Ernst Mayr,
among others. While Mayr decried the reduction of evolution to “beanbag
genetics,” he also feared Waddington’s epigenetics left a door open to the same
Lamarckian interpretations promoted by Lysenko and his ilk. Waddington even
visited Lysenko’s Leningrad laboratory and spoke with the villain in 1962. 4 No
wonder that, at the mid-1970s celebration of the modern synthesis, Hilde
Pröscholdt’s old lab mate Viktor Hamburger reflected, regretfully, that Mayr and
others had relegated Waddington’s work to little more than a “Missing Chapter.”
5 Epigenetics, and the organic philosophy in which epigenetics had taken root,
hinted of Lamarckism. During the Cold War, “Lamarckism” indicated sympathy
for Soviet Lysenkoism.

From the 1930s onward, Soviet agronomist T. D. Lysenko (1898–1976)

advocated “vernalization,” a process discovered in the nineteenth century and
brought to the world’s attention by the work of German botanists after Armistice
in 1918. Vernalization was not controversial. After being exposed to very low
temperatures, a process known as “cold shock,” agronomists planted various
grains; cold shocked seeds tended to show higher degrees of reproduction and
yield per seed than nonshocked seeds. By Lysenko’s time, vernalization was
known as “the acquisition or acceleration of the ability to flower by a chilling
treatment.” 6 Lysenko merely adopted vernalization and attempted to apply it to
his own agronomic work.
His big break came in 1928 at the Leningrad All-Union Congress of Genetics,
Selection, Plant and Animal Breeding. A well-known 1927 Pravda article had
already introduced Lysenko to the Soviet public; the article portrayed him as a
sullen, barefoot sage, a Tolstoy of the wheat field. After an especially poor
winter wheat harvest in 1927, Lysenko promoted the idea of keeping the kernels
of wheat moist and cool through the winter rather than burying them in the earth
in the autumn, where they would be destroyed by the exceedingly dry, frigid
Russian winter. His plan succeeded on a small scale, hence his invitation to the
All-Union Congress.
Pravda picked up the story of the 1928 All-Union Congress as well.
Coincidentally, the Soviet Union had just launched its first Five-Year Plan.
Again Lysenko appeared as an almost mystical figure, promising to increase
yields across the Soviet steppe. His practice of vernalization became official
policy surprisingly quickly—with mixed results. On individual farms, monitored
carefully, Lysenko’s process seemed almost miraculous. Years later, Soviets and
Westerners alike admitted vernalization was too difficult to control and
implement properly on the gargantuan scale required by Stalin. Yet early in the
1930s, few knew that the problems of scale would be so pronounced. At first,
everything worked, and these early successes resulted in an appointment for
Lysenko to head a whole department at the Odessa Breeding and Genetics
Institute.
In 1931, Lysenko came into contact with a biologist at Leningrad University,
Isaak Prezent. Prezent read more broadly than Lysenko and was versed in the
history and philosophy of biology. The two formed a close partnership and
together launched the Journal of Vernalization. It was under the tutelage of
Prezent that Lysenko began to look for the deeper biological processes
governing vernalization. Almost immediately, Lysenko and Prezent encountered
resistance from “orthodox” Mendelians in the Soviet Union. They chided
Lysenko for failing to apply rigorous statistical methods or even attempting to
keep abreast of the latest developments in breeding emanating from the UK,
Germany, and the United States. In many Western biology labs in the early
twentieth century, Mendelism (or the science of “hard” quantitative heredity) had
replaced “soft,” qualitative heredity associated with Lamarck or even Darwin.
Lysenko seemed blissfully unaware of any of these developments. 7
Ironically, Nikolai I. Vavilov (1887–1943)—who was by the 1930s an
internationally renowned plant breeder and geneticist at Leningrad—was not one
of Lysenko’s critics. As co-organizer of the Sixth International Congress of
Genetics held at Cornell University in the summer of 1932, Vavilov invited
Lysenko to present the vernalization work. It would have been an excellent
opportunity to convince mainstream Western biologists of vernalization. But for
whatever reason Lysenko ignored Vavilov’s invitation. Vavilov magnanimously
brushed off the slight. Later that summer, Vavilov toured North and South
America, collecting specimens for the Soviet global seed bank. Despite being
rebuffed by Lysenko, Vavilov openly praised Lysenko’s vernalization during his
trip as the method for “mastering individual variation.” Mastering variation had
been one of the major problems of breeding in the early twentieth century;
Vavilov recognized Lysenko’s legitimate scientific contributions to the issue. But
even while Vavilov traveled abroad, praising the Soviet agricultural program and
Lysenko’s vernalization, Prezent and Lysenko prepared their campaign against
Vavilov’s Lenin All-Union Academy of Agricultural Sciences (VASKhNIL). By
1935, Josef Stalin himself expressed pleasure with Lysenko’s work; not
surprisingly, the open political attack by Prezent and Lysenko on Vavilov and his
VASKhNIL commenced soon thereafter.
Vavilov’s VASKhNIL held a special session late in December 1936 to deal
with the swelling conflict. The two sides published their positions in a collected
volume entitled “Debatable Issues of Genetics and Breeding.” Here, Lysenko
made the audacious claim that his group had transformed a form of winter wheat
into a very successful spring varietal. The transformation happened nearly
accidentally and to a single related set of seeds. Lysenko’s group then more
deliberately transformed that now-spring wheat back into winter wheat. It was a
small sample, but from these results Lysenko and Prezent concluded that there
could not be a gene in the wheat that determined once and for all that it could
only grow under winter conditions, since it could switch back and forth. Nor
would it be logical to assume a gene for winter development randomly mutated
to become a gene for spring development and then randomly mutated back in the
other direction. For Lysenko, the apparent directionality of the change was the
important feature, and directionality was practically impossible under the
orthodox Mendelian regime. If orthodox Mendelism could not account for the
directionality, perhaps Mendelism was wrong. As Lysenko insisted, “The
hereditary basis does not lie in some special self-reproducing substance [e.g., a
gene]. The hereditary base is the cell, which develops and becomes an
organism.” 8 With this statement, Prezent and Lysenko were harkening back to
the timeless statement—first uttered by German physiologist Rudolf Virchow in
1858—that all cells derive from other cells and not from some less complex
thing.
Taken at face value, the differences between Vavilov and the Prezent-Lysenko
camp were not pronounced. Or, at least it would be difficult to predict given only
these statements that “Lysenkoism” would become among the dirtiest words in
American biology within a few years. That surprising transition came about
because of the heinous political machinations of Lysenko and Prezent, which
would result in the imprisonment and eventual death of Vavilov, followed by the
reactions of some of the most important and well-connected biologists in the
world at the time: H. J. Muller, Julian Huxley, and Theodosius Dobzhansky.
Not incidentally, all three of these men were deeply connected to each other
long before the “Lysenko affair.” As the high-flying grandson of T. H. Huxley,
Julian Huxley began his professorial career at the William Marsh Rice Institute
(now Rice University) in Texas. In 1916, Huxley lured Hermann J. Muller
(1890–1967) to Rice. Muller had already made a name for himself as a protégé
of T. H. Morgan and, backed by copious amounts of Texas oil money, Julian
Huxley tasked Muller with the creation of a world-class genetics laboratory.
Neither man could stand the Southern United States for very long, but their
friendship survived over a half century and traveled across several continents.
Huxley would return to the UK. Muller grew so tired of the culture and politics
at Rice that he returned to Morgan’s Columbia University in 1919. A few years
later, money and the promise of his own laboratory lured Muller back, this time
to the University of Texas at Austin. While at UT, Muller assisted with the
distribution of The Spark , a socialist-leaning student publication. Negative
sentiment surrounding Muller’s activities led him to leave the United States
entirely in the fall of 1932. From Austin, he landed in the Berlin-Buch laboratory
of Nikolai V. Timofeev-Resovskii, Russian director of the Department of
Experimental Genetics at the Rockefeller Foundation–supported Kaiser Wilhelm
Institute for Brain Research. Soon after the Nazi takeover in 1933, Muller fled—
once again with Huxley’s help. This time, Muller moved to N. I. Vavilov’s lab in
Leningrad. There, Muller witnessed both Vavilov’s legendary work ethic and the
inexorable turn of the tide against Vavilov’s Mendelist-Morganist, i.e.,
preformationist, genetics.
Understandably, Muller grew close to his Russian host. Because of Vavilov’s
influence, Muller was able to oversee the rigorous application of Anglo-
American genetic practices and theory to Vavilov’s VASKhNIL laboratory.
Muller also began preparing a journal, The Soviet Journal of Genetics ,
promoting the new work coming out of Vavilov’s lab. Furthermore, as much of
the official agricultural policy for the country passed through Vavilov in the mid-
1930s, Muller had a measurable impact on the direction of future research in
Russia. Or at least he might have. The presence of an American pushing
Mendelism raised the hackles of Lysenko, Prezent, and their followers; the fact
that Muller professed a strong affinity for Soviet principles made no difference.
Ironically, Muller’s own words became the ammunition that Lysenko and his
supporters co-opted to gun down Vavilov’s entire genetics program at Leningrad.
In 1936, Muller delivered his recently published novel, Out of the Night (1935),
to political and scientific figures across the USSR. Out of the Night is an
exuberant work of science fiction that praises the spirit of Bolshevism; Muller
naively believed Stalin would approve of it, and in fact had one personally
delivered to Stalin. It was rapidly translated and read, but Stalin was not pleased
with the novel. In it, Muller vigorously endorsed eugenics through a breeding
program reminiscent of—we can only assume coincidentally—that depicted in
Aldous (brother of Julian) Huxley’s Brave New World (1932). Stalin apparently
intimated that Soviet biologists should confront Muller at the 1936 VASKhNIL
meeting. There, Muller was permitted, even encouraged, to defend his eugenic
proposals. When given that opportunity, Muller denounced the fascist eugenics
program already being implemented by the Third Reich and tried to distinguish
his Communist eugenics program from the Reich’s. He insisted that with the
absence of class distinctions, as in the Soviet Union, eugenics could be properly
employed to make a fitter society. According to Muller’s hastily penned 9 March
1937 letter to Huxley, Soviet academicians responded favorably to his defense. 9
In reality, however, Muller’s eugenics was so closely tied to his genetics that
Stalin and most Soviet biologists conflated them—and rejected both. Muller
soon sensed that he was no longer welcome in the USSR and, early in 1937,
made plans to leave. Yet again, Julian Huxley pulled strings with the
international life science community and Muller was given a visiting position in
F. A. E. Crew’s Animal Genetics lab at the University of Edinburgh (the
laboratory that Waddington would later lead).
At Edinburgh, on the eve of the Second World War, Muller organized the
Seventh International Congress of Genetics (1939). He placed at the centerpiece
of the conference a “Geneticists’ Manifesto” written primarily by himself and
Huxley. That manifesto promoted exactly the same points he had promoted at the
VASKhNIL. First, he insisted race and class divisions must be put aside and
“economic and social conditions which provide approximately equal
opportunities for all members of society” must be adopted. Secondly, Muller’s
manifesto advocated that a “socialized organization” should be given appropriate
authority in the “conscious selection” of future human progeny; in the future,
humans should be bred to optimize health, intelligence, and “temperamental”
qualities. For now, that could only be done through aggressive reproductive
counseling and birth control, though in the future Muller trusted that more direct
genetic engineering would be developed. 10 The manifesto also introduced an
attack on Lysenko, Prezent, and their followers. According to the manifesto,
Lysenko’s major error was that he endorsed the fallacious notion that “the
children of parents who have had better opportunities for physical and mental
development inherit these improvements . . . in consequence the dominant
classes and peoples would have become genetically superior to the
underprivileged ones.” 11 As Muller revealed here, he saw the gene level as
permanent and deterministic; if the environment could mold the genes, the
temporary social and environmental conditions would become biologically
fixed. In other words, if Lysenko was right, unfair conditions imposed by the
dominant class would become fixed biologically into the very genetics of the
oppressed. Injustice would grow as a biological trait as well as a sociocultural
one. Muller could not stomach such a biology.
Vavilov’s delegation from the USSR was set to attend the 1939 Edinburgh
conference; Vavilov had been elected president of the International Congress, in
fact. But at the last minute, his team was barred from leaving the Soviet Union.
Tragically, only a few months later the schemes of Lysenko and Prezent came to
fruition: NKVD agents drove to Vavilov’s research station and beckoned for him
to get into their car. Once detained, he was interrogated, subjected to one of
Stalin’s show trials, and sentenced to death. Given his service to the Soviets, that
death sentence was commuted to twenty years of imprisonment and hard labor.
In a bitter irony, given that his lifelong mission was to fight future malnutrition
by preserving every extant crop species, Vavilov died in prison in 1943, likely
due to complications from malnutrition. 12
As their correspondence testifies, Muller, Huxley, Dobzhansky, and other
Western biologists close to Vavilov would not learn of these details for many
years. They did hear about Vavilov’s show trial in 1940 and feared the worst. By
the spring of 1947, they had accumulated enough evidence to deduce that
Vavilov had indeed died. Understandably, they were upset. Dobzhansky wrote to
Muller that “One should prepare and publish an appreciation of Vavilov . . . as a
martyr of genetics.” 13 Muller penned a few personal notes, but it was
Dobzhansky who prepared Vavilov’s moving obituary mostly by himself; it was
published in major scientific journals around the globe in the summer and fall of
1947. Upon reading the finished version, Muller praised Dobzhansky for “the
great service you have done in giving this record of this great man to the world.”
14
In the same letter, Muller also expressed to Dobzhansky his vehement desire
to scrub from the annals of biology the ideas of Lysenkoism that contributed to
the liquidation of Vavilov. For starters, Muller would turn his attention to the
flagship journal Science and root out any “anti-genetic” thinking he found there.
Muller felt his “gene theory” was under attack even among colleagues in the
United States, just as it had been at the 1936 VASKhNIL meeting. “It would be
intolerable for us,” he wrote, “to be defeated by Lysenkoism even in this one
publication, for it is an important one, and it would be, I think, well worth
fighting out the principle.” So in 1947—a year before Lysenkoism became the
official biological policy of the USSR and a decade after Muller left Vavilov’s
Leningrad laboratory—he began to seek out friends and colleagues to attack
Lysenkoism in scientific and popular publications. Waddington’s epigenetics
would suffer collateral damage as an unintended consequence.
Three divergent responses to Muller’s anti-Lysenkoism emerged. The first
group, mostly Americans, supported Muller. These individuals had no particular
love for the Soviet Union anyway and criticized Lysenko’s vernalization and the
state planning of science in any context with equal fervor. A second group,
mostly UK biologists, took a more nuanced approach. Several of these
individuals had actually advocated state planning of science during the Tots &
Quots years. Prolific Julian Huxley produced several articles and two books on
Lysenkoism before 1949; these largely concurred with Muller’s assessment but
were written in a far less vitriolic tone. Though Huxley dismissed Lysenko’s
scientific contributions, he was not led to the conclusion that state-planned
science automatically blazed the trail for bad Lysenkoist science—the
conclusion that so many of his American colleagues had deemed the only logical
lesson to be gleaned from this story. Contrast these positions with the third set:
the opinions expressed by four outspoken British scientists who were also
Communist Party members—J. B. S. Haldane chief among them. These four
appeared on a 1948 BBC radio debate regarding Soviet science. Haldane asked
his audience to suspend judgment about both Lysenko’s biology and the status of
state planning of science even while disagreeing with the Soviet government’s
treatment of Vavilov. Ultimately, J. B. S. Haldane insisted, the Soviet scientific
establishment had shown itself to be rational and science centered in the past. If
Lysenko produced practical improvements to agriculture, then the anti-
Mendelian ideas that backed up his work must have some value, even if it was
not obvious from the West’s standpoint. So said one of the foundational figures
of the neo-Darwinian synthesis.
Muller was outraged with Haldane and the British Left, and he promptly
shared his reaction with Dobzhansky in the United States. 15 Muller’s take on the
situation became the standard line of attack for many American geneticists in the
wake of the Lysenko affair—and remains so for many scientists and historians of
science. 16 According to Muller, the problems with Lysenkoism couldn’t be
overcome even if the Soviets could demonstrate agricultural improvement over
the short run. Evidence wasn’t enough: The issue was the philosophy itself.
Lysenko’s claim that the environment could manipulate the growing process
permanently was merely Lamarck reincarnated, the inheritance of acquired
characteristics concept from the early nineteenth century. No real scientist could
believe in that, according to Muller, though he feared that some geneticists might
be corrupted by Lysenkoism, as Haldane clearly was.
Muller’s gene theory stood in direct contradiction to the biology promulgated
by Lysenko and his followers. First, Muller’s genes had to be discrete chemical
packages, likely made of chromatin, strung like pearls upon a string and wound
up into chromosomes, then crammed inside the nuclei of cells. Secondly,
Muller’s theory demanded that these genes were the necessary and sufficient
determinants of traits including, for example, eye color, race, intelligence, or
grain yield during certain seasons. The particular environment provided the
context for expression of the trait, but not the information for making that trait. 17
Thus, as one later retelling of the story goes, Lysenkoism lacked “connection
with any facts whatsoever” and was instead “an empty, superficial piece of
publicizing” primarily because it disagreed with Muller’s two ideas about genes.
18
Muller’s fears notwithstanding, little evidence existed that the American
biological community had warmed toward Lysenkoism at all, let alone become
as tolerant as Haldane. Quite the opposite in fact. Even advocating that the
Soviet concepts should be weighed thoughtfully before being dismissed out of
hand became professionally dangerous. In 1949, two Oregon State University
professors were dismissed for asserting that Lysenko’s vernalization “had not
had a fair examination in the US.” 19 Even this hint of open-mindedness toward
Lysenkoism drew an angry reaction from Oregon State University president A.
L. Strand:

Many men in Soviet Russia . . . have died in concentration camps, or by other means, because they
would not accept the untruths that Dr. Sprizer [one of the dismissed Oregon State scientists] has
chosen to espouse . . . Dialectical Materialism! A better name would be dialectical murder. . . . Any
scientist who has such poor power of discrimination as to choose to support Lysenko’s . . . genetics
against all the weight of evidence against it is not much of a scientist, or, a priori , has lost the
freedom that an instructor or investigator should possess. 20

President Strand may have been speaking hyperbolically, but clearly it was
unwise for any American scientist to come out too strongly in favor of any of the
main features of the Lysenko story.
By the 1950s, the mention of Lysenko invoked three features that many
American biologists, funding organizations, political figures, and university
administrators rejected out of hand: centralized planning of science in the social
interest, an attachment to dialectical thinking, and more emphasis on the role of
environmental conditions in the process of development and evolution.
Advocating any one of these things might get a scientific concept labeled as
friendly toward hated Lysenkoism. For better or worse, C. H. Waddington
argued explicitly for all three of these tenets in the 1940s. He continued to
support some version of these three concepts through the ’50s, even after he had
visited Lysenko’s Leningrad laboratory—even after he had begun to promote
epigenetics as the science of the “third way.” 21

Just as he had ten years earlier, Waddington appealed to Whitehead’s notion of

canalization in his best-known paper of his first years at Edinburgh, entitled
“Genetic Assimilation of an Acquired Character.” 22 Given the political context
of the early Cold War, his title alone signaled that he might be willing to at least
investigate the scientific claims of the Lysenkoists, despite the fact that he made
no direct claims about Lysenko at all. He did place some experimental data from
his laboratory in Edinburgh alongside the older reports about ostrich callouses he
had cited before the war. The evidence wasn’t terribly new by that point; over
the early 1950s, Waddington’s lab group essentially recapitulated some of the
developmental biology experiments he and Needham had conducted in the 1930s
—mostly interfering with the development of embryos and studying the effects.
After his experience working with Dobzhansky, among others, at Caltech in
1938, Waddington’s group adopted the American laboratory custom, using
Drosophila as their model species along with birds and amphibians.
Waddington’s goal during the 1950s was not to define the nature of a
particular region of the embryo as it had been in the 1930s, but to understand
two seemingly opposing processes: (a) how dramatic changes could arise in
adult forms from small manipulations in the embryos, and (b) how
manipulations to genotypes and from the environment might produce little or no
changes in the adult form. Of course, work on process (a) was nothing new or
even noteworthy in the 1950s. In Morgan’s lab, Alfred Sturtevant had crossed
and recrossed mutant flies since the early decades of the twentieth century;
Muller had bombarded them with x-rays to see the same thing.
Unlike Sturtevant and Muller, however, Waddington was less convinced that
genes were discrete “molecules of life.” So he put dramatic change, process (a),
aside for a moment to study process (b), the ability for organisms to produce
“normal” versions of themselves even when environmental and genotypic
alterations suggested there should be some alteration in traits. What he learned
was that subjecting Drosophila pupae to high heat for a few hours—heat shock
of fruit flies, comparable to Lysenko’s cold shock of wheat grains—produced
abnormalities in the veins of their wings. Not all flies produced veins across their
wings when subjected to the heat shock. In his control group, Waddington
selected the flies at random, breeding the next generation and the next, all the
while subjecting them to heat in the pupal stage. In one experimental group he
selected those individuals that did display the wing abnormality and subjected
some of their offspring to the same environmental extremes. This experiment
continued for dozens of generations. Interestingly, after selection and reexposure
over a period of only fourteen generations, the wing abnormality appeared
regardless of the heat being administered to the pupae. In other words, after
several generations—but still relatively few—the “acquired” character of wing
abnormalities (known as crossveinless ) became fixed, the norm.
Perhaps the disappearance of veins in the wings of fruit flies did not seem
like anything dramatic. Waddington even admitted that he was unsure what the
survival value of crossveinless would be in the wild. That hardly mattered,
however. What these experiments showed was that ordinary phenotypes seemed
to resist change to perturbations in their environment, even large ones introduced
during the course of development; yet, if environmental change was large and
persistent enough, organisms can be altered quickly and decisively. But why this
way? According to the traditional Darwinian understanding, traits should change
almost imperceptibly slowly over eons. Neo-Darwinians in the wake of Muller
and Watson and Crick insisted that organismal change occurred due to mutations
in the purine and pyrimidine rungs of the twisted ladder of DNA. Genes had to
change, in other words, for traits to change, and they did so indiscernibly.
Waddington believed the crossveinless experiments showed something else was
happening. The full component of genes—the genome—worked together to
create not one phenotype but a range of possible phenotypes. Only close and
constant interaction with the environment during development determined the
actual phenotypic fate—the path taken on the route to adulthood. Environmental
conditions might shift an organism dramatically away from its typical phenotype
to an alternative one. Genes provided the possibility for this, but they did not
dictate which form an organism would actually take. Only an investigation of the
epigenetic factors involved in translating the genotype would unlock the
phenotypic possibilities.
This interpretation stood in opposition to the implicit preformationism held
by nearly all geneticists working at the time. When, toward the end of his life,
Waddington reflected on the situation in the 1950s, he realized that there were
practical effects to the philosophical differences between his gene-centric,
mechanistic colleagues who worked in genetics and molecular biology and his
own organicist perspective. The mid-twentieth century was the heyday of the
neo-Darwinian synthesis and the adolescence of molecular biology, of course.
Epigenetics fit a different mold than those pursuits: “Two rather radically
different lines of approach were followed. The one which was—and in fact still
is—favored by most geneticists depended on what I think may be called
‘atomistic’ metaphysics. It set out from the assumption of genes, and it asked, at
first, what does ‘A’ do, and later, what controls whether gene-A is active or not?”
23 A huge group of studies were conducted in this vein, several of which led to

the award of a Nobel Prize for the scientists involved. H. J. Muller, one of those
Nobel winners, specified that genes are “autocatalytic,” that is, autonomous self-
starters and self-replicators. In the 1940s, George Beadle (who penned the
introduction for Asimov’s Intelligent Man’s Guide ) and Boris Ephrussi
identified the substance produced by any given gene as an enzyme. This led to
one of the most lasting of the manifold functional definitions of genes: “one-
gene = one-enzyme.” Hans Grüneberg at University College London
manipulated mice genes to study their development and, by the middle of the
Second World War, concluded that one-gene might also equal a number of
developmental consequences. As the work continued along these lines through
the Watson and Crick work and afterwards, Waddington could fairly say that the
approach focusing on discrete particles of life had proved to be a fruitful one.
It didn’t describe nature as found outside of the laboratory very well,
however. What we actually observe are cells, cells becoming specific things,
performing specific functions in the kidney or brain or blood. Convinced that the
organic philosophy gave a more faithful rendering of the world as we encounter
it, Waddington “began developing the Whiteheadian notion that the process of
becoming (say) a nerve cell should be regarded as the result of the activities of
large numbers of genes, which interact together to form a unified ‘concrescence.’
. . . [By the late 1930s] it had become apparent that the ‘gene-concrescence’
itself undergoes processes of change; at one embryonic period a given
concrescence is in a phase of ‘competence’ and may be switched into one or
other of a small number of alternative pathways of further change.” 24 The
alternative pathways of further change were the canals first rendered artistically
by John Piper in Organisers and Genes (Figure 11.1 ). The concrescence space
was Waddington’s “epigenetic landscape.” It ended up being among the most
lasting concepts of his career. 25
Given the resistance to thinking about fields and wholes rather than particles
that Waddington sensed during his visit to T. H. Morgan’s lab in 1938, he knew
that he would have to cloak his support for organicism. Geneticists, he learned,
were reluctant to pry open the black box of development. When he spoke with
those on the other end of the spectrum, embryologists like Yale University’s
Ross Harrison and Oxford’s Gavin De Beer, he noted that they were reluctant to
cede any additional intellectual territory to genetics—to say nothing of funding.
The two groups stood opposed to each other in technique and philosophy. Over
the 1950s, Waddington worked to bridge both genetics and embryology as he
and the Needhams together had attempted to build common ground between
biochemistry and embryology during the organizer years. In 1957, he published
The Strategy of the Genes , a summary of the two decades since Organisers and
Genes ; in it he included the most well-known depiction of the epigenetic
landscape (Figure 11.2 ).

Waddington’s epigenetic landscape would have to serve as a neutral epistemic

object, allowing geneticists and embryologists to work together on the
mechanics of development. His hope was that the different levels of organization
at the genetic and embryological/developmental levels would retain their
distinctiveness while still allowing for interlevel analysis—just as Woodger had
proposed in TBC meetings and Needham had codified in Order and Life (1936).
The epigenetic landscape would serve as a model for truly organicist research:
Genetics would have a key role, but so would development, so would the
environment. Harkening back both to discussions with Dorothy Wrinch at the
TBC and to his own Cambridge training as a geologist, Waddington usually
presented the epigenetic landscape as a three-dimensional topographic drawing
with peaks and valleys. The whole surface was then given a slight tilt toward the
viewer. In his more extensive treatments, through the 1950s and ’60s,
Waddington illustrated through a number of panels the movement of a ball down
the surface through various switching points toward one of a number of eventual
conclusions. 26 The “ball” indicated a developing system, a whole organism, or
large portion of an organism. The “valleys” and “hills” were the “canals”—
pathways that the developing system was likely to take toward its conclusion as
an adult organism unless perturbed. The height of the hill and depth of the valley
indicated the degree of “buffering” against perturbation, i.e., the likelihood that
the organism/system would revert to its “normal” adult phenotype despite
environmental stress (Figure 11.3 ).

It was only a small step from the epigenetic landscape to an alternative theory
of evolution. While other neo-Darwinian biologists emphasized the changing
aggregates of genes, with evolution being based on fortuitous mutations to those
particles (Muller) and allowed to flourish by reproductive isolation (Ernst Mayr),
Waddington stressed that what got passed along from parents to children was a
set of developmental patterns, of potentialities : “You do not inherit fair or dark
hair, blue or brown eyes, from your parents; what you inherit is something which
endows you with the capacity for developing eyes of a particular colour under
certain particular circumstances, and perhaps a different color under other
circumstances. . . . Nowadays we use the word ‘genotype’ for the collection of
potentialities which are inherited. . . . Any one genotype may give rise to many
somewhat different phenotypes, corresponding to the different environments in
which development occurs.” 27 The developing organism should be thought of as
a field, a possibility space, claimed Waddington. Genes laid out the structure, the
geography of that possibility space, but they alone did not define traits. The
passage between genotype and phenotype was much more complex than what
was commonly taught in genetics courses or promoted outside of the laboratory
in the mid-twentieth century. Moreover, as Whitehead suggested in the 1920s, as
the TBC members had emphasized through the 1930s—and as Bertalanffy was
now sketching it in the 1950s—the concepts of physics itself had also shifted
away from atomic concepts and toward what Waddington called “continuum
theories, that is to say, theories in which the basic constituents of phenomena are
not thought of as separate and discrete entities which enter only into external
relations with one another.” 28
Thus even if biology could be reduced to physics and chemistry, as Nagel
hinted it might one day be, that reduction would be toward continuum theories,
concepts where properties of the space arise from the relationships, not from
properties intrinsic in this or that atom. The epigenetic landscape would survive
reduction. It would survive Lysenko.
 12

ERNST MAYR, NEO-DARWINISM, AND

BEANBAG GENETICS

W addington’s epigenetics did not sweep the field after the publication of The
Strategy of the Genes. It did make an impression on some significant individuals,
including one of the foremost neo-Darwinians, Ernst Mayr (1904–2005), who
had just put his hand to the plow writing the history and philosophy of
evolutionary biology. Influenced by Waddington’s work, Mayr delivered one of
the most influential warnings against the gene-centric mechanistic thinking that
he saw spreading into evolutionary theory. 1 Yet Mayr also found Waddington’s
work too near to the same Lamarckian interpretation of evolution highlighted by
the Lysenkoists. In the context of the Lysenko affair and the solidification of the
neo-Darwinian consensus of the 1950s, Waddington’s work seemed dangerously
out of sync with mainstream biology. Only many years later would Mayr return
to render a favorable account of organicism, never mentioning Waddington or
his epigenetics in connection to it.
The year 1959 is often remembered as the centennial of Charles Darwin’s On
the Origin of Species and there were then, as in 2009, many celebrations of
Darwin’s life and the theory he formulated. For geneticists, however, the most
memorable conference of that year was the Cold Spring Harbor Symposium on
Quantitative Biology. From June 3rd to 10th, the most prominent geneticists in
the world gathered to celebrate “Genetics and Twentieth-Century Darwinism.”
The symposium would emphasize, at least according to Milislav Demerec, the
well-known director of the Cold Spring Harbor Laboratory and one of the
organizers of the conference, “evolutionary problems related to genetics.” After
all, as Demerec reminded his audience, “Early geneticists were aware of the
powerful techniques for the study of evolution which were provided by the
discovery of Mendelian inheritance. The best illustration of their optimism about
these potentials is the fact that the first laboratory devoted to the study of
Mendelian inheritance was named ‘Station for Experimental Evolution.’” 2 And,
according to Demerec, the luminaries of evolutionary biology in the early
twentieth century were population geneticists: J. B. S. Haldane, R. A. Fisher, and
Sewall Wright.
The majority of the twenty-five papers that followed Demerec’s opening
address nicely reiterated this framing stance. In order to understand the process
of evolution that motivated Darwin, twentieth-century biologists must reckon
with and, if at all possible, experiment on issues of heredity. As modern
synthesis architect G. Ledyard Stebbins summed up the meeting, any other
interpretation would fly in the face of an international body of professionals, all
of whom “equipped with the results of a sum total of several hundred man-hours
of effort devoted to factual research and theoretical contemplation of
evolutionary problems, have reached substantial agreement on concepts
remarkably similar to those which Darwin himself held, not only about the
existence of evolution and the course which it followed, but also about the basic
processes responsible for it.” 3 Strangely enough, according to Stebbins and other
speakers, after generations of rigorous experimental research on evolutionary
problems using fantastically modern equipment resembling that found in the
laboratories of physicists and chemists, this biologist proudly asserted that their
field had returned full circle to the perspective advocated by Darwin—that
gentleman naturalist who worked with orchids and earthworms in his own
humble greenhouse a century earlier. “The only major qualitative difference
between our knowledge and that possessed by Darwin,” stressed Stebbins, “lies
in our recognition of particulate Mendelian inheritance, determined by
chromosomal genes, as the basis of nearly all of the hereditary variability upon
which selection acts.” 4 From the perspective of Stebbins, as well as the majority
of participants in the 1959 Cold Spring Harbor symposium, nothing in
evolutionary biology now made sense except in light of population genetics.
Even symposium papers having seemingly little to do with population genetics
(Finnish paleontologist Björn Kurtén’s study examining fossilized bear teeth, for
instance) were recast as reinforcing the population genetics consensus
interpretation. 5
Ernst Mayr cut across the grain of the symposium with his deliberately
contentious address entitled “Where Are We?” 6 Mayr asserted that if mid-
twentieth-century synthetic evolutionary theory seemed more authentically
Darwinian than even (paradoxically) Darwin himself had been, biology had
organismal naturalists like Mayr to thank for it rather than any set of geneticists.
Geneticists, in fact, had nearly led the whole venture of evolutionary biology
down the wrong path. While “Mendelians,” including William Bateson, Hugo
DeVries, T. H. Morgan, and H. J. Muller, assumed evolution through speciation
only occurred as often as “occasional lucky mutants” appeared ex nihilo,
“classical” population geneticists Fisher, Haldane, and Sewall Wright (who was
in attendance) attributed to each gene an “absolute selective value”—something
no self-respecting zoologist would ever attribute to a gene. In short, both groups
committed vast reductive oversimplifications. Even those who were being
honored by this very symposium were guilty of using mathematics to present
evolution “as an input or output of genes, as the adding of certain beans to a
beanbag and the withdrawing of others.” Mayr deemed their sin—and it was
cardinal—“beanbag genetics.” 7 He intended the term to parody the territory of
some in attendance at the 1959 symposium who, despite their obvious
mathematical genius, deigned to work among flowers and fruit flies while
keeping their hands mostly out of the dirt and on their calculators.
Mayr implicitly revealed his reason for using this term of derision openly
when he eventually managed to praise, backhandedly, the Fisher-Haldane-
Wright (hereafter cited as FHW) triumvirate. “Perhaps the main service of the
mathematical theory,” he speculated, “was that in a subtle way it changed the
mode of thinking about genetic factors and genetic events in evolution without
necessarily making any startlingly novel contributions .” 8 If anything, Mayr
surmised that the population genetics of the 1920s and ’30s merely provided
something of a salvageable, if wrongheaded, foundation for his own
contemporary views. Why then bother to denigrate it, if it was barely adequate?
Simple: It was the lingua franca of the 1950s and beyond—the approach lauded
by both scientific insiders such as this distinguished audience and popularizers
like Isaac Asimov. Mayr recognized that this “beanbag genetics” had become
both publicly and professionally persuasive in a way that his own whole
organism work conducted in a museum was not. Genetics in the mid-twentieth
century rendered biology sophisticated, exclusive and, consequently, more like
the physics used in the Manhattan Project or Bell Labs than anything else in
evolutionary studies. As this version of FHW genetics gained explanatory power,
Mayr’s own field, which regarded the organism and the breeding population as
the fundamental units of selection rather than the gene and the gene pool,
diminished in importance and prestige. 9
Thankfully, according to Mayr, the “newer” population genetics practiced by
Dobzhansky, among others, rescued evolutionary biology from being swamped
by abstract mathematics. This “advanced” version of population genetics
stressed the interaction of genes rather than their discrete particulate nature: “Not
only individuals but even populations were no longer described atomistically as
aggregates of independent genes in various frequencies, but as integrated,
coadapted complexes. A gene is no longer considered to have one absolute
selective value, but rather a wide range of potential values that may extend from
lethality to high selective superiority, depending on genetic background and on
the constellation of environmental factors.” 10 In addition, because Dobzhansky
had focused for a decade on naturally occurring populations of Drosophila
pseudoobscura , rather than merely the laboratory favorite D. melanogaster ,
Mayr insinuated that the new, purportedly holistic, population genetics was
closer to describing evolutionary truth than the much-touted FHW “synthesis”—
now reduced to beanbag status. 11
Yet we should not be fooled by the force of Mayr’s symposium rhetoric. In
his promotion of a kind of evolutionary theory that privileged different levels—
genetic, epigenetic, organismal, and even populational—Mayr was not joining
the ranks of Waddington, the 1930s TBC, or other 1950s “third way” biologists
like Bertalanffy. Rather, Mayr was attempting to chart a particularly delicate path
between two unacceptable possibilities. The modern synthesis as Mayr depicted
it would be neither mathematical abstraction that, by necessity, according to J. B.
S. Haldane, relied upon a recognized oversimplification of genes and
populations. Nor would it be a version of biological evolution that weakened the
role of selection or challenged the reigning (largely through Mayr’s own efforts)
gradualistic account of the origin of species in favor of jumpy, environment-
infused epigenetics. 12 “Beanbag genetics” was Mayr’s rebuttal of the former
possibility—that he probably attacked a straw man mattered little. 13 But on the
other side, Mayr saw Waddington’s epigenetics, and that seemed too invested
with Whiteheadian speculation for Mayr to even comment on, let alone
extensively attack. In fact, though Mayr shared the sentiment of Waddington’s
critique of the undue influence of FHW genetics on biology as a whole, he did
not sympathize with Waddington’s description of a complete evolutionary
biology account for “Mind, Form, and End.” 14 Mayr agreed that Waddington put
proper emphasis on the “End” part of that set; explaining the seemingly designed
“fit” between organism and environment as a wholly secular phenomenon was,
after all, one of Mayr’s chief aims. In the study of “Form,” however—a topic
Waddington had just addressed in The Strategy of the Genes —Mayr saw too
many “metaphysical concepts that have no place in our modern thinking.” Form,
as he stressed to Waddington shortly after the Cold Spring Harbor symposium,
was a peculiar philosophical hang-up of embryologists. 15 If Mayr thereby
distanced his own theories and alliances from those used by embryologists, their
metaphysics was to blame, not his modern neo-Darwinian synthesis.
 Mayr’s 1959 “beanbag genetics” jeremiad pertains to the larger story about
the decline of organicism and the marginalization of Waddington and
epigenetics. While “Where Are We?” acknowledged the technical
persuasiveness of mathematical population genetics and warned against the
allure of the oversimplified, atomistic genetic picture it generated, Mayr stopped
short of endorsing either organicism or explanations of evolution in terms of
epigenetic change, in part because of his hardened commitment to gradualism
and rejection of what he called “essentialism.” 16 Only years after Waddington’s
death, Mayr would reconsider organicism—somewhat. 17

Through the urgings of Dobzhansky, among others, Waddington received an

invitation to the second major American conference in 1959, the Darwin
Centenary at the University of Chicago. This well-known meeting became the
most prominent celebration of its kind in the world and a defining moment in the
storied history of neo-Darwinism. There, Waddington rubbed shoulders with
prominent biologists, geologists, paleontologists, philosophers, and
anthropologists from across the globe. Yet the attacks on the organic philosophy
were increasing, and Waddington’s philosophical interests in process, relations,
fields, and the workings of the organism as a whole still found too little purchase
among his peers and the public alike—at least the public that could do
something about it.
Throughout the twentieth century (and now into the twenty-first), Darwin
anniversaries have served as points of disciplinary soul-searching. Scholars at
the 1909 Darwin celebration held at Cambridge University, for instance,
wavered in their advocacy of Darwinian natural selection, with William Bateson
playing the role of Mendelian dissenter. Even so, all members taking part in that
commemoration agreed that Darwin’s advancement of the theory of the
fundamental unity of all organisms through “descent with modification”
represented a monumental philosophical shift in biosocial thought. And all
relished the enlargement of this point to unexplored areas, especially in
“hereditarian” biology, which was in the middle of its own Mendelian
revolution. 18
The University of Chicago’s Origin of Species centenary in 1959 likewise
underscored the future trajectory of evolutionary biology while shoring up one
particular historical narrative of its conceptual roots. As in 1909, this “Darwin
Year” celebration billed itself as something beyond an insular, academic
scientific endeavor. 19 Though the celebrations were localized at particular
universities, it is true that representatives from both inside and outside the walls
of the academy attended and played major roles. In contrast with their centenary
predecessors a half century before, however, the organizers of the 1959 Chicago
celebration did not draw attention to the fundamental disagreements between the
various evolutionists. Dissensus reigned in 1909. By 1959, sessions were
organized primarily along the lines of consensus established by Sol Tax, the
University of Chicago anthropologist, after numerous consultations with his
committee. That committee included Alfred Emerson of the Society for the
Study of Evolution (SSE) and the less formal but just as influential evolutionary
biology groups surrounding Mayr, Simpson, and Dobzhansky. 20 Having
organized a massively well-attended event—in truth, a heavily advertised public
spectacle—Tax and his committee did everything in their power to avoid major
confrontations between the scientists. 21 For the most part, they succeeded—
almost too well.
The 1959 Chicago Darwin centenary featured several panel discussions
comprised of major scientists interspersed between large public lectures and
smaller presentations. But the emphasis of the conference was clear: “The
Celebration was the Panel Discussions,” quipped Tax. Seeded with “highly
individualistic” scientists, they might have been “dangerous and unpredictable
firecrackers.” But by taking certain experts out of the areas of their expertise,
Tax and the conference organizers forced specialized scientists to pronounce as
generalists. The problems they were asked to speculate on were “beyond the
specialties in which any of them [felt] comfortable.” Then these scholars were
asked to arrive quickly at more or less set topics for conversation upon which
they would largely agree. They were paraded onstage under the glare of intense
stage lighting in front of well over a thousand expectant people. 22 Under these
conditions, the likelihood that these eminent scholars would pronounce
forcefully on topics they knew little about was minimal. Perhaps without even
realizing the effectiveness of their tactics, the Tax’s Committee defused potential
conflicts.
Convocation speaker Julian Huxley’s address delivered on Thursday,
Thanksgiving Day, November 26, 1959, from the pulpit of Rockefeller Memorial
Chapel, did raise eyebrows, however. A vision of his famous grandfather,
Huxley furiously extolled the “evolutionary vision”—the recognition that natural
selection had created enculturated creatures, partly inhabiting the “nöosphere,”
capable of going beyond the limitations of natural selection to create and then
challenge “alarming monsters” of war, overpopulation, Soviet communism,
poverty, and the ability to produce enough food for all but without the
concomitant ability to deliver that food to those who need it. As many skeptics
noted, it was a secular sermon delivered with more passion, and more socialist
subtext, than usually rang from the Rockefeller Chapel rostrum. It was also
shamelessly atheistic (or maybe pantheistic), embarrassing the University
chancellor, Lawrence Kimpton, who had introduced Huxley. Even in this secular
university, gathered to celebrate Darwin, the public responded tepidly at best. 23
Waddington left the Chicago meeting mildly frustrated, but it had nothing to
do with Huxley’s “sermon.” As he wrote friend Margaret Mead just after the
event, he found the “party line” foisted on participants from all directions at
Tax’s event irritating. His own presentation on “ethics” went off marginally
better than he expected. Or at least his contribution was “original, which
practically nothing else there was.” 24 But the impact of his contribution was
muted. Perhaps not surprisingly, Waddington was not included in the headlining
discussions on heredity. Instead he was assigned to two panels still further from
his expertise, one on paleoanthropology, the other on the rapprochement between
science and religion. Waddington had little professional experience with either of
these issues—just as Tax’s committee had planned. 25 On the G. G. Simpson–
chaired panel on “Man as an Organism,” Waddington spoke exactly three times.
Not once did the other participants follow up on his comments. In fact, following
his last comment, which included a brief exposition of epigenetics, Simpson
deliberately and decisively changed the topic. 26
Panel members were asked to contribute to the three-volume proceedings.
Waddington’s written contribution to the Tax compendia published in the early
1960s by the University of Chicago was to be on “Darwinism and Sociology,” a
discussion he initially based on Margaret Mead’s insights on the “human [i.e.,
psycho-cultural] transmission mechanism.” 27 But his actual submission to the
first Tax volume turned out to be something closer to his epigenetic work.
“Evolutionary Adaptation” was included in the thick first volume, including
papers by the other modern synthesis architects who dominated the conference.
Not surprisingly, his essay contrasts boldly with the articles bracketing it. Just in
front of Waddington’s piece, Ernst Mayr expounded upon his views of
gradualism, geographic isolation, and pan-selection; following it, Dobzhansky
rehashed stripped-down neo-Darwinism, stating categorically, “The elementary
evolutionary events are changes in the incidence of genes in a population.” 28
Waddington’s contribution, on the other hand, warned that biologists of all
stripes

have perhaps been tempted to oversimplify our account of the mechanism by which evolution is
brought about. This mechanism—the evolutionary system, as it may be called—has often been
 envisaged as consisting of no more than a set of genotypes which are influenced, on the one hand,
by a completely independent and random process of mutation and, on the other, by processes of
natural selection which again are in no way determined by the nature of the genotypes submitted to
them. Perhaps such a simplification was justified when it was a question of establishing the
relevance of Mendelian genetics to evolutionary theory, but it can only lead to an impoverishment
of our ideas if we are not willing to go further, now that it has served its turn. 29

Waddington then followed with his discussion of four interrelated systems—
genetic, natural selective, epigenetic, and exploitive—to help theoreticians
“escape” from the “uncomfortable position” that too heavy a reliance upon
single reductive factors had forced upon evolutionary biology (Figure 12.1 ). 30
Examining the other contributions to the Tax volumes, it is clear that
Waddington’s plea for the inclusion of epigenetics in the larger evolutionary
synthesis fell on deaf ears, as Viktor Hamburger would point out in the late
1970s. 31
Why did Waddington change the topic of his essay submitted to Tax for
inclusion in the three-volume proceedings? And why was he placed on panels far
from the issues he wrote about in his written contribution? Two events may have
altered the course of both the conference itself and the multivolume proceedings:
(1) an abrupt change in conference leadership, leading to (2) greater involvement
of Ernst Mayr in molding the panels. During that leadership transition,
Waddington’s critique of neo-Darwinism was disregarded.
The first event transpired because of the death of Karl Schmidt. Schmidt, an
anthropologist at the Chicago Natural History Museum, had contacted
Waddington in September 1956, inviting him to two Darwin conferences: The
first, held “in advance of the Centenary proper late in 1957 or early 1958,”
would allow evolutionary theorists to work out the variations in their
interpretations of evolutionary theory in advance of a major publication. The
resultant volume would be a collection of thirty-five essays drawn from
“biology, anthropology, and the history of science” published with
commentaries. These preparations would permit the 1959 Chicago Darwin
centenary to “be devoted largely to an exploration of trends of investigation and
of thought to be expected in the century to follow, and to an evaluation of
Darwin’s work in the light of modern knowledge.” 32 Schmidt had a difficult
time procuring funds and, by early 1957, he dropped the idea of a specialized
pre-conference. Nevertheless, Waddington remained “high on our list on account
of the specific bearing of your own work on the basic problem of adaptation to
environment.” 33

 The structure of the Chicago Darwin celebration and Waddington’s role in it
began to change immediately after Schmidt’s death late in 1957. Sol Tax alerted
the University of Chicago Darwin celebration participants to the abrupt change
in the structure of the entire project in December. Rather than a major meeting of
scientists and others to discuss the status of evolutionary theory preceding the
public celebration, Tax informed participants that the Chicago celebration would
be the sole event, that it would follow a more typical conference with set paper
readings, and that the collected papers would follow, rather than anticipate, the
staged event. Also, rather than following Schmidt’s list exclusively, Tax hinted
that the 1959 meeting would draw participants familiar to him from the
Washington Anthropological Society’s 1957 meeting. For the most part, these
were physical anthropologists who belonged to a different generation than
cultural anthropology’s Boas-trained luminaries, such as Waddington’s friend
Margaret Mead (who was not invited). 34
Schmidt’s death and Tax’s assumption of leadership led to a second event,
namely, an increasingly influential role in the conference for synthesis architect
Ernst Mayr. Waddington made two hasty notations directly on Tax’s December
1957 letter explaining conference changes: “embryology” and “ethics.” In his
correspondence, he expressed disappointment that these two areas—“the
influence of evolutionary theory in embryology, and the impact of evolutionary
thought on ethical philosophy”—were not directly addressed in Tax’s new
scheme, as they had been in Schmidt’s. 35 Few concessions were made to address
Waddington’s concerns, though the “Darwin Year” planning committee
eventually put together a panel on “man’s unique characteristics.” (Julian Huxley
asked Tax to put Waddington on this one. 36 ) From that platform, Waddington
found it inopportune to offer a cogent “third way” critique of neo-Darwinism. He
did make other attempts to strike up debate around these issues. After reading
Mayr’s contribution to the collected papers of the Darwin conference,
Waddington sent a copy of his critical comments to Mayr and Tax. But these
never became public and, due to Tax’s loose oversight of the three-volume paper
collection, Mayr never acknowledged, much less addressed, Waddington’s
comments. These two features help to account for some of the reason why
alternative views of evolutionary theory (such as that forcefully promoted by
philosopher Marjorie Grene just before the Darwin celebration) received little
attention. 37
Tax’s real hope for the ’59 Chicago Darwin centenary was not merely to offer
a stage for the architects of neo-Darwinism. He strongly desired his version of
anthropology to have a place at the table in the inevitable unification of social
science with the life sciences under neo-Darwinism. He believed that nature
ultimately dictated nurture—a statement he suspected was included in neo-
Darwinism—and that most anthropologists would slowly adopt this view. 38
Tax’s prediction was both closer to, and further away from, the actual
developments in anthropology and biology proper following the 1959
celebration.
One omen of the future appeared at the ’59 centenary panel devoted to
anthropological questions. Here, H. J. Muller challenged the most sensitive
tenets of the creed of human cultural exceptionalism implicitly held by
generations of Boasian anthropologists. Though some members of his Panel
Five, “Social and Cultural Evolution,” assented to the notion that “cultural
evolution in Homo sapiens is essentially independent of gene differences,”
Muller roundly condemned this sort of thinking. 39 And, as a final punctuation
mark, the panel discussion closed with Julian Huxley’s pronouncement that the
main thrust of future anthropology should be toward “improving our genetic
heritage.” 40 It is indicative of how far the gene-centric view had penetrated even
the social sciences by midcentury that the last word regarding human evolution
should be given not to a cultural anthropologist but to two biologists with
unconcealed eugenic aims. 41
 13

HISTORY OF SCIENCE IS WRITTEN BY THE

LAUREATES

E pigenetics and the organicism upon which it was based faced more than
oblique snubbing by conference organizers and otherwise sympathetic
evolutionists like Mayr. In the decade after he published The Strategy of the
Genes , Waddington experienced two disconcerting and very specific dismissals
of his work. Two of the most prominent scholars at the time, both his
acquaintances, inspired these critiques. In the first case, Peter Medawar
encouraged Harvard historian June Goodfield Toulmin to write a history of
biology that would contextualize the Cambridge organizer experiments as
failures specifically because of the organic philosophy motivating them. In the
second, Francis Crick sparred with Waddington over the degree of importance of
epigenetics in the face of Crick’s molecular biology. Together, these episodes
reveal the extent to which the grand narrative of the history of biology in the era
of molecular genetics had already been set midcentury—and the extent to which
Waddington, epigenetics, and the “third way” more generally was relegated to
the “losing” side of that narrative.
In March 1965, June Goodfield Toulmin wrote Joseph Needham and
Waddington inquiring about the “growth and downfall” of the organizer project
in the 1930s. She was initiating research for a “history of experimental
embryology from about 1920 up to 1950,” and this rise-and-fall narrative would
feature as its central theme. Waddington reacted defensively, writing first to
Needham for advice and support, then to Goodfield Toulmin in April. 1 He
reacted especially to the chauvinism implicit in her approach:

There never was any downfall of the concept of the organiser. . . . The only thing that ever fell
down was a straw man erected by some American biologists for the express purpose of pushing it
over. They performed this simple task so much to their own satisfaction that they had some success
in persuading American biologists in general that the problem of embryonic induction is not
important; and this was a major contribution to the extremely unfortunate divorce that has grown up
 between genetics and embryology in the States.

Rather than simply being a stylistic division between genetics in Europe
versus the United States, Waddington viewed this division between genetics and
embryology as a fundamental blind spot in the whole of postwar molecular
biology:

It is because the essential connection between these two subjects (which I was one of the first to
formulate in modern terms in my “Organisers and Genes” [sic ], 1940, although Morgan had of
course done so at an earlier stage) has been obscured for the last quarter of a century, that molecular
biologists are now saying that they will have to move in and solve the problems of differentiation
from scratch, as though nothing of value had yet been discovered. And of course when they do turn
their attention to the subject, they light immediately on the relations between one of their own main
topics, enzyme induction in bacteria, and the phenomena of embryonic induction, i.e., “the
organiser.”

This notion that development and genetics might have something in common
was, of course, the impetus behind Waddington’s decision to travel to Morgan’s
Caltech lab in 1938. In Organisers and Genes , Waddington postulated that the
very earliest stages in development, e.g., the differentiation of the mesoderm into
neural tissue, proceeded by a kind of preset reaction to a trigger. The patterned
reaction was part of the genetic apparatus of the cell, and Waddington connected
this system to his “masked evocator”; the trigger—the organiser—might not
have conveyed much information, but it was not unimportant or without
function.
Indeed, Waddington saw a direct correlation between the “organiser (or
evocator)–masked evocator–genes–proteins” model he formulated just before
and after World War II and the more recent molecular biological terminology of
“inducer–repressor substance–structural gene–induced enzyme.” This similarity
alone was enough to render a “fall of the organiser” narrative false: “If you start
dealing with anything as alive as the problem of gene activation in higher
organisms—which is what the organiser problem amounts to—you can’t
possibly get deep enough into it not to appear foolish unless you are active in it
yourself.” Goodfield Toulmin’s approach, implied Waddington, was simply too
infected with the hubris of American molecular biology to properly assess the
history and significance of his organizer work with the Needhams.
Joseph Needham’s response was likewise critical, though much less caustic
than Waddington’s. 2 “[You] will undoubtedly fail,” he wrote to Goodfield
Toulmin, “unless you can achieve a more open mind than is to be inferred from
the phrase [‘downfall’].” He saw the “drama” in quite a different light: “the
perennial struggle of conceptual opposites; here stimulus vs. reactivity, as
perennial as the continuity vs. discontinuity antithesis in physics.” Needham also
guessed—correctly as it turned out—that Goodfield Toulmin formulated her
rise-and-fall narrative not through conversations with other historians of science
but via Peter Medawar, who was an acquaintance of Goodfield Toulmin’s
husband, philosopher of science Stephen Toulmin. 3 Needham recognized the
shadow of Medawar’s influence because of their conflict over the status of the
Cambridge organizer project more than a decade earlier.
Back then, the conflict between Medawar and Needham circulated around
Medawar’s new book, Scientific Thought in the Twentieth Century: An
Authoritative Account of Fifty Years (1951). In it Medawar made only oblique
and unflattering remarks about the work of Needham and others—all former
supporters of Medawar when he was at Oxford. Needham, who rarely responded
to these sorts of slights after he had transitioned to Science and Civilisation in
China , immediately dashed off his complaint to Medawar:

I feel hurt by the treatment which you have given to the work of Waddington, Holtfreter, and
myself in your essay.
. . . [T]he fact that you refer to us only obliquely and indirectly, with what seems a studied
suppression of our names, I find very distressing. That you make no mention of either Chemical
Embryology or Biochemistry and Morphogenesis (works on no mean scale, and not without
originality) will in the end be regarded, I believe, as a reproach to you and not to me.
. . . I have not changed my opinions as to the value of the organiser concept, which I believe
further advances in biochemistry applied to embryology will only strengthen and elucidate. . . .
What I criticise in your article, however, is not so much a superficial approach to the problems of
morphogenesis, as the failure to accord any recognition to the whole subject of chemical
embryology, which includes much more than the study of organiser phenomena . . . originating
during the period which [Scientific Thought in the Twentieth Century ] purports to cover.
. . . The way in which you have treated us all seems to me to indicate an unfriendly spirit which
I had no idea that you entertained. 4

Medawar responded that he offered the essay as “a dispassionate statement of a
widespread and purely objective opinion.” 5 But Needham remained
unconvinced. Medawar seemed to have made an intentional about-face from his
youth when he needed the support of Needham and Woodger, who were only too
happy to give it:

I am still rather curious to know who “read into the work of the Cambridge school” [quoting
Medawar’s 3 May letter] all the significance which in your opinion, ought not to be read into it
now, and what exactly that significance was. If you are thinking of the chemical nature of the
primary organiser, we had our beliefs at the time, and other workers had theirs; but I would like to
remind you that it was Waddington and I who gave the first evidence of what is now called
“indirect induction,” with dyes; and also that I was scrupulously careful to explain the difficulties
caused by these phenomena in Biochemistry and Morphogenesis . 6

 By the time Goodfield Toulmin wrote to Needham and Waddington in the
mid-’60s, Medawar was a Nobel laureate and the director of the National
Institute for Medical Research. Though correspondence with Woodger had
largely ceased, he retained some interest in the history and philosophy of
biology. In fact, he entitled his 2 September 1965 address delivered at
Cambridge to the Zoology Section of the British Association for the
Advancement of Science “A Biological Retrospect.” When he published the talk
in Nature a few weeks later, he joked that the talk “has not been well received.” 7
But he meant only that the notion of looking backwards for the philosophical
underpinnings of molecular biology at all was such a relic of biology’s
disciplinary past that he feared few wanted to hear him; his assignment at the
BAAS was to speak of the future of biology, as Jan Smuts had done at the BAAS
four decades earlier.
Despite his bit of mild self-deprecation, Medawar’s address proved to be
anything but controversial. Completely disregarding everything he had once
believed, Medawar framed twentieth-century history and philosophy of biology
as a series of victories for mechanism over vitalism. Not surprisingly, at the
vanguard of this successful charge against the bugbear of vitalism stood his own
biology aided by new molecular approaches. Even the practitioners of the
important field of ecology had gradually learned to regard molecular biology as
the paragon of modern life science: “[Anyone] working to understand the
agencies that govern the structure of natural populations in space and time needs
much more than a knowledge of natural history and a map. He must have a good
understanding of population genetics and population dynamics generally. . . .
[A]n unreasonable feeling that he ‘ought’ to know something about it is more
likely to be found in a good ecologist.” 8 Medawar insisted on the importance of
molecular mechanics for those working at the level of systemic environmental-
organismal relationships because these interactions were, at bottom, based on the
playing out of genetic code scripts. “Genes are messages,” Medawar explained.
This notion of bottom-up genetic information had reorganized the older
“‘dynastic’ conception of evolution” based on “pedigrees or family trees.”
Biology was no longer properly a “vertical” system wherein one specialist would
study, say, deciduous trees as a botanist. Instead, “horizontal” studies at the
population, cell, and molecular level uncovered the gratifying irrelevance of all
that once counted as biology, now disparaged as mere classification and natural
history—that is, stamp collecting. Once sloughing off the significance of the
organism, evolution could be understood as change in the “genetical structure of
a population” rather than alterations to morphology, adaptations to new niches,
and so forth. 9
Horizontal approaches, especially at the molecular level, “revolutionized”
and “freed” biology from its earlier flirtation with philosophy. As an example of
this earlier philosophy-laced biology, Medawar targeted the work of theorists in
the prewar era like Sir Frederick Gowland Hopkins, Needham’s old mentor in
biochemistry. Hopkins held the now heretical view that the cell was a highly
significant center of irreducible colloidal organization rather than a mere holding
tank for the true puppeteers: the genetic elements in the nucleus. Furthermore,
Medawar caricatured the experimental embryology of the 1930s following the
biochemical work of Hopkins—by which he strongly implied the organizer work
of Waddington, Brachet, and the Needhams, though he only named Hans
Spemann—as a hunt for the fabled “philosopher’s stone.” 10 Thankfully, at least
according to Medawar, molecular biology had progressed away from searching
for the actual instigator of developmental change. Development under his
molecular scheme was preformationist: “an unfolding of pre-existing
capabilities, an acting-out of genetically encoded instructions; the inductive
stimulus is the agent that selects or activates one set of instructions rather than
another.”
To be sure, his “Retrospect” scrupulously avoided any mention of the
scientific work of TBC members. But upon reading his address published in
Nature a few weeks later, Waddington and Needham immediately recognized
that their work was being forced to play the role of historical foil to modern
scientific progress. Each commented on Medawar’s article, scribbling in the
margins of their personal copies of Nature. Their marginal remarks underscore
the extent to which they felt their work was being misjudged, avoided—and,
where found to be useful, misappropriated as the work of others. 11 Waddington
lamented years earlier that geneticists “never had much of a clue what
determination and induction were all about and, in fact, still haven’t”; Goodfield
Toulmin’s historical research only confirmed this fact. 12 But Medawar was an
Oxford biologist and a personal acquaintance. It seemed inexcusable to
Needham and Waddington that he would denigrate or ignore the importance of
their science and philosophy.
Goodfield (no longer Goodfield Toulmin) incorporated the organizer work in
her essay, delivered to the Boston Colloquium for the Philosophy of Science in
1968 and published in the volume of proceedings shortly thereafter. 13 Her tone
had changed somewhat. Spemann’s organizer—she did not mention Waddington
and the Needhams’ collaboration, but Huxley and De Beer’s—was one of those
“theoretical entities” that scientists employ from time to time. Like “gene,” the
organizer played an important heuristic role even when there was little evidence
for its existence as a thing. It lost its appeal when the organizer became reified as
a substance that had to be identified with other already known biochemicals. Of
course Goodfield’s article only explored the understanding of an organizer as a
particle or group of particles. That did fail—there is no entity. But at least for
Waddington and the Needhams, it was the state of order inside the system of the
fertilized egg that led to organization, not the chemical itself. Waddington would
critique the gene concept for the same reason.
Three years after this episode, in 1968, the National Science Foundation
(United States) asked Waddington to evaluate Goodfield’s application to renew
the grant that funded her “rise and fall of the organizer” history. Waddington
explained in detail what he took to be the precise missteps by American
biologists in the 1930s and ’40s that, combined with the myopia of molecular
biologists, had condensed all contributions of embryology from before the war
into the “decline and fall of the organiser” narrative. 14 Waddington expressed
reticence to address the issue at all, regarding it as more appropriate “for a
novelist concerned in directions of human personalities and human ambitions”
than for a historian and philosopher of science. But, given that Goodfield was
committed to a “balanced view,” and observing that she had appeared to take his
earlier criticisms to heart, he supported the renewal of her grant.
He didn’t stop his commentary there, however. Waddington used Goodfield’s
recommendation as a platform to deliver his own history of the organizer
project’s reception by life scientists—the reason why her historical account
would necessarily be incomplete. He pointed in two directions at once:
embryologists like Ross Harrison and Paul Weiss, who Waddington believed
were afraid to really engage with genetics, and the “simpleminded viewpoint” of
American molecular biologists who sidestepped the real complexities of the
developing organism. The European successors to Spemann, including
Waddington and Needham, recognized that the problem of cell/tissue
“determination” provided a fundamental conundrum that molecular biology had
not even begun to address. Molecular biologists were mechanists focused on the
behavior of particles. Determination was a field concept—a continuum rather
than a particle. Moreover, determination only masked the real issue: “the origin
of ‘competence,’ which decides which groups of developmental tendencies
become available for determination.” For Waddington, modern molecular
biologists and their disciples had taken their own fascination with the molecule
of life idea too seriously. Now the whole field was blind to the basic problems of
development, form, and structure that the organizer biologists were beginning to
address in the 1930s.


Case in point for Waddington was Francis Crick. As one of the two main
figureheads of the new DNA revolution in biology, Crick billed himself as able
to pontificate confidently on the relevance of molecular approaches to age-old
problems like the existential struggles of human beings. Around the same time
Medawar gave his address and Goodfield approached Needham and
Waddington, Crick gave a series of three important lectures at the University of
Washington. Though he preferred the title “Is Vitalism Dead?,” he agreed to
name the lecture series “Of Molecules and Men” for his John Steinbeck–attuned
American readers. Waddington reviewed the book for Nature , and though he
was not critical of Crick’s arguments against vitalism, he accused Crick of
choosing only the simplest cases—perhaps not attacking straw men, but fairly
close to it. 15
Crick wrote directly to Waddington, confessing that, as he was forced to
publish quickly, his work might have been a bit slapdash. In the face of
Waddington’s critique, Crick re-construed his classification of vitalism into three
camps: (1) substance dualists, (2) those who believe in “so-called biotonic laws,”
and (3) those convinced, like Max Delbrück, that “radically new laws of physics
or chemistry are likely to be discovered from studying biology.” Crick did not
mention which camp he believed Waddington occupied. 16 In his reply, however,
Waddington chose the third camp, with strong leanings toward the second
variety:

There is a lot of biology which is at present as far from basic physics as the gas laws are from the
dynamics of the individual gas molecules. As you say, the field of natural selection is one example
and I should myself suggest that the morphogenesis of large scale structures such as bones will
quite likely turn out in the same category. This means, I think, that new bodies of theory will have
to be developed to deal with such phenomena, but this does not imply in any way that the new
theories cannot be finally incorporated into an expanded body of physics. 17

Waddington accused Crick of twin errors: oversimplifying the positions of his
opponents, and placing too much stock in his own “conventional” approach. As
physics became more sophisticated, assured Waddington, its most mature
thinkers realized that what they took for granted as finished, objective
knowledge often turned out to be anything but. An expansion of physical law
might be needed to explain the esoteric folding patterns of proteins, for instance.
Waddington clearly regarded the philosophy behind Crick’s molecular approach
as far too unsophisticated; certainly, it did not warrant the preformationist and
mechanist “genes-for” language that Crick deployed confidently in “Of
Molecules and Men.”
A few years later, Crick wrote again, disputing Waddington’s entire notion of
an epigenetic landscape. Unlike their correspondence over “Of Molecules and
Men,” this 1974 exchange turned sour. Perhaps that sourness was due to
Waddington: In the intervening time, he had called into question Watson and
Crick’s claims to originality for the double-helix discovery. He had unearthed
memoranda and correspondence of Lord Halsbury dating to 1951 and ’52
describing the “mould-casting-mould” process of protein translation. Halsbury
had even predicted the “geometry and thermodynamics” of the double-helix
discovery, including the “A and B chain” association and “zipper-wise” process
of DNA replication. 18 Crick had not responded when Waddington made these
discoveries. Still, with this set of events in the background, Crick opened the
1974 correspondence with a notably negative tone—inquiring what in the world
the epigenetic landscape was supposed to mean, with the implication that
Waddington’s most well-known model was “rather vague”—that it was difficult
to render “the analogy precise.” 19
Expressing surprise that Crick could find such a “very simple and perfectly
clear idea” confusing, Waddington supplied a detailed three-page exposition. In
it he emphasized the epigenetic landscape’s perspicuity when describing (“not
explain[ing],” Waddington underscored) evident phenotypic stability:

This is perhaps the major implication of the landscape description, since it states that the very
numerous genes will interact in such a way as to produce only a smallish number of more or less
clearly distinguished cell-types. Just how many, or how clearly distinguishable, and what
intermediates can persist, are facts to be discovered empirically. But in most biological systems, it
seems that it is justifiable to think of only a limited number of combinations being produced out of
the astronomical number which are potentially possible. 20

To this defense of the epigenetic landscape, Crick replied that he did not
believe the ball as “cell, tissue or pattern” helped to make the overall concept
useful. “What I had expected you to say was that the path of the ball represents
the lineage of a single cell of the adult animal,” he explained. Crick then pictured
the single egg cell dividing, as typical, and each individual daughter cell
traveling down a separate path toward its eventual functional end as liver,
kidney, eye, hand, and so on. “Otherwise,” he surmised, “the analogy seems to
me to be vacuous.” 21 If the epigenetic landscape could account for any level of
analysis from cell to tissue, Crick asserted, the model would be unfalsifiable and,
therefore, unscientific—a damning critique. 22
Though still cordial, Waddington began his rejoinder by attacking Crick’s
quasi-Popperian appeal to falsifiability: “Now you can easily postulate a
developing system which involves no stabilities etc., but in which every effect
was a linear function of its causes; no dominance or epistasy [sic ] of genes, no
distinctness between issues, a genotype with 100 loci with two alleles at each
locus . . . would have 2100 phenotypes, and so on.” 23 Fisher, Haldane, and the
other population geneticists employed just such a description of an organism,
asserted Waddington. And “orthodox evolutionary” biology relied upon a
Newtonian “billiard table landscape” that worked when applied to agriculture or
laboratories with fixed environments—and only then with “fudge factor[s]” like
“epistatic variance” occasionally thrown in—but could not account for
“evolution in unpredictably variable nature.” 24
“What I object to about the ball representing a group of cells,” revealed Crick
in the next letter, “is that, in brief, a fertilised egg (where it all should start) is
only one cell.” Crick accused Waddington of accommodating the epigenetic
landscape to his “egg only” explanation solely through “fast talking and waving
your hands,” but this hardly counted as accommodation. As it stood, the
epigenetic landscape was a “useful idea in the Thirties [sic ] but I think it has
long outlived its usefulness.” Molecular biology had long since demonstrated
that elements of Waddington’s model, “switches” in development (represented
by hills in the epigenetic landscape model), for instance, could be reduced to
mutations and translation errors in DNA. And with that, Crick cajoled
Waddington, “Throw it away and start again!” 25
“[F]or purely formal reasons,” Waddington retorted a few weeks later, “I
should not leave you talking such nonsense without putting some reply on
record; and perhaps the fact that your letter contains . . . no argument at all, but is
mere abuse of the idea, suggests that you may not be so happy at a total rejection
of it as you make out.” After three months of back-and-forth objections and
counter-objections, Waddington had grown impatient. Crick seemed determined
to remain ignorant of the complexities of development, and that irritated the
“third way” epigeneticist. What’s more, Crick had dismissed the philosophy and
science that had emerged from it as ideas that had outlived their usefulness. But
if Waddington’s concepts belonged to the 1930s, as Crick asserted, “You are
going back to something which was already discarded as not useful in the
Twenties,” Waddington insisted. He knew that Crick’s Medical Research
Council Laboratory of Molecular Biology in the Cambridge Medical School had
been conducting cell lineage studies. Crick thought this work acted as the very
bridge between genetics and embryology that Waddington had always longed for
—and better than what Waddington had accomplished over the last four decades.
Crick finally let on that he thought the cell lineage studies were more suitable
than the epigenetic landscape. Certainly, Waddington admitted, cell lineage
studies could “be very precise and rigorous.” However, they rarely addressed the
real problems of development: How are changes in biochemistry and organismal
pattern caused? He confronted Crick, somewhat bitterly, in closing:

In my sceptical opinion, the whole of this line of thought has arisen because the Drosophila
genetic-engineers have invented a wonderful way to label single cells and trace its clonal
descendants, and are desperately—and not very successfully—looking for some questions that
technique can answer. It’s your choice to follow that lead. I’d just remind you—sorry to bring up
this beastly past again—that the last chap who went in for cell lineage in a big way was Woodger.
He mostly used logical algebra rather than geometry, and he finished up by defining a gene as

mend = Df Aeq’xy (x,y) E whz.(Apr’Zyg’x[left-restrict]K[right-restrict] Apr’Zyg’y E 1->1)

which I consider a fate worse than death. 26

Crick declared a cease-fire a week later. 27
The Waddington-Crick exchange over the meaning and utility of the
epigenetic landscape and all that was connected to it reveals the extent to which
Crick and Waddington held two distinct and, to an extent, competing views of
biology. Crick viewed life through the operation of unfolding sequences of pre-
formed genetic instructions. Waddington saw interacting genetic and epigenetic
systems responding de novo to a plethora of inputs. While genes and their
products played a fundamental role in Waddington’s epigenetics, the structural
“buffering” of this system through feedbacks against disruption, genomically
(internal) and environmentally (external), could not be discounted. To Crick and
to Medawar, for instance, this concern with levels and patterns, cell collectives
and whole organisms instead of single eggs and their DNA endowments, could
only be “vacuous,” a bit of metaphysics smuggled into science.
Maybe Medawar and Crick were correct. They were Nobel laureates, after
all.
 14

THE 1960s REINCARNATION OF THE DEBATE

For each of us, as for the robin in Michigan . . . this is a problem of ecology, of interrelationships,
of interdependence. We poison the caddis flies in the stream and the salmon runs dwindle and die. .
. . We spray our elms and following springs are silent of robin song, not because we sprayed the
robins directly but because the poison traveled, step by step, through the now familiar elm leaf-
earthworm-robin cycle. These are matters of record, observable, part of the visible world around us.
They reflect the web of life–or death.
Rachel Carson, Silent Spring

I n the early 1960s, Rachel Carson’s Silent Spring warned that our nonchalant
disregard for organic interconnectedness and multicausality was resulting in
dead birds and worse. It is well known that Carson’s book kick-started an
environmental movement and inspired a suite of legislative responses from the
highest levels of the American government. But Carson’s was only one of a
number of critiques made by scientists in the 1950s and ’60s of the dominant, if
implicit, philosophical perspective that pervaded the scientific-industrial
complex. Recent discoveries regarding DNA had granted molecular biologists
the kind of cultural cachet formerly reserved for physicists and chemists. But
even biology’s insiders began to express worries about the direction of scientific
progress and its social, economic, public health, and even ethical costs.
In many ways, critics merely repeated concerns from much earlier in the
century. Like Bergson, Driesch, and their followers, critics in the 1960s feared
the dehumanizing overextension of mechanism, which they saw mirrored in the
gene-centric, reductive explanations for life on offer by many prominent
biologists, Medawar and Crick included. Just as in the years following the First
World War, mechanistic science a half century later delivered many
technological and medical benefits, and promised even more. Yet it also seemed
to be endangering the natural world and, ultimately, human flourishing. Among
those who rejected the new gene-centric mechanism, some called for something
quite like vitalism. Thus, though many of the terms had changed, the
conversations around theoretical biology in the middle of the century felt very
much like they had in its first quarter. Waddington and fellow travelers must
have experienced a kind of déjà vu as they again tried to articulate an organic
alternative.
Importantly, contrary to the suggestions of some historians of science,
organicism did not disappear or go underground in the middle of the twentieth
century, decades after its 1930s peak. Though certainly not mainstream, “third
way” theorists continued to announce their alternative and gain new followers in
the midst of a new debate between mechanism and a quasi-vitalistic rejection of
mechanism. 1

In his 1961 critique of Isaac Asimov’s Intelligent Man’s Guide , biologist Barry
Commoner (1917–2012) decried the epistemological problems that would result
if biologists continued down their current path. Biology would divide into castes:
a molecular-biology caste flush with cash, postdocs, and gizmos, and an
organismal caste that would be seen as atavistic and consequently driven to
extinction. And this was only the beginning of his critique of molecular biology.
Though a friend of renowned protein biochemist Linus Pauling, Commoner
remained skeptical of Asimov’s triumphant claim that a biology focused on
molecules erased any real distinction between the living and nonliving: “Since
biology is the science of life, any successful obliteration of the distinction
between living things and other forms of matter ends forever the usefulness of
biology as a separate science. If the foregoing [statement of Asimov’s] is even
remotely correct, biology is not only under attack; it has been annihilated.” 2 In
fact, it did appear to Commoner that the discipline of biology as it functioned
before the 1960s—the biology that appreciated organisms, in other words—was
sliding gradually toward annihilation. Instead of taxonomists or embryologists,
or even geneticists, biologically curious and experimentally competent students
migrated toward more lucrative biochemistry and biophysics. Instead of plants
or even Drosophila , more and more biologists used bacteria and viruses as their
model organisms. The very meaning of “organism” had changed. The effects of
this seemingly academic change resulted in an obvious, appreciable social shift,
since those trained in physics and chemistry exhibited an “increasing tendency to
ignore the facts of life”—more specifically, the facts of evolution and
development. 3
Without a doubt, modern science had created and would continue to create
wonderful things: antibiotics, pesticides, electricity from uranium, transportation
from oil, and so on. But Commoner feared that the public regarded technological
know-how as a tangible indicator that our knowledge of the natural world was
far more comprehensive than it actually was. Scientists, too, had been blinded by
their confidence in the magic of technoscience. Thus our useful technoscientific
devices cut two ways: “Unwittingly we have loaded the air with chemicals that
damage the lungs, and the water with substances that interfere with the
functioning of the blood. Because we wanted to build nuclear bombs and kill
mosquitoes, we have burdened our bodies with strontium-90 and DDT, with
consequences that no one can now predict.” 4 Even the seemingly unmitigated
good of antibiotics had a dark side. Given our underappreciation of evolution,
we had created mutants, microbes once able to be controlled through traditional
means, now virulent, resistant to our antibiotics. The problem wasn’t merely that
chemists and physicists threw these harmful products into the air, the water, and
our bodies without knowing what they were doing; that was merely the
symptom. The larger problem was philosophical, rooted in that old mechanistic
hope, reduced now to the level of biochemical and, eventually, molecular
engineering. “The dominance of the molecular approach in biological research,”
warned Commoner, “fosters increasing inattention to the natural complexity of
living systems.” 5 Not only were the immediate symptoms problematic—the
radiation, antibiotic-resistant bacteria, pollution, and so on—the whole reductive,
mechanistic worldview would lead to oversimplified, short-term fixes, which in
turn would lead to greater and greater problems.
Commoner mirrored concerns that others had voiced generations earlier, but
his standing in the scientific community was much different than the advocates
of the 1930s and before. Commoner was a respected plant physiologist, chair of
a committee on molecular biology, and a voice for reform within the American
Association for the Advancement of Science. 6 This level of skepticism
regarding the modern reductionistic worldview had rarely been expressed by an
“insider” before, nor had sentiments like these been highlighted on the pages of
the prestigious journal Science.
Others repeated Commoner’s misgivings about the allmacht of gene-centric
mechanism. Bertalanffy continued to speak out against what he saw as the
geneticists’ tendency for casual synecdoche, which of course privileged their
own subfield. 7 Hans Jonas, a former student of Rudolf Bultmann and Martin
Heidegger, published The Phenomenon of Life (1966), another work that
highlighted the enormous complexity of living things, the organism-centric
structure of the universe, and decried the mechanistic impulse to simplify. 8
Philosopher Marjorie Grene took the gradualistic gene-centrism of the neo-
Darwinian synthesis to task in her groundbreaking “Two Evolutionary
Theories.” 9 Well-known and respected physical chemist Michael Polanyi
insisted that, while a scientist must account for the physical and chemical
contributors for any biological phenomenon, physics and chemistry were not
sufficient explanations. An organism might be fruitfully described as “a
mechanism founded on the laws of physics and chemistry but not determined by
them.” 10 Even DNA, as powerful and important as it is to life, was only part of
the physicochemical “boundary system.” If scientists could recognize organisms
as examples of “dual control” systems, in need of “irreducible higher principles”
of order, claimed Polanyi, then the long wrestling match between vitalism and
mechanism would be over. A third class of “organismic” principles could be
applied to living and perhaps even nonliving things. 11 In the biological
laboratory, a small but persistent group of biologists pursued alternatives to the
notion of an executive gene by investigating the importance of nonnuclear
inheritance. One of the most prominent, Tracy Sonneborn, pronounced, “[A]n
isolated nucleus could not make a cell even if it had all the precursors, tools, and
machinery for making DNA and RNA and the cytoplasmic machinery for
making polypeptides. Self-assembly of genic products can only go so far. . . .
Strong evidence now confirms the old dictum that only a cell can make a cell.” 12

Perhaps more vocally than anyone else, political firebrand-turned-journalist-

turned-historian of science Arthur Koestler (1905–1983) captured the discontent
in the 1960s with what more than a few molecular biologists were touting as the
final triumph of mechanism. 13 Koestler identified this disconcerting mechanistic
“prevailing philosophical bias” among scientists and influential intellectuals of
his acquaintance. 14 In fact, Koestler would lean toward a renewed vitalism,
opening another window into the again-contested fate of the philosophy of
organism in the third quarter of the twentieth century.
An agnostic Hungarian Jew who, like Needham and Bernal, vocally
supported Marxism while decrying Stalin’s brutal and autocratic version of it,
Koestler believed that science as practiced in the West took concepts of
individuality and cutthroat competition from business and read them into the
natural world. This kind of worldview went beyond what any empirical evidence
warranted. 15 Instead, Koestler was convinced that something closer to neo-
Lamarckism acting on an integrated organic system must be invoked to explain
the diversity of living things. But he was neither a vitalist nor a fuzzy-thinking
romantic, sentimental for the days before images of “nature red in tooth and
claw” forced themselves upon the scientific mind. If anything, Koestler felt that
the hearty Darwinian worldview of the late nineteenth century had fallen victim
to a kind of lazy mechanistic dogmatism in the twentieth. 16 Organisms, and
especially humans, simply responded too quickly and too effectively to
perturbations by the environment (including other organisms) for the “random
walk” of evolution postulated by population geneticists and their neo-Darwinian
compatriots to give the complete picture. 17 It was as if the advocates of neo-
Darwinism wished to retain their cake while eating it: No purpose could be
attributed to anything in the universe—even the bearers of life itself—yet
everything was adapted and served some function. That sentiment spread well
beyond biology:

As a result, man’s destiny was no longer determined from “above” by a super-human wisdom and
will, but from “below” by the sub-human agencies of glands, genes, atoms, or waves of probability.
This shift of the locus of destiny was decisive. So long as destiny had operated from a level of the
hierarchy higher than man’s own, it had not only shaped his fate, but also guided his conscience and
imbued his world with meaning and value. The new masters of destiny were placed lower in the
scale than the being they controlled; they could determine his fate, but could provide him with no
moral guidance, no values and meaning. A puppet of the Gods is a tragic figure; a puppet
suspended on his chromosomes is merely grotesque. 18

As Koestler saw it, the preoccupation with excavating the purposeless,
meaningless bedrock of chemicals and genes and then projecting these entities as
bottom-up causes—that is, as the only truly significant actors in the world—was
the only acceptable theory, meta-theory, or scientific worldview on offer by
midcentury.
Beginning in the 1950s, Koestler looked for trends in the sociology and
history of science that might make sense of the apparent blind devotion of
contemporary scientists and philosophers to mechanism. When studying the
history of the sciences, he found that biology had once followed the broader
historical trend in most of the Western sciences: Sciences had attempted to
deliver meaning as well as explanation. Even if one took a more skeptical stance
—for instance, that the sciences lacked the drive for conveying the meaning of
the world—at least then the sciences would refrain from despoiling the search
for metaphysical meaning that other meaning-making institutions found in the
universe. 19 By the nineteenth century, however, this had all changed. Meaning
was no longer central to science. If anything, natural sciences “un-meaninged”
the world. There were a variety of reasons for this change in approach on the
part of physicists and chemists. But in biology, Koestler singled out the early-
twentieth-century Mendelians as the group who had emptied biology of any
contribution to a meaningful universe. This move was problematic, according to
Koestler, because as the study of life itself, biology seemed to be closer to a
meaning-making science in its very essence than some other sciences. Though
biologists purported to prescind from making claims about meaning, they
paradoxically made an implicit claim that there was meaning; the meaning was,
There is no meaning. He especially fixated on the individual he took to be the
era’s chief Mendelian, William Bateson.
Using Bateson as his surrogate for a larger attack on meaning-destroying
biology, Koestler resurrected a piece of nearly forgotten lore from the early
1920s. Koestler titled his book with a telltale wink to Hardy Boys fans: The Case
of the Midwife Toad. Just after the First World War, a persuasive Viennese
herpetologist, Paul Kammerer, toured Europe advocating the inheritance of
acquired characteristics. Kammerer eventually came to Cambridge—to speak at
the Biological Tea Club, attended by Gregory Bateson, Joseph Needham, and
others. According to Koestler, Gregory’s father, William Bateson, unjustly
accused Kammerer of fraudulently passing off his evidence supporting
Lamarck’s theory. William Bateson’s accusation of fraud and Kammerer’s
subsequent—though Koestler claimed unrelated—suicide seemed to close the
case in 1926. The story, in other words, more or less died with its main
protagonists. Neo-Lamarckism, too, was thought to be dead. But Koestler
remained unconvinced. During a lengthy investigative process, he came to
believe that William Bateson’s dogmatic support for the worldview supported by
Mendelism and Darwinism, rather than actual empirical proof, was what had
decided the case against Kammerer and the direct influence of the environment.
With the success of William Bateson’s hard, particulate inheritance, evolution
would inevitably lend scientific credibility to a much harsher view of humans
and the rest of the biological world. For this audience, Koestler skillfully, if
anachronistically, recast the conflict between the elder Bateson and Kammerer as
a gripping, tragic, metaphysically motivated struggle between vicious
“Mendelian-Darwinism” and virtuous neo-Lamarckism. 20
Not only was Koestler an ambitious polymath and an avid reader, he had a
sharp eye for people’s inner inclinations. He had long sensed discontent on the
part both of scientists and the informed public, who suspected the picture of
evolution they were being sold had major flaws in its philosophical foundations.
Moreover, his reading of Whitehead, whose works he frequently cited,
convinced Koestler that the multiple dustups between mechanists and vitalists
through the nineteenth and twentieth centuries had already produced a feasible
alternative, a more philosophically robust synthesis than that offered by the neo-
Darwinians. Though he had few, if any, formal scientific credentials, he believed
he could harness this stream of latent frustration and turn it into an actual
movement to address the sicknesses of isolating modernism, in both their
capitalist and their communist versions. Over the course of the 1960s, he
successfully convinced three publishers (Macmillan in New York, Hutchinson in
London, and Molden in Vienna) to fund an expeditionary event: an international
conference of scientists who would convene in a massive chateau in the Swiss
Alps to attack gene-centric mechanism. Koestler asked the publishers to pay the
scientists half of a roughly $13,000 honorarium in advance (the Ford Foundation
also offered a $1,000 grant). In return for their investment, he promised they
would profit from the book of prepared papers and conference proceedings, with
comments, questions, disputes, and so forth transcribed directly from state-of-
the-art tape recordings of the entire multiday event. 21 Koestler obviously felt
there was enough interest inside and outside the academy to undertake such an
ambitious venture.
This academic conference, Koestler’s 1968 Alpbach Symposium, and the
resulting volume, Beyond Reductionism , is a milestone of this upsurge in anti-
mechanism. The attendees hailed from a wide variety of disciplines:
anthropology, sociology, and psychology, as well as biology. A new appreciation
for Carson-inspired ecological thinking colored many of the presentations and
discussions. But many of the themes were familiar to those who were familiar
with “third way” thinking. So were many of the participants. Ludwig von
Bertalanffy, predictably, called for more effort to be put into understanding
organic levels, from gene to individual to population, in terms of interlocking
systems. Respected University of Chicago and Rockefeller University
embryologist Paul Weiss identified an age-old “analytical-reductionist
orthodoxy”—the circle of those who resisted the notion that wholeness had
anything to offer the “role of pars pro toto ” molecular biology had assigned
itself. 22
Koestler invited Waddington, too. Waddington missed the first day of the
Alpbach Symposium, the day that included the Weiss and Bertalanffy papers,
because he was receiving his doctor honoris causa from his friend Mischa
Fischberg’s University of Geneva. 23 When he arrived, he adopted a somewhat
skeptical mien; he had, of course, heard it all before—three decades earlier in the
TBC. And there were other reasons for him to be skeptical, reasons closer to
home.
During the retreat, Waddington argued extensively with Koestler. The host
had co-opted a number of Waddington’s terms and put them to uses he did not
approve. “Genetic assimilation” and “chreod,” Waddington insisted, did not
imply somehow that Lamarck offered a kinder, gentler depiction of evolution, as
Koestler purported. Nor did Waddington buy Koestler’s notion that all modern
genetics was a conspiracy cooked up by William Bateson and the early
Mendelians (who continued to be Koestler’s favorite targets well after Case of
the Midwife Toad ). Waddington certainly concurred with the other Alpbach
participants that something was rotten in the state of molecular biology. Yet
genetics was not its source. Genetics did have something to offer in the larger
context of development and evolution; it just could not grant a sufficient
explanation, as gene-centric mechanists like Francis Crick seemed to defend. In
other words, Waddington resisted the firebrand talk of Koestler and some others
at the Alpbach Symposium not because he disagreed with their diagnosis of the
problem, per se, but because he believed their prescriptions for revolution
disregarded too much good science.
And there was something more: Waddington regarded the anti-mechanism
espoused at Alpbach as occasionally hewing too close to vitalism. The “third
way,” for which he continually argued, had converted many of the biological
rank and file. The foundations did not need to be torn up and replaced by
something closer to vitalism, thought Waddington; one must stay the course in
articulating the alternative. The lessons of epigenetics—a true alternative to
mechanism and vitalism—merely needed to become more widely adopted in the
life and social sciences.
A brief episode after the conference involving Waddington, Bertalanffy, and
Koestler reveals more about his suspicions regarding their positions. Waddington
mailed Koestler a multipage critique entitled “Comment on Bertallanfy [sic ],”
an attempt to disentangle his organicist ideas from the direction he feared
Bertalanffy was taking them with his general systems theory (GST). 24
Waddington had missed commenting on the paper in person because of his
honorary doctorate ceremony at Geneva. 25 Waddington argued in his
“Comment” that Bertalanffy’s GST did not exclude certain theoretical work that
he felt was clearly derived from the same reductionist metaphysic undergirding
molecular biology—though now it was clothed in the new “digital information”
style that would eventually become the lingua franca of the computing and
artificial intelligence realms. Waddington clearly parsed “reductionism”
according to the ways he had come to think about it during the TBC years. In the
UK, reductionism means “the attempt to find some simpler elements in terms of
which the complex can be understood.” Waddington took this to be
unproblematic, almost common sense; one might even call it “analysis.”
Bertalanffy’s critique of this perspective was misguided and made GST seem
just another piece of semantic fluff. The enemy, claimed Waddington, was
American reductionism, which

implies that we start with a full knowledge of the simple elementary units, and, on the basis of this
knowledge, can determine what the character of the apparently complex really must be. The critical
point in the [organic] paradigm is that it starts at the other end, as it were; from the basis that we
know very little about such elementary units as atoms, molecules, or genes, and what little we do
know has been learnt from the study of the complexes [e.g., organs, tissues, cells, ecosystems, etc.]
into which they enter. We must therefore be prepared to accept additions to our knowledge about
them; but equally it is good tactics to refuse to accept any alleged new insight into the properties of
our basic units until the evidence really forces us to do so. It is this point of view, which began to be
adopted by leading biologists in the thirties (Needham, Woodger, Weiss), that is the real mark of the
new paradigm. 26

Before Waddington sent this essay off to Bertalanffy, however, he sent a copy to
Koestler. Waddington also implicitly attacked Koestler’s position, but Koestler
offered a gracious response to Waddington. Because Bertalanffy was in such a
precarious academic position in Canada, and because most of GST had been co-
opted by computer scientists uninterested in biology or philosophy anyway,
Koestler recommended Waddington politely bow out of the conversation without
sending or publishing the “Comment.” Waddington complied; Koestler never
alerted Bertalanffy to the criticism. 27

Waddington’s reluctance to fully embrace Koestler’s version of anti-mechanism

at Alpbach was due to several factors. Certainly, it had something to do with
Koestler’s presentation of his views and with the unhelpful fungibility of
Bertalanffy’s GST. It also may have been due to the differences between
Alpbach and the other, more extensive series of symposia with which
Waddington had been intimately involved since 1966. The International Union
of Biological Sciences (headed by Waddington at the time, not coincidentally),
with the financial support of the Rockefeller Foundation, sponsored a series of
four annual symposia on theoretical biology. These meetings brought together
two dozen or so biologists, mathematicians, chemists, computer engineers,
physicists, and neurologists—largely through Waddington’s single-handed
efforts. They were held at Villa Serbelloni, the stunning fifteenth-century estate
overlooking the town of Bellagio, Italy, on the southern shore of Lake Como,
near the Swiss border. It was such a perfect setting for the meetings that, once
the Rockefeller Foundation withdrew its support for the building’s use, the IUBS
theoretical biology club ceased. 28 The experience was as close to Needham and
Woodger’s original Theoretical Biology Club as Waddington had experienced in
the years since the war. As good as Koestler’s Alpbach conference was, the four
IUBS symposia were better.
For starters, Waddington found the IUBS symposia more focused. Koestler’s
agenda had followed in the wake of those general fears regarding the extension
of reductive mechanism inspired by Carson, among others. Waddington did
share those fears—though he would come to feel them more acutely in the 1970s
—but he found Koestler’s “holon” concept, introduced in Ghost in the Machine
(1967) and promoted at the Alpbach symposium, both vague and derivative.
Whitehead had rendered a more sophisticated version of this notion almost a half
century earlier. 29 Waddington centered the discussion at the first IUBS
symposium, held from 28 August to 3 September 1966, around a pair of pre-
circulated texts dealing with the finer points of biological theory. The first text
was his own “intentionally provocative” Ballard Matthews Memorial Lectures
given at University College North Wales earlier that year, entitled “The Basic
Ideas of Biology.” The second was a companion piece submitted by co-organizer
Ernst Mayr, “Cause and Effect in Biology.” Waddington hoped these essays
would evoke long-standing questions in biology and that the diverse
backgrounds of the attendees would shed new light on them (rather than merely
heat, as Waddington felt the Alpbach conference tended toward).
Before the meetings convened, Waddington and Mayr had been in lockstep
on that particular organicist balance they wanted to strike. Initially, they hoped to
attract professional philosophers—Ernest Nagel, for one. But both agreed that
Nagel and his followers did not, as Mayr put it, “truly understand the special
problems of biology.” There were likely others, less resistant to organicism;
“[Michael] Scriven” might be one. Mayr tried to convince Waddington that,
since “third way” arguments had already persuaded most working biologists—
molecular biologists and biophysicists excepted—it was the mathematicians,
physicists, chemists, analytic philosophers, and so forth, who remained blinded
by the power of gene-centric mechanism and who needed to hear the truth: “Let
us take the leadership in showing that . . . the existence of highly complex
historically evolved systems, etc. etc. adds such a dimension of complexity to
the simple systems normally studied by physicists and chemists that it is entirely
legitimate to develop an entirely . . . independent philosophy of biology. Let us
show the irrelevance of all the time honored arguments such as mechanism and
vitalism . . . by showing that most of these . . . concepts are meaningless in the
light of our modern understanding of historically programmed systems.” 30 Yet
Mayr cautioned that among the variety of “third way” adopters, there were still
“vitalistically tainted biologists.” He hinted at individuals who would be given a
platform at the Alpbach conference in ’68. If Waddington invited them in the
hopes of expanding the ranks of anti-mechanistic thinkers, Mayr and others
would be forced to “distance” themselves, perhaps to the extent that he and
others would “unwittingly be driven into the reductionist camp.” 31 This
sentiment certainly may have impacted Waddington’s attitude when he arrived at
Koestler’s Alpbach meeting.
Tensions between the perspectives of Waddington and Mayr sharpened during
and after the IUBS symposium, however. In “The Basic Ideas of Biology,”
Waddington criticized a definition of evolution given earlier by Mayr (who,
incidentally, was citing Waddington in that definition) as not going far enough.
Evolution was not merely gene change, even complex gene change in a
population, as Mayr held. Evolution was about the production of phenotypes—a
dance between genes, cells, morphology, the organism, the environment, and
time itself. Mentioning “complexity” in regard to evolution was only the first
step; Waddington wanted organized complexity over time. And where did that
organization come from? No one had a clear answer for that. 32
Waddington took Mayr’s suggestions and sought out like-minded younger
biologists such as Richard Lewontin and Brian Goodwin. Mayr approved of
them. But Waddington also attracted mathematicians attempting to map
extraordinarily complex networks and systems, and the interaction of those
networks with a kind of inanimate environment, such as the French recipient of
the 1958 Fields Medal, René Thom (1923–2002). Thom’s conversations with
Waddington on topology covered similar ground to that Waddington had earlier
shared with Wrinch during the heyday of the TBC. Thom was just then refining
his “catastrophe theory,” a depiction of the process by which discontinuous
phenomena could arise from otherwise smooth, continuous conditions.
Waddington hoped these individuals would help him get greater traction on the
confluence of development, genetics, and evolution—something beyond neo-
Darwinism.
That was too much for Mayr. He would bow out of future meetings, claiming
that Waddington’s concepts hewed too close to “theoretical biology,” whereas
Mayr preferred “the conceptual basis of biology.” He saw as much difference
between these two foci as between “the theory of physics and the philosophy of
physics” and, though he didn’t fault Waddington’s theoretical biology angle,
Mayr preferred something going in the other direction. 33 Later, Mayr would
explain to philosopher of biology Marjorie Grene that participants at the first
IUBS symposium were too interested in “what I like to call biological
engineering.” 34 It wasn’t too close to the gene-centric mechanism he opposed, to
be sure, but it felt far too skeptical about the methods of biology that Mayr held
dear. There was too much math, too much interest in modeling, and too much
criticism of the very neo-Darwinian synthesis that Mayr had helped to establish.
Moreover, with Thom present, the conversation seemed always to veer toward
the discontinuity observed in biological development—in Waddington’s
epigenetic experiments, for example—and away from the gradualism traditional
in evolutionary biology and vehemently defended by Mayr. It may have been his
experience at the IUBS meeting in 1966 that led Mayr to put together his famous
conference on the modern synthesis in the early 1970s; at that conference,
discontinuity was ruled out of court. 35
The three following IUBS conferences were just as well attended. Though
Mayr dropped out, he was replaced by a veritable who’s who of biologists,
including Francis Crick and John Maynard-Smith (1920–2004), philosophers
such as Marjorie Grene (1910–2009), theoretical physicists such as David Bohm
(1917–1992), and mathematicians, including Thom’s equally talented and better-
known colleague Sir Erik Christopher Zeeman (1925–2016). Though
Waddington found sympathetic comrades among the physicists and
mathematicians, he did not always see eye to eye with his colleagues in biology.
Maynard-Smith, for instance, ardently defended neo-Darwinian orthodoxy,
admitting its ties to the old Weismannian formula segregating germ and soma
modernized as the central dogma. Crick worried aloud about how far the
concepts raised during the IUBS meetings departed from what mainstream
biologists like him believed—all but guaranteeing the ideas generated at IUBS
would be treated as fringe. But Waddington had plenty of “third way” supporters
too. Brian Goodwin and Richard Lewontin attended more of the meetings than
anyone else and would themselves become champions of a new wave of
organicist thought.
Epigenetics was clearly at the heart of Waddington’s hope for this new
generation of organicists. Definitions, methods, and social hierarchies within the
approach of epigenetics pepper the four-volume proceedings that emerged from
the IUBS meetings. 36 René Thom attempted to describe in careful mathematical
language the epigenetic landscape, going so far as to create a physical model out
of clay with liquid running down the slopes; unintentionally, it turned out to be
eerily similar to John Piper’s painted rendering in Waddington’s Organisers and
Genes (1940). Clearly most attendees believed this was where many of the
mysteries in biology continued to hide. How do genotypes become phenotypes?
How do phenotypes then impact genotypes such that evolution is possible? In
the neo-Darwinian account so deeply impacted by Crick’s central dogma that
even Mayr—with his leanings toward organicism—could not seem to get around
it, the organism is a black box. Development in that regime is reduced to the
unrolling of a program in the genes without reference to the environment, except
to say that it provides certain basic conditions for the unrolling to happen. But
the concept of epigenetics coined by Waddington and given a full expression at
the four IUBS theoretical biology meetings moved away from the notion of
programs and the unidirectionality of information exchange from DNA to RNA
to protein to phenotype to population to phylogeny. Both top-down and bottom-
up explanations and exchanges would be required.
A new language, in fact, would need to be developed for “third way” biology.
This was the only consistent way to avoid the faulty assumption that Stanford
University physicist Howard H. Pattee detected in all of the molecular biology
literature he had read. Tacitly, at least, he found molecular biologists like Crick
enamored with “the classical idea of a . . . machine [as] a good physical analogy
to living matter.” 37 The problem originated in their misunderstanding of modern
physics and their unexamined theoretical assumptions, combined with imprecise
language:

What [physicists] find is that even the lowest interesting example of a hierarchical interface is beset
with precisely those difficulties that we find in all hierarchical structures, namely, that each side of
the interface requires a special language. . . . [L]iving organizations are not distinguished from
inanimate matter because they follow laws of physics and chemistry, but because they follow the
constraints of these internal, hierarchical languages. . . .
[A] large part of what is called theoretical physics is a study of formal languages, searching for
clear and consistent interpretations of experimental observations. Biologists have never paid this
much attention to language, and even today most molecular biologists believe that the “facts speak
for themselves.” . . . All these facts tell us at present is that life is distinguished from inanimate
matter by exceptional dynamical constraints or controls [that] have no clear physical explanation.
We will not find such an explanation by inventing new words for our description of each level of
hierarchical control. Instead, we will have to learn how collections of matter produce their own
internal descriptions. 38

Unbeknownst to Pattee, this was a cry that members of the Theoretical Biology
Club had uttered repeatedly since the 1930s: Convict the individual biologist of
his own intellectual confusion, and then resolve interdisciplinary contradictions.
Create an intelligible language to better encompass the complexity of the
evolving, developing organism. Pattee was channeling J. H. Woodger and Ian
Suttie.
Thus the IUBS theoretical biology meetings demonstrated to Waddington that
the organicist impulse was still alive and well in the mid-twentieth century, even
if prominent members of his own field failed to pick up epigenetics—even if
most biologists were implicitly committed to gene-centric mechanism. Barry
Commoner and participants at Alpbach in 1968 may have been justifiably more
concerned about the pervasiveness of mechanism and its potentially perilous
effects, but for Waddington, their attempts at addressing these worries
occasionally skirted into vitalism.
Within the two years between the Alpbach Beyond Reductionism symposium
and 1970, something changed for Waddington. Perhaps it was his own health
problems that began to crop up toward the end of the 1960s—he was an avid
pipe smoker and began to suffer various complaints seemingly related to that
behavior. Or perhaps it was merely impending retirement. In any case, any hint
of dismissiveness he expressed at Alpbach disappeared. Like Commoner—like
Carson, in fact—Waddington began to recognize more acutely the connections
between the dominant, if usually implicit, philosophy of mechanism and the
degradation of ecological and social resources. Needham, too, lifted his head
from Science and Civilisation in China to contribute, sporadically, to works
challenging the consensus mechanistic position. But it was again largely
Waddington who spearheaded the social-biological critiques in the 1970s that
emerged from the old organicist group. Whatever the potential concerns about
vitalism he expressed earlier, gene-centric mechanism now seemed the clearer
threat.

A copy of Waddington’s Behind Appearance: A Study of the Relations Between

Painting and the Natural Sciences in This Century (1970) towers over nearby
books on a library shelf. 39 The tome is simultaneously a rich amalgam of
modern abstract painting tessellated with photographs of bubble chambers and
jagged seismograph readouts. The book also contains one of Waddington’s
clearest expositions of Whiteheadian process philosophy. (Sadly, this particular
example rarely circulates. Some past patron or student who valued the full-color
prints displayed within, pearls within this gray-clad oyster, carefully excised all
its most famous reproductions—the Kandinskys, Magrittes, and Dubuffets—and
replaced it on the shelf. This copy of Behind Appearance is now a carved-up,
ironic symbol of what happens when the whole is not believed to be critical to its
parts.)
Despite appearances, Waddington’s Behind Appearance was no mere
dalliance. During the end of the 1960s, he apparently became convinced that
biological science was on the wrong trajectory after all—the IUBS conferences
were perhaps anomalies. Now society itself was beginning to follow that
misguided trajectory. There was no question in Waddington’s mind that the rise
of molecular biology was extremely important to science. But the larger
sociocultural changes molecular biology seemed to instigate, and that were just
now washing over Western civilization, seemed to be threatening the very values
that gave rise to science and civilization in the first place. He witnessed modern
biology visible in biochemical and molecular biology labs and saw that work
there neither required nor encouraged anything other than the absorption of
physicochemical facts that spoke “for themselves.” The cultural consequences of
such circumscribed values would be severe, if not immediately recognized. The
age-old questions about the nature of life, human culture, and sentience would be
trivialized in light of the promises of chemical, biological, and physical
engineering. The whole motivation to apply science in the first place—to
increase the flourishing of human society and the natural world—would bow
before the throne of mechanized efficiency, commodification, and the chase after
dollars and cents, pounds and pence. Behind Appearance was, among other
things, a paean to the importance of the human arts in highlighting the hue of the
organic whole visible through the very techniques that technoscience was in
danger of hollowing out—an attempt to show why C. P. Snow’s Two Cultures
still very much needed each other.
 15

THE CONVENTIONAL WISDOM OF THE

DOMINANT GROUP, OR COWDUNG

F or a brief period, the late 1960s seemed to Waddington like another early
1930s: a tipping point in favor of the “third way.” In his correspondence with
Arthur Koestler after the 1968 Alpbach meetings, Waddington expressed
confidence. He referred to the “new paradigm” to describe the organic notions
held by himself, Needham, Woodger, Bertalanffy, and a few American scientists.
It was on its way toward being “adopted by leading biologists.” But the fight
was not yet complete. In the UK, as in America, biologists were interested in
“the attempt to find some simpler elements in terms of which the complex can be
understood.” Waddington found nothing wrong with that formulation of science
—it was what he himself did in his work in Drosophila epigenetics. If it was
reductionistic, it was of a gentle sort, compatible with the “third way.”
In the United States, however, Waddington identified a reigning alternate—
wrongheaded in his view—definition: American reductionism “now implies that
we start with a full knowledge of the simple elementary units, and, on the basis
of this knowledge, can determine what the character of the apparently complex
really must be.” An astute scientist, by contrast, admits that “What little we do
know [of basic units] has been learnt from the study of the complexes into which
they enter. We must therefore be prepared to accept additions to our knowledge
about them; but equally it is good tactics to refuse to accept any alleged new
insight into the properties of our basic units until the evidence really forces us to
do so.” 1 Organicists must first be good empiricists. Assumptions that
fundamental genetic units give us insight into behavior, for instance, need to be
rigorously tested rather than presumed to be true. And any behaviors that can be
shown to have a strong genetic basis must still be viewed as integrated parts of
“complexes” and, more generally, organic structures rather than the programs of
master molecules.
The new awareness of social problems that typified the late 1960s and ’70s
soured Waddington’s confidence. Overpopulation would soon lead to one sort of
global catastrophe or another; overpopulation influenced traffic congestion,
unemployment, natural resource degradation, pollution, and so on. Each of these
factors exacerbated the other and enflamed international and interethnic conflict,
including the nuclear-tipped standoff between the Anglo-American and Soviet
blocs. After Alpbach 1968, Waddington turned toward these sociopolitical
predicaments. He admitted they were complex issues—a “complex of complexes
,” in fact. 2 Piecemeal solutions floated to address them created as many
problems as they solved. The world could use a systemic, hierarchical approach
that privileged processes over entities, relations over things. Unlike earlier in his
career, however—even as recently as the IUBS meetings—Waddington could no
longer be as sanguine that “third way” thinking was on the rise. An entrenched
gene-centric, mechanistic philosophy stood in the way of holistic solutions to
biological or social problems. In his final publications before his untimely death
in 1975, he railed against this knee-jerk mechanism—what he called the
“conventional wisdom of the dominant group.” 3

Waddington’s posthumously published Tools for Thought is an organicist’s

manifesto. The book playfully but forcefully dances across philosophy, biology,
mathematics, statistical theory, even computer science, reiterating the serious
misgivings Waddington harbored about the trajectory of twentieth-century
science. Unlike the final, semi-autobiographical works of some of his peers—
unlike Francis Crick’s What Mad Pursuit , for instance—Tools for Thought
expressed genuine skepticism that science in its molecule-focused form would
be able to address some of the most pressing problems in the planet’s eminent
future. Under the rule of the techno-scientific regime he saw exhibited most
strongly in America and increasingly in Europe, far too much intellectual energy,
not to mention money, had been, and continued to be, poured into finding
answers to short-term problems without a deeper investigation into the
philosophical roots of those problems. As a result, those short-term answers
actually created more difficult problems in the long term. Lasting positive social
and ecological change had become so fleeting because technocrats, even when
scientifically trained, began from a deficient worldview. Complicating that fact,
Waddington doubted whether modern scientists and technocrats recognized they
were acting out of any philosophy at all. Because they had been trained to value
the unending production of white papers, research articles, grant proposals, and
other brief, seemingly fact-of-the-matter publications, he rather doubted they
would be able to step back, reflect, and identify the deficiencies in their own
worldviews. Even that practice—thinking philosophically—had been dismissed
as frivolous, bad for the business of grant acquisition. Given that prejudice, most
followed what Waddington deemed the “conventional wisdom of the dominant
group” or “COWDUNG.” This was Waddington’s “memorizable, appropriate
and accurate” (and deliberately confrontational) acronym for their worldview. 4
COWDUNG was by no means new or particularly American, however, even
if it accompanied the rise in American scientific power after the Second World
War. Waddington, like Whitehead, found this worldview evident in the
philosophy of René Descartes. Like him, most biologists in the second half of
the twentieth century held to the view that the world “essentially consists of
things , and that any changes we notice are really secondary, arising from the
way things interact with each other.” But the world, according to Waddington’s
“third way,” “consists of processes , and the things that we discern are only stills
out of what is essentially a movie.” 5 His views were born from Alfred North
Whitehead and beyond to Heraclitus. 6 He realized, however, that the
Democritean-Cartesian view dominated the sciences; and it was the
“commonsense” view held by the vast majority of the Western public in
capitalist societies. Part of the issue undoubtedly had to do with the scale of each
science. In geology, astronomy, chemistry, and the like, “It seems much simpler
to regard [chemicals, rocks, etc.] not as processes but as things, and to get down
to the practical problem of finding out how these things interact with one
another, to bring about the essentially secondary processes.” “There are,”
Waddington admitted, “many contexts in which the ‘thing’ view is the sensible
one to adopt.” But the Democritean-Cartesian scientists did not remain strictly
chemists or astronomers. In the early twentieth century, they migrated into
biology and the social sciences; their numbers increased rapidly during the
middle of the century in which molecular biology germinated. These migrants
applied their own COWDUNG to living organisms and, unsurprisingly, saw
organisms “as a set of chemical and physical interactions going on between
essentially unchanging things.” Waddington had, of course, been surrounded by
advocates, witting and unwitting, of Democritean-Cartesian COWDUNG for his
entire post-WWII career. 7
The problem in biology was that there were so many processes and networks:
“But still, powerful though this [thing] approach is, it has so far really only been
successful in connection with some of the questions we want to ask about living
things, not all of them. It has given us little understanding of embryonic
development; little except some rather empty theories about evolution; and
hardly anything at all about the mind.” 8 Theories about evolution and humans,
minds, and the interactions of minded humans had been just the sorts of topics at
which the “third way” had been aiming during the twentieth century.
COWDUNG predominated, however, and Waddington could not pretend in this
context that his views had been persuasive.
Aside from the COWDUNG invective, Waddington rarely acknowledged in
print his own intellectual distance, even isolation, from his colleagues after
World War II. He did, however, wonder openly about the resistance to the “third
way.” Why had the organicism held by Whitehead, Woodger, Needham,
Haldane, Ritter, Russell, and Lloyd Morgan and so many others not been
successful in displacing or at least muting the pull toward Democritean-
Cartesian thinking? It was no body of experimental data that determines the truth
of that philosophy. “[R]eductionism is a recipe for action . . . if you are
confronted with a complex situation, for instance a living system, your best bet
to get some sort of pay-off or other is to look for the physical or chemical factors
which can influence the phenomenon in question.” 9 The Democritean-Cartesian
worldview, in other words, made for “lousy philosophy,” but when what is called
for is making a “quick (scientific) buck by discovering some useful practical
information,” it trumped the slow, complex search for multiple levels and a
plurality of inputs. Even if, according to Waddington, organic philosophy
remained the “method for making major advances in human comprehension”—
those of “Darwin, Freud, Einstein [and] the quantum physicists,” for instance—
reduction to simple mechanisms promised finality, predictability, and the
supposition that scientific discoveries grasped “the whole of what is contained in
the reality independent of ourselves.” It also promised the ability to commodify
biological products. Waddington regarded these promises as hollow, since they
rested on the “Fallacy of Misplaced Concreteness in its simplest form”—a fact
most scientists refused to see. 10 For them, the persuasiveness of reductive
mechanism for action, control, patenting, product development, and so on, was
hard to refute. 11
Waddington repeated these points in a review of E. O. Wilson’s Sociobiology
for the New York Review of Books in August 1975. 12 In one of the most ironic
episodes of his career, the anti-sociobiology Science for the People (SftP)
organization took Waddington’s review to be largely supportive of sociobiology
and cited Waddington’s review as their reason for attacking Wilson. 13 What
Waddington’s critics missed, and what subsequent reprintings of his New York
Review of Books review leave out, is that Waddington’s review included two
books. One was Wilson’s Sociobiology ; the second was Biogenetic
Structuralism by Charles Laughlin and Eugene G. d’Aquili. 14 Once read in its
proper context—treated as a whole organism, in other words—one can see that
Waddington was actually critical of Wilson’s gene-centric mechanism. In fact,
Sociobiology: The New Synthesis was, for Waddington, a sterling example of
COWDUNG. Unfortunately, Waddington died on 26 September 1975, days after
SftP published their own condemnatory review of Wilson’s Sociobiology in the
New York Review of Books. Waddington did not live to extricate his own essay—
tellingly entitled “Mindless Societies”—from Wilson’s promotion of
sociobiology. Unfortunately, historians of this episode have ignored both the
substance and the real nature of Waddington’s review and, therefore, have
promoted the concept that he was largely supportive of Wilson. “Mindless
Societies,” however, is properly read as the product of an older but still active
scientist struggling to have his biophilosophy taken seriously by those at the
center of the scientific mainstream.
A series of heart attacks in the late 1960s forced Waddington to cut back on
his very full traveling and speaking schedule, but he continued to write reviews
of books and popular articles even from his hospital bed. 15 He found motivation
through the fear that the scientific adoption of organicism that he had once
assumed would come to pass was in fact slipping away. Needham, though still
active, wrote little for the scientific community. Woodger was now wholly
engaged in analytic philosophy and far from the network of scientists with whom
he had once interacted. Once that generation had quit the field entirely, would
the “pre-reductionist Whiteheadian thought” and the deep-seated implications
the “third way” had for “general equality [and] freedom” find no champion in all
of science? 16 The consequences, he thought, would be profound. The
conventional wisdom ignored holism in favor of the conformity of “units of
personnel” that could be more efficiently processed and moved from point to
point. In “The Age of Conformity: The Autocatalytic Loss of Freedom,” for
instance, Waddington stressed that this push for control over the human and
nonhuman environment inherent in the mechanistic conventional wisdom was
already leading to depersonalization and the curtailing of optimum human
flourishing in megalopolis environments. 17
As Waddington began to recover in 1970, he and Margaret Mead gathered
social and life scientists, urban planners, even musicians, to a symposium in
New York called “Biology and the History of the Future.” Their goal: to discuss
how an organicist worldview could combat the rash of social problems that
seemed to be sweeping over the Western world. 18 Consensus at the conclusion
of the conference, not surprisingly perhaps, was that the COWDUNG of
reductionistic mechanism, though necessary to deal with enormous aggregates of
genes and atoms, led scientists and technocrats to ignore individualities,
development, networks, and patterns. While probably acceptable, even crucial,
when dealing with physicochemical units, this Democritean-Cartesian
philosophy could not really grasp the intricacies of growing living things, let
alone behavior, social dynamics, or ecological systems constantly in flux.
This acute awareness of what conventional wisdom brushed aside motivated
Waddington’s misunderstood critique of Wilson in 1975 as well. In a tone that
may have proved to be too gentle for the SftP group, Waddington faulted Wilson
for merely waving his hands at epigenetic mechanisms such as genetic
assimilation and phenotypic adaptation. This was a mistake, according to
Waddington, because epigenetic adaptation is merely “another way of referring
to learning, by a phrase broad enough to encompass modification of physical
structure as well as of behavior.” In Sociobiology , Wilson completely
disregarded epigenetic “learning” and symbolic (as opposed to chemical)
communication. How then could sociobiologists claim that colonial insects such
as ants and termites, Wilson’s exemplars, tell us anything about learning and
communication among more complex species, as Wilson insisted they did? For
Waddington, organismal development and symbolic communication were more
hazardous to Wilson’s sociobiology than even the problem of altruism on which
Wilson and other sociobiologists usually focused their attention. Waddington
believed Wilson was “guilty of at the very least a certain incoherence, in
indulging himself in a delicious mental frisson of Camusian alienation, while
denying [higher organisms] . . . even a hint of mentality.” 19 For Waddington,
there was a telling reason why Wilson and the other sociobiologists were
deliberately obviating direct discussions of symbolic communication, epigenetic
adaptation, and learning, as well as any mention of “mind.” If they took these
obvious aspects of organisms more seriously, sociobiologists would be forced to
concede the inadequacy of their gene-centric, mechanistic claims. They would
be forced to acknowledge that there was an alternative.

Despite his broad connections across biological subdisciplines, Waddington

received no formal Festschrift, and no major biography was ever published of
his life and works. A colleague at the University of Edinburgh, Alan Robertson,
handled the single large memorial that was published in the Biographical
Memoirs of the Royal Society. Robertson was not especially familiar with
Waddington’s prewar work and evidently did not think highly of organic biology.
20 Other colleagues were less skeptical of theory in general, but still did not see
Waddington as a major figure. David R. Newth, biologist at the University of
Glasgow and editor of the Journal of Embryology and Experimental Morphology
, found Waddington’s theoretical musings interesting. But he was not uniformly
admiring of Waddington. He later apologized to Joseph Needham for his
statement, published in the Waddington “remembrance,” that “Waddington was
casual to the point of irresponsibility in matters that failed to interest him.” 21 In
part, Newth accounted this lack of focus to Waddington’s multifaceted
background in geology, philosophy, and experimental embryology. Waddington,
to put it another way, did not have the narrow disciplinary pedigree of a typical
biologist by the 1970s.
In the absence of a grand acknowledgment by his peers, Waddington
assembled and edited many of his papers that seemed most definitive of his
legacy. He entitled the collection Evolution of an Evolutionist —an intentionally
assertive title given his peripheral standing in the evolutionary biology
community at the time. With the exception of a brief introductory
autobiographical sketch where he pronounced the influence of Cambridge
biology and organic philosophy on his subsequent interests, Waddington selected
the papers that would position himself simultaneously as orthodox biologist and
heterodox challenger of the gene-centric neo-Darwinian establishment. He
interleaved, for example, his pre-WWII canalization paper with others from the
mid-1950s explaining genetic assimilation of acquired characters. 22 Strangely,
he omitted his 1942 essay “The Epigenotype”—the ur-paper for epigenetics and
the article for which Waddington is most remembered today. His collection was
conservative; even in his autobiographical highlights, he downplayed some of
his most radical contributions. Those unfamiliar with the rest of his career could
be forgiven for regarding him, as philosopher Marjorie Grene apparently did, as
a not “wholly orthodox mechanical materialist”—but little else. 23
Broadly speaking, the image of Waddington impressed upon the world both
by his colleagues and by himself at the end of his life and career was the image
of a philosophically astute biologist with a plethora of “social parts” that were
kept “in more-or-less water-tight compartments.” 24 His other, more unorthodox
interests—his devotion to “third way” philosophy, his promotion of modernist
architecture and artwork, his concern for global environmentalism, and so on—
were relegated to footnotes. In other words, Waddington was just a biologist;
largely because of his marginal status to the conventional wisdom of the neo-
Darwinian synthesis, his work retained only historical interest.
Tragically, during his last days, he seemed to sense the possibility that his
work was drifting toward irrelevance. In his final article, published
posthumously, Waddington decried the widespread popularity of the
conventional wisdom and his inability to tarnish its popularity. He was grateful
that he had read Whitehead when he did, before the organic philosophy had been
“pushed out of fashion by Wittgenstein and [Bertrand] Russell.” With the
encouragement of Woodger, Needham, and Gregory Bateson, he had heeded
Whitehead’s warning regarding “‘The Bifurcation of Nature’ into Mind and
Matter, set against one another as two totally separable and incomparable
essences.” 25 He had avoided the philosophical naïveté Whitehead had
condemned. If he had bequeathed anything to the future, it was the hope of
epigenetics, perhaps rooted in a “third way” rejection of mechanism. When
realized, the science of epigenetics would serve as an admonition to consider
wholes, intrinsic relationships, inseparability, cells, tissues, processes, the
environment, and the organisms in that environment—in a word, the vastness of
biological complexity, a complex of complexes—as much as the folding,
unfolding, and copying of tangled DNA threads. But upon his death in the mid-
1970s, these hopes seemed unfounded, the dream of those, like Woodger and the
Needhams, who had ignored the conventional wisdom of the dominant group.
 CONCLUSION

A THIRD WAY AFTER WADDINGTON?

P erhaps now I can attempt to answer the questions raised in this book: Why did
scholars pursue organicism in biology over the better part of the last century?
How did it become so important in the interwar period? Why did organicism
become a marginal concern by the last quarter of the century? What does this
alternative to mechanism and vitalism have to do with epigenetics? In this
conclusion, I would like to address these questions more explicitly and then
return to ask yet another one: Why does this story matter for the history and
philosophy of science?
The answer to the first two questions—why did it originate and why did
biologists from multiple nationalities and over multiple decades arrive at such
similar concepts—has to do with the answer to the third question: Why did it
seem to decline in the last three decades of the twentieth century? To illustrate a
possible answer to that third question, I turn to one of the most ambitious books
of the late twentieth century written by one of its most important science writers
—a work that addressed the philosophy of biology generally and the mechanism,
vitalism, organicism debate more specifically: Douglas Hofstadter’s Gödel,
Escher, Bach: An Eternal Golden Braid (1979).

At first glance, it seems improbable that Gödel, Escher, Bach won the Pulitzer
Prize in 1980 or that Time magazine placed the book among the All-TIME 100
Best Non-Fiction Books since 1923, alongside Keynes’s General Theory and
Shirer’s Rise and Fall of the Third Reich. 1 Hofstadter’s book was nothing like
these famous works. The young computer programmer adopted the meandering,
almost nonsensical style sported by Lewis Carroll in Through the Looking Glass
to pirouette over challenging puzzles from music, art, physics, entomology,
neurology, and mathematics. He made no larger political points—the book isn’t
Richard Wright’s Black Boy or Dee Brown’s Bury My Heart at Wounded Knee
(both also on the list). If Hofstadter made a central argument, it was that artificial
intelligence (AI) was possible and should be pursued.
In the core chapter of the book (he called it a “metaphoric fugue,”
reminiscent of Bach) he attacked the single most persistent objection to AI:
Human minds are irreducibly complex and thus minds cannot be reduced to
machine behavior. To refute this objection, Hofstadter employed an Anteater, a
Crab, a Tortoise, and the warrior Achilles. These four characters comically
rehearsed the dispute between those who believed life and mind were
exceptional and those who regarded them as fully reducible to physics and
chemistry. 2 It was another performance of the old conflict, this time written in a
dizzying, palindrome-filled voice.
Upon further examination, Gödel, Escher, Bach does seem to belong among
those classic works that have captured and defined a moment in time. This
metaphoric fugue—the core chapter of Gödel, Escher, Bach —reflected the
conventional wisdom of the dominant group. Hofstadter had been trained in
physics and interested in AI rather than organisms, so he turned to sociobiologist
E. O. Wilson for assistance in finding an apt biological illustration for his
argument about the construction of order out of disorder. For Hofstadter’s
mechanistic account to work, he would have to explain the possibility of
complex behavior as a straightforward outgrowth of simple behavior. In other
words, he would have to show how the apparent complexity of human thought
could be attained by much simpler computer mechanisms if AI was to be
successful. Wilson suggested that Hofstadter use the analog of ants and their
colonies: Ants were ideal for the central fugue in Gödel, Escher, Bach because
they exemplified the potential for great complexity, yet individual ants were
clearly mindless machines. When grouped, these ant-machines produced
sophisticated behavior resembling human-like planning, preparation, and
problem solving.
Hofstadter ran with Wilson’s ant suggestion. Like ants, he said, human
neurons do not think; they are tiny wet machines. But like ant colonies,
collections of neurons in a brain do think, or at least appear to. Hofstadter
veered away from any account of origins, a direct discussion of genes, for
instance—his interests lay in using computers to replicate human thought rather
than explanations in biology per se. The broader implications of his illustration
were obvious, however. It is within the realm of possibility that mindless ants
could create complex, seemingly mind-possessing wholes by aggregating
random behavior across the slightest of hierarchical distinctions (e.g., worker,
soldier, queen). Using Wilson’s ant analogy and his own unconventional style,
Hofstadter offered what was basically a very traditional mechanistic answer to
complexity. While anti-mechanists—some of the participants in Waddington’s
IUBS symposia in the late ’60s, for example—pointed to the empirical evidence
of intricacy and coordination, mechanists maintained belief in underlying
simplicity and randomness. Ant colony behavior only seemed to be
superorganic, more advanced than the sum of its parts. In truth, each member
merely worked out its programming without a greater plan or organization. By
analogy, a brain only seemed to be irreducibly complex. In actuality, each neuron
was a simple electrochemical switch. This whole-is-merely-the-sum-of-its-parts
argument could be extended to amazingly complex things—even Bach’s fugues
and Escher’s mind-bending sketches. Advanced human culture required only
small tweaks to cellular machinery. Connecting these dots, Hofstadter led his
audience to see AI as attainable—not far off, in fact. It merely required
differently arranged but otherwise uncomplicated computer programs.
Hofstadter showed some hesitation when discussing ant communication,
however. At first glance, this was a hard nut to crack from a strictly mechanistic
point of view. How could practically mindless ants convey meaningful messages
without an overall design or purpose? In Hofstadter’s field of AI, this would
cause no problem: The computer engineer was the efficient cause, shaping
programs to fit the final purpose. Instead of admitting his analogy broke down,
however—that ants and brains really are different than computer programs—
Hofstadter doubled down on mechanism. We speak as if there is design behind
symbolic communication, order out of disorder, brains from bundled neurons
and so on, or as if there is some larger organic principle operating in the
background, but we know there is not. The solution to the conundrum of ant
communication, thought Hofstadter (channeling Wilson), required a more
perceptive perspective: “Achilles : . . . From an ant’s-eye point of view, a signal
has NO purpose. The typical ant in a signal is just meandering around the
colony, in search of nothing in particular, until it finds that it feels like stopping. .
. . No planning is required, no looking ahead. . . . But from the COLONY’S
point of view, [the ant] has just responded to a message, which was written in the
language of the caste distribution. Now from that perspective, it looks very much
like purposeful activity.” 3 Whether ant colonies were purposeful—and by
analogy whether neurons firing in brains were purposeful—depended upon
perspective. Hofstadter’s Crab briefly protested that one needed to invoke holism
in order for these obvious differences between levels to make philosophical
sense. Crab’s line unintentionally echoed the nearly century-old sentiments of
the first organicists and, more recently, some of the IUBS participants’
comments. But Anteater, the Wilson-esque mouthpiece of Hofstadter’s fugue,
immediately ruled Crab’s appeal out of court. At the lowest level, he countered,
stochastic mechanism dominated. You certainly wouldn’t want to attribute
thinking to ants! Where would the thinking come from, given the tiny size of
their heads, the miniscule number of neurons inside their exoskeletons?
According to Anteater, order appeared out of this disorder in the same way
bubbles appeared out of boiling water. Just as no one would find any greater
meaning in those bubbles formed according to ordinary physical laws, one could
not call on “‘higher-level laws’” to explain phenomena such as ants moving to
and fro working for the good of the collective. 4 Over millions of years, nature
selected mechanisms that conferred greater benefit on each individual ant and
piled those mechanisms on top of each other until whole functioning ant
societies resulted. Communication could only be understood by understanding
the biological principles underneath it. 5
Curiously, after making this mechanistic assertion, Hofstadter partially
backed out of it. His hesitation is just as revealing for understanding the fate of
“third way” biology in the last quarter of the twentieth century as anything else.
Rather than taking a more straightforward position on whether the whole is
greater than or exactly equal to the sum of its parts—that is, whether an organic
alternative appealing to higher-level laws or ordinary reductionistic mechanism
provided the most complete explanation—Hofstadter waved his hands. Instead
of deliberating about the issue further, his characters pronounced mu. “Mu” was
a reference to the Zen doctrine that “only by not asking such questions can one
know the answer to them.” 6 Only by avoiding the philosophical knot presented
by organisms and minds could one cut through it, he implied.
Gödel, Escher, Bach was awarded the Pulitzer Prize because it presented
weighty puzzles in computer science, mathematics, and the philosophy of mind
via lighthearted literary devices, vignettes of Bach, and Escher’s provocative
illustrations. Hidden at the whimsical book’s core, however, was the serious
point that mechanistic science ensured progress and that statements to the
contrary were retrogressive. This claim alone was not new or surprising. Jacques
Loeb had leveled similar charges against Driesch at the beginning of the century;
Francis Crick’s Of Molecules and Men repeated Loeb’s line a half century later
with the well-funded confidence of a molecular biologist. The difference in
Hofstadter’s presentation—the reason why his book is such an apt example of
the fate of organicism—was not the appeal to mechanism itself, but the
conjunction of this claim with a general skepticism about philosophy’s ability to
judge these sorts of scientific claims. Hofstadter dressed up his skepticism in the
garb of Zen Buddhism. Zen was a red herring, however, meant to soften a
relatively standard late-twentieth-century anti-philosophy.
 In effect, Gödel, Escher, Bach portrayed the mechanistic story as the right
one. But rather than face the century’s worth of philosophical challenges to
mechanism—some of which have been detailed in this book—Hofstadter
declared Western philosophy incompetent to deal with the issue at all. By taking
this stance, Hofstadter anticipated E. O. Wilson himself. As the elder statesman
of entomology quipped in a 2009 interview, “[I]f I were a good philosopher, I’d
say it’s an unanswerable question. But I’m a biologist, and I say it’s answerable.
. . . One by one, the great questions of philosophy, including ‘Who are we?’ and
‘Where did we come from?’ are being answered. . . . So gradually, science is
simply taking over the big questions created by philosophy. Philosophy consists
largely of the history of failed models of the brain.” 7 Though once fruitful
territory for inquiry, fundamental philosophical questions in biology had passed
out of the realm of philosophy—a field probably incompetent to deal with them
anyway, according to one of the best-known biologists of the late twentieth
century. Thus mechanism through the action of selfish genes could be assumed.
If philosophical argument from those who believed an anti-mechanistic
alternative existed questioned that position, it could be safely ignored.
The answer to why organicism briefly faded from view, then, is not merely
that mechanism provided a watertight explanation. Rather, philosophical
questions such as these were judged irrelevant to biology. In that environment,
mechanism could be tacitly promoted—assumed, in other words—without
having to be proved or even argued.
Waddington, if he had been alive, would have pointed out the poverty of this
COWDUNG assertion. “Philosophy of one kind or another cannot be avoided or
evaded or given up like sin in Lent,” stressed Waddington. We should prime
ourselves to look for concealed philosophical commitments perhaps especially
because the “Conventional Wisdom of the Dominant Group [says] that we don’t
have to fuss about philosophy.” 8 In other words, the more the dominant group
proclaims it has no philosophy or that philosophy is irrelevant, the more insistent
we should be that it is in fact promoting some philosophy.
Why, then, did a “third way” rise and come to the attention of scientists and
philosophers across Europe in the first place if it could be so easily dismissed by
the late twentieth century? There are several promising reasons to explore here.
First, many scientists expressed disappointment in the overconfident statements
of the mechanists themselves, who assured their audiences at the turn of the
century that simple mechanistic explanations for every organic process would be
forthcoming in the very near future. 9
Secondly, proponents of organicism from J. S. Haldane to Joseph Needham
repeatedly appealed to empirical observations for support for their position.
When it came to organisms, complexity certainly did not appear to spill forth
from physicochemical simplicity, as mechanists claimed. Their own repeated
observations—the very empiricism enshrined in the so-called “scientific
method,” in other words—militated against the mechanists. As E. S. Russell put
it: “[S]tart from conceptions which are tacitly followed by most biologists (I
except biochemists!) in the practical conduct of their researches, and, accepting
the broad facts of observation at their face value, to generalize these without
paying much attention to ultimate questions of philosophy. We thus acquire a
provisional concept of the organism which gives us certain principles for
biology, the final test of which must lie in their success or failure in practice.” 10
Back in 1919, William Ritter, too, stated this desire to make empirical
evidence the foundation of biological theory—to uphold the concrete rather than
elevate the abstract, as mechanists did, or the completely intangible, as vitalists
did:

Ideas or psychoids or entelechies or “principles” of any kind conceived as independent of, or even
separable from, sensible objects are quite as repugnant to me, an organismalist, as they are to any
[mechanist]. The essence of my contention is that the natural substitute for these imponderable
things are the living, individual organisms themselves , and not the particles of which they are
composed. Each and every individual organism is a natural reality by exactly the same criteria that
the atoms, molecules, cells, and tissues of which it is composed are natural realities. 11

There should be no reason, these theorists believed, to regard observations about
the wholeness of the organism’s interdependent systems as somehow illusory.
Thus, from an empirical adequacy standpoint, advocates of the “third way”
believed they, not mechanists, held to the traditional standards of science.

Pursuit of the “third way” did continue after Waddington. Notably, the loudest
arguments for an alternative to mechanism and vitalism came from the pillars of
the neo-Darwinian synthesis and their students: paleontologist George G.
Simpson (1902–1984) and his student Stephen Jay Gould (1941–2002), and
geneticist Theodosius Dobzhansky (1900–1975) and his student Richard
Lewontin (1929–). 12 Simpson adopted the language of other “third way”
thinkers arguing for an expanded set of physical laws, much like J. S. Haldane
and Joseph Needham had:

Insistence that the study of organisms requires principles additional to those of the physical
sciences does not imply a dualistic or vitalistic view of nature. . . . [L]iving things have been
 affected for . . . billions of years by historical processes. . . . The results of those processes are
systems different in kind from any nonliving systems and almost incomparably more complicated. .
. . Biology, then, is the science that stands at the center of all science . . . and it is here, in the field
where all the principles of all the sciences are embodied that science can truly become unified. 13

Dobzhansky, too, inveighed against the creep of genetic determinism he saw
toward the end of his life: “Heredity is not a status but a process. Genetic traits
are not preformed in the sex cells.” 14
Influenced by sentiments such as these and the direct influence of
Waddington, Gould and Lewontin worked over the course of their careers in the
last quarter of the twentieth century to challenge conventional wisdom wherever
they encountered it. Their landmark essay, “The Spandrels of San Marcos and
the Panglossian Paradigm,” was one clear example of this commitment. The
article undercut the dominant neo-Darwinian perspective that emphasized
gradual evolution based on gene mutations. Using the flying buttresses of
Venice’s most famous landmark as their grand metaphor, they pointed out that
most change is decorative—not adaptive, not structural. Important evolutionary
change, by contrast, is a whole organism change in form. The epigenetic
landscape model illustrated in Needham’s Biochemistry and Morphogenesis
(1950) and Waddington’s The Strategy of the Genes (1957) depicted what
“punctuations” in phenotypes would look like at the individual organism level.
The notion of punctuated equilibrium upon which Gould and Lewontin built
“Spandrels of San Marcos” scaled this model to the phylogenic level. Likewise,
they insisted that evolution occurs through switches, rapid punctuation events
followed by long durations of coherence and stability. 15 Thus the model of
phyletic change for which Gould became well known had its roots buried in the
soil of the organic philosophy cultivated by the Theoretical Biology Club.
Inspired by Waddington’s IUBS theoretical biology symposia in the late
1960s, Lewontin went on to become one of the most vociferous proponents of a
third generation of organicism. He wrangled with sociobiologists and attacked
those who would tie human intelligence to race through the 1970s and ’80s. In
books such as Biology Under the Influence, Biology as Ideology , and Not in Our
Genes , Lewontin and his coauthors extended the critiques of “third way”
proponents going back through the entire century: “It is not that the whole is
more than the sum of its parts. It is that the properties of the parts cannot be
understood except in their context in the whole. Parts do not have individual
properties in some isolated sense, but only the context in which they are found.”
16 Between the two of them, Lewontin and Gould carried the legacy of

organicism through the end of the century.

 Given that individuals like Lewontin and Gould were hardly marginalized
biologists in the last few decades of the twentieth century, and that they argued
just as fiercely against the conventional wisdom of the dominant group in their
day as Woodger, Waddington, Needham, and others did in theirs, how can we
continue to believe that organicism died out entirely?
There is at least one reason—well supported by historical evidence—to
believe that indeed the “third way” declined in a substantial way after the death
of the original TBC members. This reason has to do with changes to biology
education and funding. Few cited this reason directly; it became apparent to me
in the research for this book—through reading late-career memoirs, obituaries,
and scientists’ reflections upon the history of their fields. Those biologists that
came of age in the interwar period and earlier—certainly the first generation of
Lloyd Morgan, Russell, and Haldane, but also the Needhams, Waddington,
Woodger, Mayr, Hamburger, Bertalanffy, Medawar, Zuckerman, Raymond Pearl,
and so on—were exposed to history and philosophy hand in hand with their
scientific training. Scientific education changed dramatically over the course of
the twentieth century, especially in the United States—a country that has had one
of the most extensive and influential scientific programs in the world. It is fair to
say that few working biologists today have educational experience akin to their
disciplinary ancestors. Few practicing biologists know much, if anything, about
the history and philosophy of their science outside of the pantheonic names and
stories directly related to that work. This tradition of avoiding history and
philosophy of science is not completely new, of course; recall Raymond Pearl’s
warning to Woodger in the 1920s that “in America, [biologists] do not like to
think and look with a very fishy eye on anything which savors of philosophy.” 17
Moreover, it is almost cliché to point out that by the late 1950s, C. P. Snow—
incidentally, a close friend of Bernal and Julian Huxley, among others—worried
that the humanities and the sciences represented “Two Cultures” pursuing
divergent goals, viewing the world through incompatible lenses. 18
That being said, two decades after Snow’s famous address, Waddington
wondered if familiarity with the Whiteheadian philosophy of organism had
rendered him an outsider in his own field—as if because of his interests in
history and philosophy of biology, his work was less likely to attract attention. 19
At roughly the same time, Tots & Quots organizer Solly Zuckerman commented
to Peter Medawar that laboratory biologists discouraged their doctoral students
from looking back at “anything older than five years.” 20 The trend of dismissal
by biologists of the history and philosophy of their field continues to rise. E. O.
Wilson manifested this pattern of balkanization in his patronizing comment from
2009 that philosophy is “largely the history of failed models of the brain.” This
sort of philosophical know-nothingism is alive and well in the twenty-first
century, reiterated by famous science popularizers ranging from Neil deGrasse
Tyson to Bill Nye. 21
Maybe the most telling evidence in support of this claim about the lack of
historical and philosophical perspective in the life sciences is visible through
those cases that are exceptions to the rule. Those modern scientists who are
intimately familiar with the history and philosophy of their fields, such as Scott
Gilbert, Mary-Jane West-Eberhard, Brian K. Hall, Eva Jablonka, and Brian
Goodwin, typically find alternative accounts more appealing than the
conventional wisdom of the dominant group.
There could be good reason for this bifurcation of biology from the history
and philosophy of biology. In a less-often-quoted passage from The Structure of
Scientific Revolutions , Thomas Kuhn elegantly summarized the tendencies in
the natural sciences that render them more or less separate endeavors from fields
such as history and philosophy. Take physics, for example: There is little
incentive for the physicist to read the deeply philosophical works of Newton, or
even Einstein and Schrödinger. As Kuhn insisted, “Everything [a physicist]
needs to know about these works is recapitulated . . . in a number of up-to-date
textbooks.” 22 These textbook treatments invariably strip away explicit
philosophical claims at the cores of these historical works. Adopting the
questioning stance of those natural philosophers who created modern science—
Michael Faraday, William Whewell, Joseph Priestley, and so on—might in fact
erect an unnecessary roadblock in one’s early career, according to Kuhn:
“Because the unit of scientific achievement is the solved problem and because
the group knows well which problems have already been solved, few scientists
will easily be persuaded to adopt a viewpoint that again opens to question many
problems that had previously been solved.” 23 Confidence in solved problems
pays dividends in a way that continued questioning does not. Thus, as Kuhn
pointed out, science is profitably insulated from the overly critical, even
skeptical, spirit that typifies history and philosophy. If scientists make
philosophical claims, they usually do not do so intentionally.
Nevertheless, history and philosophy of science matter for scientists because
sometimes scientists do make broader claims—claims that require historical and
philosophical knowledge or at least an understanding that there are historians
and philosophers who have a better understanding of this material. Even if we
disregard off-the-cuff remarks like Wilson’s, even if Kuhn is generally correct
that scientific education properly insulates up-and-coming scientists to the
benefit of scientific practice, scientists contextualize the relevance of their
scientific work by pointing at history and philosophy. As physicist Pierre Duhem
stressed, for instance, “The physicist is compelled to recognize that it would be
unreasonable to work for the progress of physical theory if this theory were not
the increasingly better defined and more precise reflection of a metaphysics: the
belief in an order transcending physics is the sole justification of physical
theory.” 24 Moreover, scientists are often utilizing some sort of philosophical
approach in the laboratory whether or not they know it.
A half century after Duhem, just before Kuhn published Structure , physical
chemist Michael Polanyi emphasized the fundamental embeddedness of
theoretical concepts at the earliest stages, even in supposedly “non-theoretical”
laboratory work. We should certainly not envision biology, for example, to be an
intellectual edifice built solely from detached knowledge stacked like bricks.
Polanyi saw scientific knowledge as perpetually renewing, facts as ornate,
handcrafted by scientific artisans. 25 While perhaps the cold regularity of physics
makes it easier to craft theory, we find that many major biologists through
history understood clearly the philosophical ramifications of their benchwork—
those attending Waddington’s IUBS theoretical biology meetings were only a
small sample. When Hofstadter pronounced mu on the conflict between
mechanism and anti-mechanism—after advocating Wilson’s mechanistic
philosophy—he severed this discussion from its historical and philosophical
roots. He not only denied that this was an issue biologists could solve, he turned
to sweep away his footprints, denying that he was endorsing a particular
philosophy at all. The clear advantage of such an argument—a philosophical
argument that one is not making a philosophical argument—is that it is
practically unassailable. You might be conjuring a philosophical argument to
attack me, but I am just doing science.
While the structure of scientific education may mask the claims scientists
make and discourage them from learning the history and philosophy of their
fields in the first place, changes in funding serve as an even more powerful
practical reason why the bifurcation persists to the possible detriment of careful
theory construction. Scientists must justify the potential relevance of their work
to receive funding to pursue it, often appealing—hat in hand—to bureaucrats.
Waddington complained about the “quick (scientific) buck” in the 1970s.
Perhaps his younger colleagues in that era were taking the path of least
resistance rather than attempting to understand living systems in “their more
complex and richer forms”—rather than imitating the science of Charles Darwin,
in other words. 26 At our point in history, however, quantity seems clearly to
have bested philosophical sophistication. Our increasingly neoliberal age
demands that scientific publication rates increase in lockstep with financial
investment. 27 The broader intellectual content or, rather, the ramifications of a
scientific study to an old philosophical question such as, “How are living things
different from nonliving?” no longer appear significant to funding agencies such
as America’s National Science Foundation (NSF), if they ever did. Presently,
when the NSF asks a grant applicant to identify “broader impacts,” as it does in
every grant application, the NSF means it wants to know the project’s feasibility
to be made applicable to an existing sociocultural, economic, health-related, or
technical issue, not necessarily its relevance for long-term theoretical questions
in that field. 28
The answers to the first three questions I posed above, then, are relatively
clear if not encouraging. The rise of the organic philosophy occurred at a time
when, at least in training and funding, biology and the history and philosophy of
biology were not so far removed from one another in their guiding questions.
The decline of organicism in the 1970s mirrors a global trend toward
balkanization via changes in education and funding, pinned to still larger social
values. These values dissuade scientists from pursuing historical and
philosophical training or addressing broad, contemplative theoretical issues.
These large theoretical issues do not easily fit a rapid publication pace nor enable
biologists to easily compete for funding.
“Scientist” was once William Whewell’s nineteenth-century neologism for
“natural philosopher.” There are few obvious indicators left that philosophers,
historians, and scientists share such a recent common ancestor.

What is the significance of the present historical study? Does organicism offer a
truer depiction of the world than either mechanism or vitalism? While the central
figures explored in this book uniformly rejected vitalism, they thought of
mechanism quite differently. All wanted to reform mechanism, not dismiss it. As
a methodological approach, they believed, mechanism was crucial. Waddington
and Needham, for instance, expressed this point quite clearly in their final
works; in this, they echoed their predecessors, from Henderson, E. S. Russell,
and J. S. Haldane. For biology to grasp a phenomenon, just to get a handhold on
a problem, we often must break complexities down into simpler parts. We then
reassemble what we’ve broken apart. This kind of analysis is fruitful; it gives us
a more complete picture of the world. However, when that impulse expands
beyond this sort of gentle methodological stance—when it hardens and spreads
into metaphysics—it conflicts with the conviction that the whole complex
organism matters and that the environmental context matters to that organism.
For better or worse, we confront a world full of whole things, whole organisms.
While methodological mechanism helps us understand those whole things,
metaphysical mechanism implies that wholeness is not crucial, that a different
formulation with the same parts would produce only superficial differences, and
that form as a function of the parts is not intrinsic. Ultimately, it proclaims
wholeness as illusory—just as Hofstadter’s Anteater did. The whole human
brain, just like the whole ant colony, really is just the outworking of random
physicochemical collisions between mindless, haphazardly arranged parts. What
this metaphysical mechanism reflexively implies about the content of the ideas
that emerge from that wet-machine human brain—about the idea of mechanism
itself, for instance—is a topic that neuroscientists do not often explore.
Defenders of the organic philosophy, Waddington and Needham chief among
them, believed they ended the mechanism-vitalism debate by kicking the old
conflict over on its side. Or, to put it another way, they thought they showed the
irrelevance of the debate as traditionally posed. Vitalism, as Waddington
explained, has always been predicated on the notion that living things required
the postulation of an entity or fluid outside the bounds of physics and chemistry.
As knowledge in both physics and biology developed over the twentieth century,
however, it became more honest to proclaim ignorance: “Since we do not know
in full what entities are demanded by the laws of physics, the distinction
[between mechanism and vitalism] is more or less inapplicable. The question to
ask about biological theory is not whether it is vitalist or mechanist, but whether
it is a useful scientific theory or not. Many useful biological theories cannot yet
be expounded in terms of conventional physics.” 29 It was no longer simply a
duel over the uniqueness of life, vitalists versus the mechanistic perspective that
organisms are wet machines. Life did not require a vital substance, but it was a
denser web of relationships that presented itself to us as a new and distinct class
of things emergent from the physicochemical. Organisms were not exceptional
because they contained extraphysical forces but because the rules at the
organismic level were more expansive than those demanded by ordinary physics
and chemistry. As Smuts and Lloyd Morgan hinted earlier, the rules themselves
had evolved, resulting in a biology safe from encroachment. Physicists interested
in biology such as Max Delbrück, Michael Polanyi, Walter Elsasser, and David
Bohm would repeat this claim in the middle of the century.
What’s more, the organismal perspective asked for a reorientation of the
approach taken by biologists and other scientists in the past: Regard the
experiment as a unique moment, pregnant with the perspective of the observer.
In order to “re-see,” however, a biologist would need to think carefully about his
or her own epistemology even in the laboratory. When given a triangle and an
additional point, “third way” biologists should see a tetrahedron rather than a
trapezoid.
This epistemological stance surely contributed to the challenges that the
organic philosophy faced after the Second World War. It meant biologists would
have to think seriously about epistemology—a dubious proposition. Worse,
stopping to think at all seemed to threaten the speed of biological discovery. It
did not matter to most biologists that, as Whitehead and others seemed to claim,
replicability did not mean objectivity nor, as Woodger pointed out, that biology
was rife with theoretical sloppiness and “poaching.” It did not matter much that
the organic philosophers proclaimed that the exact same event could not happen
in a universe where every moment was new, where organisms were “more nearly
comparable to a river than to a mass of solid rock.” 30 Paradoxically, they
proclaimed that objective knowledge was only gained in the context of an
observer who altered the possibility of objective knowledge. Mechanists
appealed to objectivity in their denunciations of vitalism, but that could only be a
mask, the followers of the organic philosophy claimed. At best, organicists
implied, biologists could offer only rough probabilities—and those were not well
defined, perhaps, not predictive. In other words, when they attempted to answer
the old question, “Knowing only the behavior of nonliving components and their
relationships, would it be possible to predict the occurrence of a whole living
organism from those components?,” organicists answered, “Not by a long shot”;
mechanists tended toward, “Yes, soon.”
For these reasons, among others, a debate continued through the middle of
the twentieth century, after “third way” theorists in the 1930s had seemingly
provided an answer to the mechanist-vitalist controversy. It continued in
basically the same form, in fact: an unresolved dialectic between, for instance,
the thesis of Crick’s gene-centric mechanism and the antithesis of Waddington’s
organicism. Though the names of the opposing sides had changed over the
century, they still represented a perpetual dance of worldviews. Biology
followed a swing of a conceptual or meta-theoretical pendulum, rather than the
replacement of one intellectual paradigm with another.
In making this claim about the non-paradigmatic nature of the history of
biology, I am echoing an argument made by Hilde Hein at the very moment it
was happening, a half century ago. In the late 1960s, while Thomas Kuhn was
busy responding to the criticisms of his paradigm model made by Karl Popper,
among others, Hein published an observation about the field of biology that
should have been incorporated into the broader conversation. Unfortunately, it
slipped by practically unnoticed. There is a pervasive myth, noted Hein, that
science progresses by discarding wrong ideas, that the history of science is a
history of conceptual refuse. A theory persists into the present, it is assumed,
because it has withstood tests. A concept’s contemporaneity is, in this sense, its
justification. 31 Certain issues in the history of science, however, have never been
discarded. In fact, they appear again and again. These, Hein decided, should be
called “meta-theoretical commitments,” since they are as much psychological or
cultural as they are evidentiary. She did not use the term “paradigm”—her
description of meta-theoretical commitments veered closer to what Woodger’s
friend W. V. O. Quine indicated by his term “auxiliary assumptions.” 32 In
biology, Hein argued, the two major meta-theoretical commitments have long
been mechanism and vitalism. 33
While this debate seemed long dead by the time Hein wrote, all of its features
were being recapitulated in the contemporary clashes between what she called
“molecular biology” and “organicism.” These two concepts, according to Hein,
transcend what we usually mean by “theory” in science. They are entire
networks of theories and observations, distinct ways of seeing. Unlike Kuhn’s
stipulation in his notion of paradigms, Hein intimated that meta-theoretical
commitments refuse to be counted among the refuse of science’s past. They are
not, in other words, replaceable the way that Kuhn specified paradigms must be.
They do not disappear.
There is a cynical way of reading the perpetual struggle of meta-theoretical
commitments such as mechanism, vitalism, and the organic philosophy. Not long
after her tousle with Waddington and Joseph Needham, June Goodfield
pronounced that, “The arguments for reductionism and antireductionism seem
irrelevant to what is actually done in the laboratory, mere echoes from the
sidelines whose impact and influence are effectively nil.” 34 As in the case of her
original critique of the organizer experiments years earlier, Goodfield had again
taken her cues when making this statement from none other than Peter Medawar.
She had asked Medawar whether theory mattered even in selecting concepts,
research questions, and so on: “I made the suggestion . . . that the attitude with
which a man approached biology might help him to focus on certain problems
that otherwise might be ignored.” Medawar’s response was “a flat negative . . . it
made no difference at all. A man was a good scientist or he was not.” 35 Of
course Medawar had trouble articulating just what these qualities were that made
a good scientist, a scientist like himself.
Decades earlier, mechanist Jacques Loeb offered up a more reflective answer
to a similar question, an answer opposed to Medawar’s—an answer much closer
to that given by Loeb’s contemporary Pierre Duhem. When Raymond Pearl
questioned Loeb about the influence of these meta-theoretical commitments on
his own scientific work, Loeb answered in the affirmative: “[T]he idea of
proving purely mechanistic explanations of all life phenomena has been the
guiding idea of all of my work. If this idea were taken out, I do not see how I
would have had interest enough to go on working.” 36 And though advocating a
different meta-theoretical commitment, Waddington concluded the same about
his own work: There were practical consequences to a biologist’s philosophical
perspective. In her 1968 article, Hein interviewed embryologist Paul Weiss and
geneticist Gunther Stent. They likewise reported, contra Goodfield and
Medawar, that meta-theory definitely did matter to their scientific work. Perhaps
those biologists who continue to believe like Medawar are simply unaware of
their tacit commitments. Or, like Hofstadter, they believe they should just
proclaim mu and move on, Zen-like. More likely, given the state of biology
education and the tradition of disciplinary balkanization that has grown up over
the twentieth century—the “Two Cultures” problem—they do not think about
these issues at all.
Awareness of the scientists themselves aside, the present book underlines
Hein’s assertion that “the same prejudices do recur in every generation and at
every stage of scientific inquiry.” 37 Mechanism might overturn vitalism to be
itself challenged by organicism, but these are not paradigms replacing one other.
Mechanism returns to be met again by an anti-mechanistic alternative. These are
repeating patterns of thought, rather than paradigms that can be accepted or
rejected. These are webs of concepts glued together by occasionally intense
emotional convictions. Those who hold a particular meta-theoretical
commitment often refuse to see the evidence proffered by the opposing side as
convincing—or even to regard what the opponent brings to the table as
“evidence” at all. Thus anti-mechanists like Walter Elsasser can claim that
“There is no series of actual experiments . . . such that it would be possible to
demonstrate the way in which all the properties of an organism . . . can be
reduced to consequences of molecular structure and dynamics,” while
mechanists like Jim Watson retort “Complete certainty now exists among
essentially all biochemists that the other characteristics of living organisms . . .
will all be completely understood in terms of . . . small and large molecules.” 38
One meta-theoretical commitment may temporarily gain the upper hand in the
work of one scientist and may capture an entire subfield of biology, even for
years; invariably the pendulum swings away from the dominant perspective. The
yin and yang continue to spin.


In those days just after the end of the Great War, idealistic young men like Ian
Suttie and Joseph Woodger looked around the blasted, gas-soaked landscape of
Europe and the Middle East and imagined a better civilization rising up from its
ruins, guided by a more organism-friendly science than Fritz Haber’s and
Jacques Loeb’s. If only someone would stand up, they thought, and, like
Socrates long ago, convict those self-purported to be the intellectual elite of their
own conceptual confusion. 39 Like the Greek original, the twentieth-century
scientific Socrates would have to question earnestly those who claimed to hold
the keys to truth and authority. Like the original, the new Socrates would probe
those answers to be sure they were not just assumptions allowed to stand
because they went unchallenged or because they legitimized existing wealth and
power. The ancient Socrates believed evil had its roots in ignorance and that, if
ignorance was exposed and swept away, goodness could result. The modern
Socrates, they thought, would have to sweep away ignorance and provide a path
out of it by resolving antimonies and by pointing to an alternative, a “third way.”
It was no accident, then, that Woodger’s friends and colleagues, even Karl
Popper, nicknamed Woodger “Socrates.” His axiomatization project, his
promotion of Whitehead and the first generation of organicists, his reason for
drawing the Theoretical Biology Club together—all of it was done with the
Socratic conviction that a science unexamined was not worth trusting.
Some, like Joseph Needham, J. D. Bernal, Dorothy Wrinch, and L. L. Whyte
were convicted. Others, like Waddington, followed Woodger’s trajectory but
ultimately went beyond him, creating epigenetics as a means of attacking the
persistent controversy in theoretical biology regarding the value of an organic
perspective in evolution. Still others, Peter Medawar and Francis Crick, for
instance, could never see the value in wrangling over theory when so many
smaller, more practical problems remained to be solved. History shows that, over
the latter half of the twentieth century anyway, the life sciences community sided
most often with those like Medawar and Crick. Consequently, Woodger’s name
has been more or less forgotten and the organic philosophy left still on the
margins in biology, just as Waddington’s epigenetics—though now touted as a
hot area of research—was once trivialized as warmed-over Lamarckism,
Lysenko with an English accent.
Yet perhaps this historical trajectory—the misunderstandings, the
marginalization, and the reinterpretation—was to be expected. After all, as
Joseph Needham playfully reminded Woodger from time to time, the Athenians
executed Socrates.
 EPILOGUE

IS MODERN EPIGENETICS ORGANIC?

D uring a nationally televised address on August 9, 2001, President George W.

Bush issued a ban on biological research using embryonic stem-cell lines.
Though it would soon be overshadowed by the terror attacks of September 11th,
the ban made international headlines and spurred debates in scientific and
philosophical journals and on university campuses across the United States and
beyond. Suddenly, the news media was tossing about terms like “totipotent,”
“pluripotent,” “somatic,” and “embryonic” to a public largely ignorant of the
science itself but deeply invested in the moral questions that seemed to lurk
behind the science, nevertheless. That was the public introduction to epigenetics
—an approach that promised insight into puzzles genetics alone could not solve,
with potential ramifications for biomedicine. 1
Like many, I first heard of epigenetics during the 2001 debate over stem cells.
It was only much later that I discovered the extent to which the story of
epigenetics and the story of the Cambridge Theoretical Biology Club were
woven together. The Life Organic is an attempt to tell the origins of epigenetics
from the search for a “third way,” and I still feel that the traditional narrative
tools used in the preceding chapters were a good way to fashion this account.
That being said, I have since recognized that there are other valuable tools
available to historians and philosophers of science if what we want to understand
is the development of a field after inception.
In this epilogue, I want to offer a glimpse into a different way to tell the
history of epigenetics as a growing field in its own right, outside the influence of
Waddington or the TBC. By analyzing hundreds of published articles and
measuring the use of certain terms, it is possible to map the contours of the
landscape of work in epigenetics. From what my team and I have collected thus
far, the data begin to address a question that has plagued me since I first
recognized the close connection between the British scientists of the first half of
the twentieth century and epigenetics of our day: When researchers say
“epigenetics” today, do they still mean the same thing? And, if not, is our
epigenetics different than Waddington’s original? In other words, is modern
epigenetics still organic?

A couple of quick notes about what follows. First, it does not have the same
intimate feel as the preceding account. It is not sourced from personal letters
secreted away in archives. Secondly, this work is only in its preliminary stages;
if it feels thin, that is because there is still so much to do. 2

Epigenetics was not new in the new millennium, of course, but likely because of
the attention being devoted to stem cells, research in epigenetics began to take
off around the new millennium. A trickle of articles in the 1990s became a
downpour during the stem-cell debates of 2001, then a torrent after the Human
Genome Project (HGP) completed major operations in 2004. Even within the
much smaller sample size surveyed by me and my team, the massive increase in
interest into epigenetics in the recent past becomes apparent (Figures E.1 and E.2
).

The HGP, especially, challenged some long-held dogmas, surprising

nonspecialists who had learned through their biology courses that all forms of
complexity were due to the number and variety of genes. That assumption came
with another implicit one: Complex organisms such as humans must contain an
astounding number of genes compared to, for instance, dandelions or worms. As
it turns out, the HGP showed that those intuitions were terribly misguided: The
human genome expresses under 20,000 genes, fewer than other seemingly less
complex organisms. 3 Moreover, there appears to be little correlation between
size of organism, number of chromosomes, size of genome, and actual coding
base pairs—traits we led ourselves to believe were obviously connected.
Something other than the number of genes must be necessary to explain organic
complexity, and where genetic explanations fall short, epigenetic ones seem to
be called for.
Epigenetics has persisted since its inception in the 1940s—that much is
uncontested. The meaning of epigenetics over that time span, however, remains
an open question. During the renaissance of epigenetics in the early 2000s,
Nobel laureate Joshua Lederberg acknowledged with some consternation that his
colleagues were imbuing the term “epigenetics” with two different meanings. He
pinned the original version of epigenetics to Waddington by dubbing it
“epigenetics-W”—that version interested in development. 4 However, according
to Lederberg, bioengineer Robin Holliday minted a different definition in the
1990s. “Epigenetics-H[olliday]” is “Nuclear inheritance which is not based on
differences in DNA sequences.” 5
Lederberg hit upon something, but his insight did not penetrate far enough.
Certainly, it appears on the face of it that there are two versions of epigenetics.
Upon closer examination, however, it is unclear just how distinct epigenetics-H
is from ordinary genetics. In other words, the research program of epigenetics-H
(as opposed to popular twenty-first-century accounts of epigenetics) seems not to
have shifted away from that of standard, midcentury genetics.
What distinctions there are largely have to do with the precise locus of
information inside the nucleus. Traditionally, geneticists have claimed that the
new information that impacts an organism’s phenotype is derived from
mutations—mistranslations of the pyrine and pyrimidine rungs of the DNA
ladder so that Cs morph into Ts, for example—or unique recombinations of
existing genetic material. In epigenetics-H, the focus on mutations and
recombinations is supplemented by an examination of other processes known to
occur inside the nucleus of cells. Currently, the four most studied epigenetic-H
mechanisms are (1) DNA CpG methylation, (2) modifications to the process by
which DNA is wrapped around specialized histone proteins in the chromosome,
(3) nucleosome repositioning, and (4) RNA-mediated regulation of noncoding
regions of DNA. Without going into detail, it is enough to note that each of these
processes potentially alters the phenotype without those alterations necessarily
originating from the Cs, Ts, Gs, and As of the DNA molecule itself. Most
epigenetics-H changes remain reversible, at least in principle. Most also regulate
the genome, meaning epigenetic mechanisms sometimes directly impact the
genotype. Effects like these—the ones that mysteriously seem to link childhood
diabetes to starving parents, for instance—are the ones that receive the lion’s
share of attention from science journalists. 6
Nevertheless, while each of these epigenetic mechanisms add new and
exciting twists to the old narrative of genetics, they do not alter the main
characters in that story. Each of these processes may expand what we commonly
mean by “gene”—the bits of inherited stuff that make organisms what we are—
but they do not replace the old reductionistic mechanism that Waddington,
Needham, Woodger, and others fought against. The research program of
epigenetics-H remains focused on the mechanism of molecules shuffling in the
nucleus of a cell, in other words. The sense remains that the complete program
for the wet machine is still situated in that spot. Lederberg identified two
versions of epigenetics. Upon closer examination, however, he was not quite
right: There appears to be epigenetics-W, and then just more molecular biology
governed by the old conventional wisdom of the dominant group.
When he introduced epigenetics in the early 1940s, Waddington meant the
study of the enormous number of inputs that create the phenotype. He had just
invested nearly a decade of his career identifying Spemann’s organizer, and he
envisioned epigenetics as the pursuit of how organismal complexity emerges
from initial simplicity—nearly the same questions raised by the first generation
of organicists at the turn of the twentieth century. It certainly had something to
do with genes. But as he repeatedly insisted over the decades, “genes” could not
be the end as well as the beginning of the story.
Gradually, epigenetics mutated. In 1958, the very year that Francis Crick
published it, geneticist David Nanney showed in “Epigenetic Control Systems”
that Crick’s central dogma did not hold in every case (Waddington, of course,
had demonstrated this even before Crick had proclaimed the dogma).
Information for the creation of proteins and, presumably, the phenotype did
proceed from DNA outward. But that explanation was insufficient, and Nanney
showed why: Cells harboring the same genotype did not always express the
same phenotype, and vice versa. This recognition led to two outcomes. First,
Nanney identified the developmental process later known as the transition from
stem cell totipotency to pluripotency to specificity—the scientific issue behind
the stem-cell debate in 2001. Secondly, Nanney warned his colleagues that they
could not always see clearly that all processes in the nucleus of a cell were
genetic. They uncritically employed faulty epistemology. As a result,
development could not accurately be described as a simple unrolling of a DNA
program in the nucleus; Crick’s central dogma could not be held dogmatically.
Thus, after Nanney, the early course of epigenetics research was simply to
clarify which inputs into a developmental system were genetic and which were
epigenetic, i.e., not coded for in the nucleus itself. By the time Crick pronounced
it, epigenetics-W was agnostic about his central dogma. This agnosticism
immediately became a problem since, according to some molecular biologists,
his central dogma has been at the heart of molecular biology’s “mechanistic
program” long before Crick titled it. 7
The agnosticism of early epigenetics researchers regarding the central dogma
dovetailed nicely with the work being conducted by Boris Ephrussi and Tracy
Sonneborn, among others, into cytoplasmic inheritance. Such research promised
to clarify an ambiguity raised by Nanney. If, for instance, it was possible to
locate components of the cellular structure outside of the nucleus that impacted
inheritance, then it should be easier to determine the degree to which
information from outside the DNA molecule itself was preserved generation to
generation. That research might also be able to answer the extent to which
Lamarckian explanations carried weight. A rigorous debate did take place over
the parsimony of cytoplasmic inheritance midcentury. Ultimately, however, the
conventional wisdom of molecular biologists, the dominant group, dismissed the
results of those who advocated cytoplasmic inheritance. It was far easier to
believe in an executive gene, a master molecule preformed with instructions to
build an organism. 8 As we have witnessed, that mechanistic master molecule
concept impacted the reception of epigenetics-W. More than that, it changed
even the meaning of epigenetics itself.

In 2015, in order to address that question—is modern epigenetics still organic?

—I set out with a small team to interrogate large data sets composed of possible
definitions of epigenetics embedded within epigenetics research publications.
We hoped to model the development of epigenetics in the second half of the
twentieth century.
Our preliminary findings (table E.1 and figure E.3 ) agree to a large extent
with Lederberg’s hunch that many researchers have transitioned away from
epigenetics-W. That transition coincided with the rise of Wilson’s redefined
sociobiology and Hofstadter’s mu in the 1970s. 9 In 1975, Robin Holliday and
John Pugh published “DNA Modification Mechanisms and Gene Activity
During Development,” a groundbreaking article that established that heritable
change did indeed take place outside of the DNA molecule itself. 10 But it also
shifted the locus of epigenetics research away from development. In under a
decade, and certainly by the late 1980s, epigenetics-H—an extension of typical
molecular biology—had decisively, if not entirely, supplanted epigenetics-W.
From now on the research frame of epigenetics research would be located at the
molecular level squarely within the nucleus of the cell.
While more research needs to be conducted on the social and economic
contributors to this transition, the redefinition correlates with two other changes.
The first stage was political and economic: funding shifts in the wake of
President Richard Nixon’s “War on Cancer.” The second was demographic: the
coming of age of a new generation of cancer researchers trained in molecular
methods. According to the recent reflections of one of the most accomplished of
these researchers—Robert A. Weinberg at the MIT Center for Molecular
Oncology—from the start, these Young Turks were programmatically inured
against complex, multicomponent explanations and smugly derisive of the
reasoning offered by prior generations of scientists. They literally refused to take
the complexity of epigenetics-W-type science seriously. 11

There is another, more hopeful result visible through our canvassing of

epigenetics publications over the last seven decades: Epigenetics-W did not
expire, nor was it wholly subsumed into the epigenetics-H borne aloft by waves
of cancer funding. Contemporary scientists including Scott Gilbert, Brian K.
Hall, Mae-Wan Ho, Eva Jablonka, Marion Lamb, and many others continue to
work in areas blazed by Waddington and his comrades long ago. Not only has
their work extended the applicability of epigenetics-W, they have challenged and
refined the conventional wisdom regarding the dominant interpretation of
evolution through new companion subfields such as evo-devo.

The point of this far-too-brief overview of turning points in the history of

epigenetics is not to scrutinize and pass judgment on research traditions per se,
but to reiterate a deeper point regarding biology’s history. Epigenetics in some
form or another is here to stay. Almost weekly news stories touting
“revolutionary” epigenetics discoveries—for instance, that trauma from the
Second World War has been etched biologically into the children of survivors—
only serve to underscore this point. 12 H. J. Muller rejected the inheritance of
acquired characteristics in part out of concern that it signaled that, over time, the
“dominant classes and peoples would have become genetically superior to the
underprivileged ones.” 13 His worries may have been overblown when he voiced
them in the 1930s. But contemporary epigenetic studies make his fears seem no
longer quite so far-fetched.
While it is possible that the present popularity of epigenetics indicates that
the metaphorical pendulum might be swinging away from the gene-centric
mechanism of the later twentieth century, it is by no means certain that
something anti-mechanistic—something organic—will replace it. On the
contrary, within this moment of change and excitement revolving around the
term “epigenetics,” we are witnessing precisely the same meta-theoretical
commitments dancing around each other as were exhibited in the debates
surrounding vitalism, mechanism, and organicism throughout the twentieth
century. Despite the fact that Waddington’s epigenetics was an attempt to
investigate the production of phenotypes from genotypes by refocusing biology’s
priorities on morphology, hierarchies, emergence, evolution, and so on, the
contemporary approach flying under the flag of “epigenetics” reflects old
mechanistic tendencies to reduce organisms to genotypes. According to twenty-
first-century epigenetics, life is still gene-centered and mechanistic; it’s just that
“gene” now means more than it used to. Rather than a new Kuhnian paradigm
replacing an outdated, anomaly-filled one, today’s struggle to define epigenetics
seems to repeat the oscillating, pendulum-like pattern visible through at least the
last century of the life sciences.
 NOTES

INTRODUCTION
1 . McGrath, “Bergson Comes to America,” 601.
2 . “Bergson Fills Hall at First Lecture,” The Sun (New York), 4 Feb. 1913, 7,
http://chroniclingamerica.loc.gov/lccn/sn83030272/1913–02–04/ed-1/seq-7/ . See also “French Philosopher
Not a Good Sailor,” New York Tribune , 3 Feb. 1915, 9,
http://chroniclingamerica.loc.gov/lccn/sn83030214/1913–02–03/ed-1/seq-9/ ; “Bergson’s Second Lecture,”
The Sun (New York), 5 Feb. 1913, 2, http://chroniclingamerica.loc.gov/lccn/sn83030272/1913–02–05/ed-
1/seq-2 ; and “Believes in Free Will,” New York Tribune , 5 Feb. 1913, 5,
http://chroniclingamerica.loc.gov/lccn/sn83030214/1913–02–05/ed-1/seq-5 .
3 . Though McGrath (“Bergson Comes to America,” 599, n. 3) disputes the authenticity of the traffic
jam on Broadway story, he admits it was “widely repeated.” The story is at least plausible, given eyewitness
details and the geographical location of Havemeyer Hall.
4 . Mayr, Toward a New Philosophy of Biology , 13; see also Steffes, “Panpsychic Organicism.”
5 . Ritter, Unity of the Organism , 1, 24 (italics in the original).
6 . Beckner, “Organismic Biology,” 55.
7 . As Randall Collins notes in Sociology of Philosophies (6), definition by rejection of other positions
(i.e., what we stand against ) is at least as common as a positive definition (i.e., what we stand for ).
8 . Butterfield, Origins , 9.
9 . Munson, general introduction to Man and Nature , xx.
10 . To be sure, a handful of works have addressed bits and pieces of this story. One of the most
important recent accounts of the early history is Esposito, Romantic Biology . Esposito’s account drops off
after World War II; thus, he only hints at a primary connection between organicism and epigenetics. A
critical addition to this account from the philosophy of science is Nicholson and Gawne, “Neither Logical
Empiricism nor Vitalism.”
11 . See, for example, Francis, Epigenetics . One of these exciting accounts came in the form of a radio
broadcast (now podcast): Radiolab, “Inheritance,” season 11, episode 2,
http://www.radiolab.org/story/251876-inheritance/ .
12 . The “central dogma,” coined in 1958, was Francis Crick’s nickname for the notion that information
flows from DNA to RNA to protein and, assumedly, to organism—and never in reverse. It was another way
of codifying the nineteenth-century belief that germ creates soma or, using twentieth century terminology,
genotype creates phenotype . Corollary to the central dogma is the dictum of geneticist H. J. Muller: “Every
gene from a pre-existing gene” (H. J. Muller, “Bar Duplication,” Science 83, no. 2161 [1936]: 530). Even
with the notion of “genes-for” traits and the language of genetic coding introduced during the late 1950s,
the actual process of development from genotype to phenotype remained an unopened black-box problem
into the twenty-first century.
13 . These claims appear in a number of otherwise indispensable sources, the most important of which
are Harrington, Reenchanted Science , and Gilbert and Sarkar, “Embracing Complexity.”
14 . Ironically, commentators offer epigenetics as a key example of one such paradigm shift in biology.
 15 . Philosopher Hilde Hein made a similar observation five decades ago, though it seems not to have
been followed up in the literature. While I dispute Hein’s categorization of organicism as synonymous with
vitalism, my historical investigation agrees with her analysis that “the controversy between organicism and
molecular biology is not an isolated historical dispute; but is a manifestation of a perennial disagreement
between two types of meta-theoretical commitment” (“Molecular Biology vs. Organicism,” 238). To be
sure, little that I am arguing in this final point is remarkable to philosophers of science, who moved on from
Popper and Kuhn some time ago (and rarely discussed Huxley). Lakatos, Laudan, and Feyerabend attracted
interest for a time, though their philosophical models are now dated as well. Some of the most exciting
work on concept change, especially in the life sciences, can now be found among the works of John Dupré
(e.g., Disorder of Things ) and Kevin C. Elliott and Daniel J. McKaughan (e.g., “Values in Scientific
Discovery”).
16 . Northbourne, Look to the Land ; and Paull, “Lord Northbourne.”
17 . The individuals of this collective feature prominently in The Life Organic , as they do in the early
work of historian of science Pnina Abir-Am. See, for instance, Abir-Am’s “Discourse of Physical Power
and Biological Knowledge in the 1930s”; “The ‘Biotheoretical Gathering’ in England and the Origins of
Molecular Biology (1932–38)”; “Recasting the Disciplinary Order in Science”; and “The Biotheoretical
Gathering, Trans-Disciplinary Authority and the Incipient Legitimation of Molecular Biology in the 1930s.”
Abir-Am used the term “Biotheoretical Gathering”—instead of Theoretical Biology Club—a name that
appears in one piece of correspondence written about the time of the foundation of the group. Her goal in
these publications was to use the Theoretical Biology Club/Biotheoretical Gathering as a lens through
which to draft a novel sociology of science. In this, her work continues to be very valuable. I am more
interested in the issue that motivated the members of the group: how to build the legacy of the organic
philosophy into an interdisciplinary scientific research program.
18 . The biology of Soviet agronomist Trofim Denisovich Lysenko (1898–1976), and the Lysenkoist
“school” that followed him, has been the subject of extensive debate in recent years. Lysenkoists opposed
the “hard” genetics espoused by Mendelians in the UK and United States in favor of something more open
to environmental influences. The story of Lysenko and Soviet biology in the 1930s through ’60s has
become a morality tale for scientists and funding agencies, a warning about the ill effects of government
interference in scientific research. See Roll-Hansen, “On the Philosophical Roots,” for a recent retelling of
this morality tale. The construction of this morality tale by geneticists, and the rhetorical force of this tale in
dampening support for epigenetics, is the subject of chapter 11 .
19 . This account fits more naturally in Hayden White’s category of tragedy rather than comedy or
romance, in other words.
20 . Gilbert and Sarkar, “Embracing Complexity.”
21 . See, for example, Pigliucci, “Do We Need an Extended Evolutionary Synthesis?” and Burggren
and Crews, “Epigenetics in Comparative Biology.” This list encompasses Scott F. Gilbert, Brian Goodwin,
Brian K. Hall, Gerd Müller, Susan Oyama, James A. Shapiro, and Mary-Jane West-Eberhard, to name only
a few. One of the most important recent essays reinforcing my claim here is Nicholson’s “Return of the
Organism.”

1. THE FIRST GENERATION OF ORGANICISTS

1 . Stephens, Hermit Philosopher of Liendo .
2 . Keeton, “Montgomery—Pioneer of Organicism,” 309, 2n. St. Thomas Hospital is the major research
hospital across the Thames from Parliament.
3 . Salter, “A New Type of Naturalism,” 91.
4 . Dielmann, “Elisabet Ney.”
5 . Owen quoted in Salter, “New Type of Naturalism,” 92.
 6 . “Review of ‘On the Formation of So-Called Cells in Animal Bodies’ by E. D. Montgomery,” 198.
7 . Keeton, Philosophy of Edmund Montgomery , 8.
8 . Ruse, Darwinian Revolution , 212–14.
9 . Darwin, Variation of Animals and Plants , 366. See Keeton, Philosophy of Edmund Montgomery ,
for a discussion of Darwin and his comrades on the issue of cell theory.
10 . Dielmann, “Elisabet Ney.”
11 . Montgomery, “Unity of the Organic Individual [1],” 326.
12 . Montgomery, “Are We ‘Cell Aggregates’?,” 106.
13 . Keeton, “Montgomery—Pioneer of Organicism,” 334–35.
14 . Montgomery, Philosophical Problems in Light of Vital Organization (London: G. P. Putnam’s Sons,
1907); Loeb, The Mechanistic Conception of Life (Chicago: University of Chicago Press, 1912).
15 . The first generation of organicists saw Claude Bernard’s milieu intérieur concept featured in his
Introduction à l’Etude de la Médecine Expérimentale (1865) as inspiration for their own organicism. See,
for instance, J. S. Haldane, Organism and Environment , 3, 1n.
16 . Esposito (Romantic Biology ) offers more extensive treatment of this first generation.
17 . As the era is characterized in Shrimpton, “‘Lane, You’re a Perfect Pessimist’”; and Payne, History
of Fascism , 23–24.
18 . Thus my account offers an important amendment to the excellent one by Esposito, Romantic
Biology .
19 . Dexter, “C. O. Whitman,” 1251–53; and Isaacs et al., “Hirohito.”
20 . Lillie, “Charles Otis Whitman”; and Mayr, “Whitman.”
21 . Maienschein, “Whitman at Chicago.”
22 . Whitman, “Inadequacy of the Cell-Theory,” 641.
23 . Whitman, “Inadequacy of the Cell-Theory,” 645–46 (italics in the original).
24 . Lloyd Morgan, “Autobiography,” 240.
25 . Richards, Darwin and the Emergence , 377; Lloyd Morgan, The Springs of Conduct: An Essay in
Evolution (London: Edward Arnold, 1884).
26 . Romanes and Lloyd Morgan, Post-Darwinian Questions .
27 . Lloyd Morgan, “The Doctrine of Auta”; and Lloyd Morgan “Three Aspects of Monism.”
28 . Lloyd Morgan, “Three Aspects of Monism.”
29 . Lloyd Morgan, Introduction to Comparative Psychology (London: W. Scott, 1894).
30 . With evolutionary incongruities in mind, Lloyd Morgan proposed what came to be called
“Morgan’s canon”: “In no case may we interpret an action as the outcome of the exercise of a higher
psychical faculty, if it can be interpreted as the outcome of one which stands lower in the psychological
scale” (An Introduction to Comparative Psychology , 2nd ed. [London: W. Scott, Ltd., 1903], 53). The
behaviorist school latched on to Morgan’s canon and repeated it ad nauseum. Yet they overlooked—if they
read his original works at all—the larger “third way” latticework that Lloyd Morgan labored to weave
underneath his psychology. Behaviorists misinterpreted the canon as just another phrasing of mechanistic
materialism: One should not attribute to something like “mind” what should be understood merely as
“body.” Since organisms were ultimately just “body,” any evidence of higher consciousness on the part of
nonhuman animals should be regarded as mere instinct, itself epiphenomenal of the meat of the organism.
As B. F. Skinner pronounced in 1938, “Darwin, insisting upon the continuity of mind, attributed mental
faculties to subhuman species. Lloyd Morgan, with his law of parsimony, dispensed with them” (The
Behavior of Organisms: An Experimental Analysis [New York: Appleton-Century, 1938], 4). If we actually
read Lloyd Morgan’s own words, however, we find that he meant no such thing. In fact, Skinner’s
interpretation demonstrates willful ignorance, as Lloyd Morgan took pains over the course of several books,
articles, and lecture series throughout the 1910s and ’20s to explain exactly what he did mean.
31 . Lloyd Morgan likely developed emergentism in light of his study of the works of George Henry
Lewes—especially Problems of Life and Mind (1875–79)—and William Kingdon Clifford, who introduced
three-aspect monism in “On the Nature of Things-in-Themselves” (Mind 3, no. 9 (1878): 57–67) and “On
Theories of the Physical Forces (1870),” Lectures and Essays (London: Macmillan, 1886), 74–84.
32 . Lloyd Morgan, Emergent Evolution , 2.
33 . Lloyd Morgan, Emergent Evolution , 7–8.
34 . “New Technical Weapons,” Times (London), 29 Apr. 1915, 9. Note the “again”—even the British
press recognized the use of chemical weapons by both German and French forces prior to the First Battle of
Ypres, though not on this scale.
35 . J. S. Haldane to Louisa Kathleen Trotter, 16 Oct. 1890, quoted in Sturdy, “Co-ordinated Whole,”
246–47.
36 . Douglas, “John Scott Haldane,” 118; and Goodman, Suffer and Survive , 2–4.
37 . Thomas F. Gieryn (“Boundary-work”) suggests that the “pure” and “applied” distinction became
especially important in Britain for professional and rhetorical purposes during this era.
38 . J. S. Haldane, “Vitalism,” 412.
39 . J. S. Haldane, “Vitalism,” 408.
40 . J. S. Haldane, “Vitalism,” 410.
41 . J. S. Haldane, Mechanism, Life, and Personality: An Examination of the Mechanistic Theory of Life
and Mind (London: John Murray, 1913); and Organism and Environment as Illustrated by the Physiology of
Breathing (New Haven: Yale University Press, 1917).
42 . J. S. Haldane, Organism and Environment , 3, 1n.
43 . J. S. Haldane, Organism and Environment , 17.
44 . J. S. Haldane quoted in Needham, Time , 204.
45 . J. S. Haldane, Respiration (New Haven: Yale University Press, 1922).
46 . Bergson, Meaning of the War .
47 . Edman, “Eighteenth Annual Meeting,” 127.
48 . Edman, “Eighteenth Annual Meeting,” 128–29.
49 . Holt et al., New Realism .
50 . Edman, “Eighteenth Annual Meeting,” 129 (italics in the original).
51 . Mayer, “L. J. Henderson”; and Cross and Albury, “Walter B. Cannon, L. J. Henderson, and the
Organic Analogy.”
52 . Henderson, Fitness of the Environment , 306. Joseph Needham claimed that he “had been brought
up on L. J. Henderson’s ‘Fitness of the Environment.’” Needham continued to find support for his belief in
“non-reductionist biological organicism” in Henderson’s work—seventy years after Fitness of the
Environment was first published. Needham, foreword to Beyond Neo-Darwinism , vii–viii.
53 . See correspondence between Loeb and Henderson, Box 6, JL-LC.
54 . “The properties of matter . . . are now seen to be intimately related to the structure of the living
being and to its activities. . . . For the whole evolutionary process, both cosmic and organic, is one, and the
biologist may now be rightly regard the universe in its very essence as biocentric” (Henderson, Fitness of
the Environment , 312); see also Fry, “On the Biological Significance of the Properties of Matter.”
55 . Henderson, The Order of Nature: An Essay (Cambridge: Harvard University Press, 1917).
56 . Haraway, Crystals, Fabrics, and Fields , 35–36.
57 . Ritter, Unity of the Organism , 1, 27 (italics in the original).
 58 . Russell seems to have borrowed this dichotomous way of setting up the problem from Jacques
Loeb’s address “The Mechanistic Conception of Life,” which Loeb delivered in 1911 to Ernst Haeckel’s
International Congress of Monism.
59 . E. S. Russell, “Vitalism,” 329.
60 . E. S. Russell, “Vitalism,” 337.
61 . E. S. Russell, “Vitalism,” 343.
62 . “Dr. E. S. Russell,” 8.
63 . E. S. Russell, Study of Living Things , vii.
64 . E. S. Russell, Study of Living Things , 41.
65 . E. S. Russell, Study of Living Things , 60–61.
66 . E. S. Russell, Study of Living Things , 61.
67 . E. S. Russell, Study of Living Things , 80.
68 . E. S. Russell, Form and Function , vi.
69 . E. S. Russell proclaimed in the 1950s, “If we look at living things quite simply and objectively we
cannot but be struck by one feature of their activities, which seems to mark them off from anything
inorganic. . . . [T]here is common to all living things this basic element of directive striving, usually
unconscious and blind, only rarely emerging into consciousness to become intelligently purposive.” (“The
‘Drive’ Element in Life,” 108).
70 . J. S. Haldane, Philosophical Basis of Biology , 140–52.
71 . Montgomery, “‘Cell-Aggregates’?,” 100–101.

2. NEEDHAM’S REVIVAL OF MECHANISM

1 . Rignano, Man Not a Machine , 10.
2 . Needham, Man a Machine , 38.
3 . Rignano, Man Not a Machine , 24.
4 . Needham, Man a Machine , 62.
5 . Holorenshaw, “Making of an Honorary Taoist,” 2 and 20.
6 . Gurdon and Rodbard, “Biographical Memoir,” 9–14. Fellows of the Royal Society are honored for
their distinguished work in science, mathematics, engineering, medicine, or technology. Fellows of the
British Academy are so honored for their work in the humanities and social sciences. A “Companion of
Honour” is a rare title bestowed upon an individual for outstanding lifetime achievements and is limited to
sixty living individuals at any one time.
7 . Werskey, “Understanding Needham,” 16.
8 . Gurdon and Rodbard, “Biographical Memoir,” 9.
9 . Holorenshaw, “Making of an Honorary Taoist,” 7; and Needham, Chemical Embryology .
10 . Goodfellow, “The Church Socialist League.”
11 . Rattlejag Morris, “The History of Morris,” accessed 12 Aug. 2014,
http://www.rattlejagmorris.org.uk/history-of-morris-dancing .
12 . Needham, “Mechanism,” 220–57; Needham, Sceptical Biologist ; and Needham, The Great
Amphibium .
13 . Needham, “Recent Developments.”
14 . Hartridge and Roughton, “Improvements in the Apparatus.”
 15 . F. J. W. Roughton, “The Use of Thermodynamics in Biology,” paper delivered to the Cambridge
Biochemical Club (May 1927), cited in Needham, “Recent Developments,” 85.
16 . Needham, “Recent Developments,” 79.
17 . C. Lloyd Morgan to Joseph Needham, 31 Oct. 1925, M.62, Joseph Needham papers, Cambridge
University Library (hereafter JN-C).
18 . Needham, “Recent Developments,” 89.
19 . Needham, Time , 10.
20 . Gould, “Nonoverlapping Magisteria.”
21 . Needham, Time , 9.

3. SOCRATES AND THE PRINCIPIA BIOLOGÆ

1 . Baker, “Norfolk Regiment.”
2 . War Office, Statistics of the Military Effort .
3 . Woodger, “Notes on a Cestode.”
4 . Gatenby and Woodger, “On the Relationship Between the Formation of Yolk.”
5 . Heard, introduction to Origins of Love and Hate , xx–xxi.
6 . See especially their April 1925 correspondence, C1/3/Suttie, Woodger papers, University College
London, London, England, UK (hereafter JHW-L).
7 . Suttie to Woodger, 7 Jan. 1922, C1/3/Suttie, JHW-L.
8 . Suttie to Woodger, 16 Mar. 1924, C1/3/Suttie, JHW-L.
9 . “Obituary: Dr. Ian D. Suttie,” 880; Suttie, On the Origins of Love and Hate (London: Kegan Paul,
Trench Trübner, 1935).
10 . Suttie to Woodger, 7 Nov. 1919, C1/3/Suttie, JHW-L.
11 . Suttie to Woodger, 7 Nov. 1919, C1/3/Suttie, JHW-L (italics in the original).
12 . Woodger, Elementary Morphology and Physiology , 487–88.
13 . Coen, “Living Precisely.”
14 . Woodger to UCL Registrar, 6 Jan. 1930, C1/3/Misc. correspondence, JHW-L.
15 . Nicholson and Gawne, “Rethinking Woodger’s Legacy.”
16 . Woodger, “Explanation in Biology”; and Woodger, “Science and Metaphysics in Biology.”
17 . Woodger to J. Needham, 28 May 1929, M.97, JN-C.
18 . Woodger to J. Needham, 28 May 1929, M.97, JN-C.
19 . Hentschel, “Review of Biological Principles .”
20 . Woodger, Biological Principles , 4.
21 . At Albert Einstein’s “Akademie Olympia,” participants read both books and found them
immensely influential on their scientific work. Hans-Josef Küpper, “The Bernese ‘Akademie Olympia,’”
accessed 13 Oct. 2013, http://www.einstein-website.de/z_biography/olympia-e.html .
22 . Woodger, Biological Principles , 229–30. He felt uncomfortable with “methodological vitalist” in
part because Woodger could not find any strong positive claims that were shared across the group. Each
anti-mechanist had different, occasionally contradictory, reasons for opposing mechanism. Ernst Mach,
Analyse der Empfindungen (Analysis of Sensations , English translation, 1914); Karl Pearson, Grammar of
Science (3rd ed., 1911).
23 . Wilson, Cell in Development , 1037.
 24 . Wilson, Cell in Development , 1117.
25 . See, for instance, the misplaced critique of Roll-Hansen, “E. S. Russell and J. H. Woodger.”
26 . Woodger, Biological Principles , 266.
27 . Woodger, Biological Principles , 291.
28 . Woodger, Biological Principles , 294.
29 . Woodger, Biological Principles , 450.
30 . Woodger, Biological Principles , 416. This line of reasoning about progress in evolution continues
to find supporters, according to McShea, “Complexity and Evolution.”
31 . Note that this is exactly the direction taken by researchers in the modern field called “evo-devo.”
32 . Woodger, Biological Principles , 480.
33 . Woodger, Biological Principles , 486.
34 . J. Needham to Woodger, 26 Feb. 1929, M.97, JN-C.
35 . Ludwig von Bertalanffy to J. Needham, 7 Oct. 1929, M.5, JN-C.
36 . J. Needham to Woodger, 12 Oct. 1929, M.97, JN-C.
37 . J. Needham to Woodger, 26 Nov. 1929; and return, 29 Nov. 1929, M.97, JN-C.
38 . Woodger to J. Needham, 25 Jan. 1930, M.97, JN-C.
39 . J. Needham to Woodger, 1 Feb. 30, M.97, JN-C.
40 . At the time, Mind ’s editor was the Cambridge philosopher G. E. Moore, one of the three
Cambridge philosophical “triumvirate” with Bertrand Russell and Ludwig Wittgenstein and someone
Woodger certainly hoped to impress.
41 . Needham, “Review of Biological Principles .”
42 . Woodger to J. Needham, 2 Jun. 1930, M.97, JN-C.
43 . Woodger to J. Needham, 1 Aug. 1930, M.97, JN-C.
44 . Woodger, “Biological Principles ; Letter to the Editor.”
45 . G. E. Moore to J. Needham, 16 June 1930, M.97, JN-C.
46 . J. Needham to Woodger, 31 July 1930, M.97, JN-C.
47 . Woodger to J. Needham, 2 Oct. 1930, M.97, JN-C.
48 . Pearl to Woodger, 1 Aug. 1929, C1/2/P, JHW-L (italics in the original).
49 . Pearl to Woodger, 7 Sep. 1929. C1/2/P, JHW-L.
50 . He wrote Harding that he wanted “to improve upon Chap. IX [ . . . ] which I felt was very
muddled.” Woodger to Harding, 1st Apr. 1930, C1/2/H, JHW-L.
51 . Woodger, “‘Concept of Organism,’ Part I,” 13.
52 . Woodger, “‘Concept of Organism,’ Part II,” 450.
53 . Woodger to J. Needham, 2 Apr. 30, M.97, JN-C.
54 . Callebaut, “Again, What the Philosophy of Biology Is Not,” is only the tip of the iceberg when it
comes to critical treatments of Woodger. Nicholson and Gawne, “Rethinking Woodger’s Legacy,”
challenges this narrative.
55 . Woodger, “‘Concept of Organism,’ Part III,” 184.
56 . Woodger, “‘Concept of Organism,’ Part I,” 14.
57 . For example, Roll-Hansen, “E. S. Russell and J. H. Woodger.”
58 . Woodger, “‘Concept of Organism,’ Part I,” 18. Ron Amundson has leveled a similar, more
extensive critique in Changing Role of the Embryo . Woodger’s accusation here is his own version of what
came to be called “Lillie’s paradox,” first enunciated in 1927. See Burian, “Lillie’s Paradox.”
59 . Woodger, “‘Concept of Organism,’ Part I,” 19.

4. THE TIPPING POINT

1 . International Congress of the History of Science and Technology, Science at the Cross Roads .
2 . Cohen, Bukharin and the Bolshevik Revolution , 350–53.
3 . Goldsmith, “Joseph Needham, Honorary Taoist,” 1–3.
4 . Hodgkin, “‘Microcosm,’” 14.
5 . Hodgkin, “‘Microcosm,’” 14.
6 . Unpublished Cofman circular, “Second International Congress on the History of Science, 1931,”
E.110, JN-C.
7 . Unpublished Cofman circular, “Second International Congress on the History of Science, 1931,”
E.110, JN-C.
8 . McDougall, “Philosophy of J. S. Haldane,” 420.
9 . Colville, Fringes of Power , 269–71; Crafford, Jan Smuts , 197–99; Dubow, “Smuts,” 45–74; and
Heyns, “Preamble of the United Nations Charter,” 337.
10 . Smuts, Holism and Evolution , 3–4.
11 . Darwin, Origin of Species , 84.
12 . Smuts, Holism and Evolution , 12.
13 . Laplace, Philosophical Essay , 2–3.
14 . Smuts, Holism and Evolution , 8.
15 . Smuts, “Presidential Address,” 3.
16 . Thomson, “Section A, Presidential Address,” 29.
17 . Smuts, “Presidential Address,” 10.
18 . Smuts, “Presidential Address,” 10–11.
19 . Smuts, “Presidential Address,” 11 (italics in the original).
20 . Crafford, Jan Smuts , 141.

5. WADDINGTON AND THE ORGANIZER

1 . Holorenshaw, “Making of an Honorary Taoist,” 7–8.
2 . Spemann, “Über Correlationen in der Entwicklung des Auges.”
3 . Gilbert, “Selective History of Induction, II.”
4 . Spemann, “Experimentelle Forschungen zum Determinations- und Individualitatsproblem.”
5 . Spemann, “Über die Determination der ersten Organanlagen.”
6 . Hamburger, “Hilde Mangold,” 4–8.
7 . Fässler and Sander, “Hilde Mangold,” 325.
8 . Spemann, “Entwicklungsphysiologische Studien am Tritonei, III.” At the time, the genus name was
Triton .
9 . Dana Campbell and Gonçalo Themudo, eds., “Triturus cristatus (Laurenti, 1768),” Encyclopedia of
Life, http://eol.org/pages/1018157 .
10 . Hans Spemann, “The Organizer-Effect in Embryonic Development,” Nobel Lecture (1935),
Nobelprize.org , Nobel Media AB 2014,
http://www.nobelprize.org/nobel_prizes/medicine/laureates/1935/spemann-lecture.html .
11 . Hamburger, “Hilde Mangold,” 10.
12 . Fässler and Sander, “Hilde Mangold,” 327, table 1.
13 . Spemann, “Organizer-Effect.”
14 . Hamburger, “Hilde Mangold,” 11.
15 . Fässler and Sander, “Hilde Mangold,” 326–27.
16 . Spemann and H. [Pröscholdt] Mangold, “Induction of Embryonic Primordia.”
17 . DeRobertis, “Spemann’s Organizer.”
18 . Spemann and H. [Pröscholdt] Mangold, “Induction of Embryonic Primordia,” 38.
19 . Spemann and H. [Pröscholdt] Mangold, “Induction of Embryonic Primordia,” 38.
20 . Robertson, “Conrad Hal Waddington,” 575–77.
21 . See Wad reminiscences by friend Gregory Bateson, MS 98, Box 36, Folder 1447, GB-SC.
22 . Waddington, “Practical Consequences of Metaphysical Beliefs,” 73–74.
23 . Gregory Bateson to Albert Robertson, 30 Apr. 1976, MS 98, Box 36, Folder 1447, GB-SC.
Robertson would publish these remarks in: Robertson, “Conrad Hal Waddington,” 578. Yoxen (“Form and
Strategy in Biology,” 312) envisions Bateson and Waddington together as “two highly intelligent, playful,
rather sophisticated young men, casting about for projects that would satisfy their aesthetic and
philosophical sensibilities.” Peterson (“Finding Mind, Form, Organism, and Person”) details the
relationship between Bateson and Waddington and their specific influences on each other.
24 . Cock and Forsdyke, Treasure Your Exceptions , 207–10 and 640; and Richmond, “Women in the
Early History of Genetics.”
25 . Waddington, “Pollen Germination in Stocks.” With his publication, Waddington joined a relatively
exclusive club in British genetics. That year, Journal of Genetics published studies by R. A. Fisher, C. D.
Darlington, Sydney Cross Harland’s groundbreaking cotton genetics paper, R. C. Punnett, Julian Huxley,
and E. B. Ford. Edith Saunders had pioneered work in this area (“Further Studies on Inheritance”). As
Waddington and Saunders noted, M. incana does not seem to conform to typical Mendelian ratios. A patient
Danish plant breeder, Øjvind Winge (“Inheritance of Double Flowers”), worked out the solution to this
apparent problem two years later.
26 . Waddington, “Notes on Graphical Methods.”
27 . Waddington, “Philosophy and Biology; essay submitted for the Arnold Gerstenberg Prize, 1929,”
(unpublished), 31, MS 3024.2, C. H. Waddington papers, University of Edinburgh Library, Edinburgh,
Scotland, UK (hereafter CHW-E).
28 . Waddington, “Philosophy and Biology (unpublished),” 38–40.
29 . Waddington, “Philosophy and Biology (unpublished),” 66.
30 . Note that Waddington wrote these words the year before R. A. Fisher published his landmark
Genetical Theory of Natural Selection (Oxford: The Clarendon Press, 1930).
31 . “The Strangeways Laboratory,” 539.
32 . Robertson, “Conrad Hal Waddington,” 578.
33 . Waddington, “Developmental Mechanics of Chicken and Duck Embryos.”
34 . Haldane and Waddington, “Inbreeding and Linkage.” Waddington recalled this experience as a
“classical neo-Darwinist” without relish. After the “strong nourishing soup” of work with whole organisms,
this tasted like “thin gruel of mathematical formalism.” Waddington, Evolution of an Evolutionist , 8. On
occasion of Haldane’s death, Waddington wrote to Joseph Needham, “I never knew him well, and I don’t
actually think very much of all the ‘mathematical theory of evolution’ stuff which he started—it is all
ultimately based on the quite falacious notion that selection coefficients belong to genes, whereas actually
they belong to phenotypes, a matter which has profound consequences.” Waddington to J. Needham, 25
Mar. 1965, M.18, JN-C.
35 . Oppenheim, “Viktor Hamburger.”
36 . Yoxen, “Form and Strategy,” 315.

6. THE ORIGINAL THEORETICAL BIOLOGY CLUB

1 . Woodger to J. Needham, 30 Apr. 1932, M.98, JN-C. Robson had recently completed his landmark
book The Species Problem (London: Oliver & Boyd, 1928).
2 . J. Needham to Woodger, 8 May 1931, M.98, JN-C. Lana Whyte helped design the jet airplane and,
in the process, formulated his own unified theory of physics and biology, The Unitary Principle in Physics
and Biology . He cited Needham, Waddington, and Woodger as among his “main teachers in biology.”
Whyte, Unitary Principle , 3. His claim about discussing unified field theory with Einstein appears in his
Focus and Diversions .
3 . Senechal’s I Died for Beauty is an indispensable, beautifully written biography of Wrinch.
4 . Wrinch to Woodger, 3 Nov. 1930, C1/3/Misc. correspondence, JHW-L.
5 . Wrinch to J. Needham, 10 Sep. 1932, J.243, “1932 notes,” JN-C.
6 . B. Russell, Introduction to Mathematical Philosophy , 136.
7 . Theoretical Biology Club meeting notes appear primarily in E.111 and J.243, JN-C.
8 . Olby, “Structural and Dynamical Explanations,” 281; D’Arcy W. Thompson, On Growth and Form
(Cambridge University Press, 1917).
9 . These discussions were the inspiration behind a major theme that was to be developed in
Waddington’s epigenetics.
10 . Davis, Dietrich, and Jacobs, “Homeotic Mutants,” 139–40.
11 . Abir-Am (“The ‘Biotheoretical Gathering’ in England and the Origins of Molecular Biology,” 178)
tracks TBC attendance up to 1938.
12 . Woodger to J. Needham, 4 Dec. 1932, J. 243, “1932 notes,” JN-C.
13 . Weaver’s diary, 15 June 1936, 401D RG 1.1, Folder 498, Box 38, RF-NY.
14 . See the MI5’s extensive file on Joseph Needham through the mid-1950s: KV2/3055, NA-K.
15 . Olby, Path to the Double Helix , 258. Olby’s suspicions have been borne out through recently
declassified MI5 files: KV 2/1811, NA-K.
16 . Waddington, “Induction by Coagulated Organisers,” 275–76.
17 . Note that Davis et al. (“Homeotic Mutants,” 139) faults Spemann for believing the organizer to be
“an irreducible, holistic phenomenon.” The Sept. 1933 TBC notes (E.111, JN-C) suggest Needham,
Waddington, and others found fault not with Spemann’s holism but his preference for vitalistic explanations.
18 . Bertalanffy and Woodger, Modern Theories of Development .
19 . Wrinch, “Chromosome Behaviour.”
20 . Wrinch, “Molecular Structure of Chromosomes.”
21 . Olby, Double Helix , 116.
22 . Abir-Am, “Biotheoretical Gathering, Trans-Disciplinary Authority,” 2.
23 . Hacohen, Karl Popper , 310.
 24 . Wrinch to Woodger, 23 May 1935, M.99, JN-C.
25 . Popper quoted by Hacohen, Karl Popper , 315. See “1936 Whitsun[day]” meeting notes, J.244, JN-
C.
26 . Hacohen (Karl Popper , 481) suggests that Popper’s ire in Poverty of Historicism was directed at
Waddington’s Science and Ethics , which Popper seems to have misinterpreted as advocating Soviet-style
state planning in science, as much as any conversation in 1936.
27 . Needham, Order and Life , 17, 20n.
28 . Needham, Sceptical Biologist , 85.
29 . Needham, Order and Life , 17, 20n.
30 . Zavadovsky, “‘Physical’ and ‘Biological.’”
31 . Marcel Prenant, “La conception matérialiste dialectique en Biologie,” Bulletin Societe
Philomathematique 116 (1933): 84, quoted in Needham, Order and Life , 46.
32 . Needham, Order and Life , 45.

7. LARGE PLANS VERSUS THE ULTIMATE LITTLENESS OF

THINGS
1 . J. Needham, “An account of the negotiations regarding financial aid for technical assistance, etc., for
studies on the borderline between Biochemistry and Embryology” (unpublished memo), 12 July 1938,
E.114, JN-C.
2 . According to Kay’s investigation of the Rockefeller Foundation’s 1938 Annual Report (Kay,
Molecular Vision , 43 and 188–89), Weaver coined the term “molecular biology” and began explicitly
channeling funding in this direction around 1938; see also Kohler, “Management of Science.”
3 . J. Needham to W. E. Tisdale of the Rockefeller Foundation, 26 July 1934, E.114, JN-C.
4 . Abir-Am, “Discourse of Physical Power,” 362.
5 . Abir-Am, “Discourse of Physical Power,” 342.
6 . Abir-Am, “Discourse of Physical Power,” 363.
7 . Abir-Am, “Biotheoretical Gathering, Trans-Disciplinary Authority,” 23.
8 . Sir Henry Dale, head of the Royal Society, was a contact of Weaver’s and knew that British
intelligence closely monitored the Needhams and Bernal, including recording telephone conversations and
intercepting mail. See British intelligence files on Bernal (KV2/1181) and the Needhams (KV2/3055), NA-
K.
9 . “Memorandum” (no author, no date [probably written by J. Needham in late 1935, given internal
dating and the fact that associated scraps are written on notepads bearing dates in October 1935]), E.114,
JN-C.
10 . “Memorandum on the position of C. H. Waddington,” enclosure with letter from Needham to
Tisdale, 20 July 1936, B.25, JN-C.
11 . Tisdale to J. Needham, 9 July 1936, B.25, JN-C.
12 . It was well known in Oxbridge scientific circles that “James Gray has a curious hostility to
Experimental Embryology & related subjects.” That curious hostility translated specifically into shifting
grant money away from Waddington even before this Rockefeller Foundation funding incident. J. Huxley to
J. Needham, 12 June 1934, M.34, JN-C.
13 . J. Needham to Tisdale (undated [I date it contextually to December 1936]), B.26, JN-C.
14 . “Memorandum on Applications for Grants” (approved by the Cambridge University Council on 1
Mar. 1937), B.26, JN-C.
15 . Dean, Donaldson, Adrian, and Dale of Strangeways Laboratory to J. Needham, 29 Nov. 1936,
B.26, JN-C. Interestingly, the RF did end up funding Waddington’s embryology but indirectly through large
grants to Strangeways Laboratory. A grant of $20,000 mostly went toward Fell’s radiation biology group in
1935. A few years later, the RF gave £32,000 to help open a new research wing. RG 1.1, Series 401D, Box
41, Folders 522 and 523, RF-NY.
16 . J. Needham to Hopkins, 26 June 1938, B.28, JN-C.
17 . Hopkins to J. Needham, 15 Aug. 1938, B.28, JN-C.
18 . Hopkins to J. Needham, 20 Sep. 1938, B.28, JN-C.
19 . J. Needham, “Memorandum on Cambridge work on Organiser Phenomena” (undated [I date it
contextually to October 1938]), B.27, JN-C.
20 . Interestingly, many years later, Waddington published on the effects of nuclear weapon radiation on
human and animal development, his work in Fell’s lab giving him some degree of expertise in this area. The
British Foreign Office found Waddington’s article “a very useful summary.” “Press extract, ‘New Statesman
and Nation’ of 6 July 1957,” FO 371/129243, NA-K.
21 . J. Needham to Tisdale, 21 Nov. 1936 [marked “draft”]; and J. Needham to Secretary of Trustees,
Strangeways Laboratories, 27 Nov. 1936, B.26 JN-C. Rockefeller continued this support from 1935 to
1938, according to Abir-Am (“Biotheoretical Gathering, Trans-Disciplinary Authority,” 32).
22 . H. M. Miller to J. Needham, 27 Jan. 1936, B.25, JN-C.
23 . Witkowski, “Optimistic Analysis,” 268.
24 . Waddington, “Studies on the Nature of the Amphibian Organization Centre—VII,” 370–71.
25 . Holtfreter, “Reminiscences”; and Gerhart, “Johannes Holtfreter.”
26 . Waddington, “Homeostasis.”
27 . J. Needham, “An account of the negotiations . . .” (unpublished memo), 12 July 1938, E.114, JN-C.
28 . J. Needham to Toivonen, 1 Nov. 1963, MS 3039.4, CHW-E.
29 . Woodger, Axiomatic Method . John R. Gregg, in the Duke University Zoology Department, did find
Woodger’s axiomatic work fruitful. He and Woodger exchanged many letters in which both tried their
hands at axiomatic statements of biological processes. See C1/G/Gregg, JHW-L. This historical fact
militates against the assertion of Michael Ruse (“Woodger on Genetics,” 2) that “there is little if anything of
value in Woodger’s work.” See Nicholson and Gawne, “Rethinking Woodger’s Legacy.”
30 . See Bernal’s obituary for Franklin, “Dr. Rosalind E. Franklin,” Nature 182, no. 4629 (1958): 154;
and Brenda Maddox, Rosalind Franklin: The Dark Lady of DNA (New York: HarperCollins, 2002).
Interestingly, Dorothy Wrinch kept up a correspondence with Franklin after both had moved away from the
Oxbridge-UCL triangle. See their 1955 correspondence, FRKN/2/33, Rosalind Franklin Papers, Churchill
Archives Centre, Churchill College, Cambridge, England, UK.
31 . Abir-Am, “Biotheoretical Gathering, Trans-Disciplinary Authority,” 68, 109n.
32 . Senechal, I Died for Beauty , 248–53.
33 . Winchester, Man Who Loved China , 36–37 and 50.
34 . Waddington to J. Needham (undated) 1938, M.94, JN-C.
35 . Robertson, “Conrad Hal Waddington,” 592–93. See also Kohler, Lords of the Fly , 271–74.
36 . Waddington to J. Needham, 15 Nov. 1963, M.95, JN-C.
37 . T. H. Morgan, Embryology and Genetics (New York: Columbia University Press, 1934);
Dobzhansky, Genetics and the Origin of Species (New York: Columbia University Press, 1937).
38 . See Waddington’s preface to Organisers and Genes , viii.
39 . “In the ’20s, T. H. Morgan had argued that epigenesis should be considered in terms of the
activities of genes, but he had little effect on the workers in this field. . . . I wanted to return to Morgan’s
idea.” Waddington, Evolution of an Evolutionist , 7.
40 . In Organisers and Genes (p. 45), Waddington emphasized (1) the genetic complex (he introduced
the term “epigenetic” here) acting concertedly and (2) the notion of process versus entity. While living with
Waddington in the mid-1930s, anthropologist Gregory Bateson introduced the notion of regulation in social
systems (he called it “schizmogenesis”). Waddington adopted his own concept of gene regulation and
stabilizing—Bateson called it “negative”—feedback in developmental genetics. Peterson, “Finding Mind,
Form, Organism, and Person,” 240, 481n.
41 . Hall, “Waddington’s Legacy.”
42 . Gerson, “Juncture of Evolutionary and Developmental Biology”; Gilbert, Opitz, and Raff,
“Resynthesizing Evolutionary and Developmental Biology”; and Sansom and Brandon, eds., Integrating
Evolution and Development .

8. AS MANY OPINIONS AS THERE ARE MEN

1 . No minutes exist for any meeting later than 1937. However, scraps of textual and artifactual
evidence suggest another meeting took place in 1938 at the Cornford’s Hunstanton mill. One such scrap I
came across entirely by chance during a visit to the Cornford mill in July 2015: a block of Chinese text
scrawled onto a discarded sheet of plywood propped against a bedroom wall inside the mill. Dr. Di Lara
Luo at the University of Alabama translated this artifact and discovered that it was a poem signed with the
Chinese name taken by Joseph Needham and naming Lu Gwei-Djen, the Chinese scholar then visiting
Cambridge who would become Needham’s second wife after Dorothy’s death. Luo also identified
references to , the concept of “Great Unity” that Needham associated with the organic philosophy (see
chapter 10 ). This interpretation was independently verified by John Moffitt at the Needham Research
Institute, Cambridge, who also confirmed that this handwriting was indeed Joseph Needham’s.
2 . “Churchill’s Scientists,” The Science Museum, Exhibition Road, South Kensington, London SW7
2DD, 9 July 2015.
3 . Woodger, “On Biological Transformations.”
4 . J. Z. Young to Zuckerman, 4 Apr. 1932, SZ/TQ/1/1/11, SZ-N.
5 . Zuckerman, “Obj[ect] of 1st discussion” (unpublished memorandum), SZ/TQ/1/7, SZ-N.
6 . Rubenstein, “Relations of Science, Technology, and Economics.”
7 . Ralph J. Desmarais, “Tots and Quots (act. 1931–1946),” Oxford Dictionary of National Biography ,
http://www.oxforddnb.com/view/theme/95704 .
8 . Levy, Science: Curse or Blessing? (London: Watts, 1940).
9 . [Zuckerman et al., published anonymously], Science in War .
10 . Waddington, The Scientific Attitude (London: Penguin, 1941).
11 . Wittgenstein, Klagge, and Nordmann, Ludwig Wittgenstein , 381–96; and Waddington, Ethical
Animal , 40–41.
12 . Waddington, Science and Ethics , 7
13 . Waddington, “Relations Between Science and Ethics.”
14 . Waddington, Science and Ethics , 18. C. S. Lewis slightly misunderstood Waddington here and
caricatured Waddington’s actual position (which Lewis might have actually had more sympathy for if he
had taken time to understand its context) in That Hideous Strength , the 1945 conclusion to Lewis’s “Space
Trilogy.”
15 . Stephen, Misuse of Mind .
16 . Waddington, Science and Ethics , 60.
 17 . Waddington, Science and Ethics , 71.
18 . Waddington, Science and Ethics , 72.
19 . Waddington, Science and Ethics , 89. Rothschild’s defense of ethology, selfish hereditary units, and
natural aggression would be taken up two decades later by Tinbergen’s Nobel co-laureate, Konrad Lorentz
in the 1960s, and sociobiologists in the 1970s.
20 . Waddington, Science and Ethics , 90–91.
21 . Waddington, Science and Ethics , 92.
22 . Waddington, “Canalization of Development,” 563–65.
23 . Robson and Richards, Variation of Animals ; and see Detlefsen, “Inheritance of Acquired
Characters.”
24 . Waddington, “Canalization of Development,” 563.
25 . Waddington, “Canalization of Development,” 563.
26 . Waddington, “Canalization of Development,” 564.
27 . Waddington, “Canalization of Development,” 565.
28 . Haig, “Epidemiology of Epigenetics,” 16–20; and Lederberg, “Meaning of Epigenetics,” 6.
29 . Waddington, “Epigenotype,” 10.
30 . Waddington, “Epigenotype,” 13.
31 . Kirby, Operational Research , 88–89.
32 . Mead, Blackberry Winter, 252.
33 . Robertson, “Conrad Hal Waddington,” 580.
34 . Abir-Am, “‘Biotheoretical Gathering’ in England,” 349–62.
35 . “I expect to go on with war work, primarily oriented toward the Japanese war. This is partly
because I want to on general grounds, but largely because I have no respectable biological job and can’t see
how to get one. Gray has definitely stated that he will not provide any resources for embryology, even if
more funds are available to his department.” Waddington to J. Needham, 3 Oct., 1944, M.95, JN-C. Sir
James Gray (1891–1975)—a cytologist and expert in tetrapod locomotion—opposed experimental
embryology at Cambridge and the collaboration between Waddington and the Needhams in particular. This
animosity was well known within the British scientific community. Gavin De Beer and Julian Huxley
sensed this hostility and, upon receiving a negative and condescending review of Elements of Experimental
Embryology (Oxford: Oxford University Press, 1934), Huxley entreated Needham to respond on his behalf
by writing to Nature : “Such an article w[oul]d obviously come very well from you, with your biochemical
position & your interest in the philosophy of biology, & would I believe help to clear the air, quite apart
from the specific issue of the review of our book. (You know as well as I do—probably better!—that e.g.,
James Gray has a curious hostility to Experimental Embryology & related subjects, which was revealed in
practical form when the Kings Fellowship was given to Scott[?] in place of to Waddington—and there are
other ‘Modern’ Zoologists who share this so-called physiological view—& to distrust the natural process of
expansion of a [place?] in the position in which Biology now finds itself).” J. Huxley to J. Needham, 27
June 1934, M.34, JN-C.
36 . According to Robertson (“Conrad Hal Waddington,” 581), there were at the time only three chairs
in genetics in the whole of the United Kingdom. Waddington’s introduction to F. A. E. Crew, the individual
in charge of appointing him to the directorship, likely came through Lancelot Hogben, a TBC and Tots &
Quots member and longtime friend of Crew’s. Witness, for instance, Hogben’s admiration for Crew in Clare
Button, “Presentation Skills,” University of Edinburgh Special Collections,
http://libraryblogs.is.ed.ac.uk/towardsdolly/2014/06/30/presentation-skills .
37 . Waddington to Mead, 17 Oct. 1973, Folder 2, Box B17 MM-LC, (punctuation in original).
9. “OFF IN ALL DIRECTIONS LIKE AN EXPANDING
UNIVERSE”
“Off in all directions like an expanding universe” is the phrase Needham used to describe the TBC
members’ trajectories after the war. J. Needham to Woodger, 27 Oct. 1955, M.100, JN-C.

1 . Margaret Mead convinced Needham to keep a descriptive journal covering his first thirty-six hours
in China. He continued this ethnographic style of writing for some time afterward. Before he left for China,
Mead, Gregory Bateson, and Needham likely met in New York for the “Institute of Problems on the War”
meeting held at the Astor Hotel on Saturday, 28 November 1942. The editors of Science and Society , with
whom Mead had a cordial relationship, funded the meeting. Needham gave a talk there entitled “The
Utilization of Scientists,” which mirrored the proposals of Waddington, Solly Zuckerman, and others in
Science in War . Mead and Bateson were working on national morale and cultural analysis at their New
York–based Institute for Intercultural Studies. C.16 and C.17, JN-C.
2 . Needham, Time , 179. Needham quoted from W. H. Mallock, The New Republic; or, Culture, Faith,
and Philosophy in an English Country House (London: Chatto & Windus, 1877), 179.
3 . Needham, Time , 180.
4 . Whitehead, Science and the Modern World , 59.
5 . Needham, “Biologist’s View,” 197.
6 . Needham, “Biologist’s View,” 182–83. Here Needham was quoting Woodger but did not cite a
specific source. Likely this was an aphorism repeated at TBC meetings and in conversations between
Needham and Woodger.
7 . Winchester, Man Who Loved China , 100–101 and 160–61.
8 . MI5 file: “Needham, Joseph Terence Montgomery,” KV2/3055, NA-K.
9 . Winchester, Man Who Loved China , 166.
10 . Werskey, Visible College , 276–77.
11 . Needham, Biochemistry and Morphogenesis , xv–xvi and 119–24.
12 . Needham, Biochemistry and Morphogenesis , 115.
13 . Needham Research Institute, “Science and Civilisation in China Series,”
http://www.nri.org.uk/science.html .
14 . Saha, “Joseph Needham,” 19.
15 . These accusations have persisted but have never been proven definitively, according to Schafer
(“Bacteriological Warfare”). Winchester (Man Who Loved China , 212–15) claims that Needham was
“duped” by Soviet agents who had “created artificially” sites of infection and then tricked Chinese and
Korean bacteriologists into believing that these had been spread by UN and American troops during the
Korean War. These scientists reported their findings, without revealing the Soviet sources, to Needham and
other members of UNESCO. MI5 reported with some concern that Needham seemed to be scoring points
with the British public with arguments that, in spite of the attempts by the British press to refute them, came
across as plausible and backed by evidence. See “Needham, Joseph Terence Montgomery,” KV2/3055, NA-
K.
16 . “Germ War Doubters Routed,” The Daily Worker (27 Sept. 1952) clipping in MI5 file: “Needham,
Joseph Terence Montgomery,” KV2/3055, NA-K.
17 . Lyall, “Joseph Needham.”
18 . Needham, History of Scientific Thought , 281 (italics in the original).
19 . Needham, History of Scientific Thought , 291–93.
 20 . Needham, History of Scientific Thought , 286.
21 . Even a year into the process, Waddington complained that he could not take on other
responsibilities because of procurement and personnel problems. Waddington to Edith Paterson
(Manchester University), 17 Sept. 1947, MS 3059.2, CHW-E.
22 . Robertson, “Conrad Hal Waddington,” 582.
23 . See notes for April 1948 and July 1949 meetings in J.245, JN-C. See also correspondence
regarding these and other TBC meetings in C1/M/22 and C1/2/“Karl Popper’s Letters,” JHW-L.
24 . R. Carnap to Woodger, 19 Aug. 1951, C1/C/5/“Carnaps,” JHW-L.
25 . Popper to Woodger, 26 Jan. 1949, C1/2/“Karl Popper’s Letters,” JHW-L.
26 . Popper, “Obituary: Joseph Henry Woodger,” 330.
27 . Medawar, Thinking Radish , 65.
28 . Medawar, Thinking Radish , 73. But in a seemingly contradictory passage, Medawar also boasted
of being made financially comfortable by way of the salary granted by the demyship in 1935: £350.
29 . Medawar, “Shape of the Human Being.”
30 . Medawar to Woodger [no date] Mar. 1943, C1/M/22, JHW-L.
31 . Medawar to J. Needham, 10 July 1942, M.57, JN-C.
32 . Gibson and Medawar, “Fate of Skin Homografts.”
33 . Medawar to Woodger, 7 Oct. 1945, C1/M/22, JHW-L.
34 . Medawar, Memoir of Thinking Radish , 89–90.
35 . Medawar, Memoir of Thinking Radish , plate 4 records a 1946 meeting of the TBC.
36 . Karl Popper, The Open Society and Its Enemies (London: Routledge, 1945).
37 . The Honorable N. Avrion Mitchison, personal email to author, 2 Dec. 2013.
38 . See their correspondence from Dec. 1945 to Jan. 1946, C1/M/22, JHW-L.
39 . Medawar to Woodger, 1 Oct. 1946, C1/M/22, JHW-L.
40 . Medawar to Woodger, [undated, probably 1949], C1/M/22, JHW-L.
41 . Medawar to Woodger, 20 Feb. 1950, and Medawar to Woodger, 8 Apr. 1950, C1/M/22, JHW-L.
Evolutionary biologist John Maynard-Smith recalled Medawar’s legendary ability to maintain a façade of
politeness even while curtly dismissing someone with whom he disagreed. Interview of John Maynard-
Smith by Richard Dawkins (Apr. 1997), “Peter Medawar: ‘He smiles and smiles and is a villain,’” Web of
Stories , http://www.webofstories.com/play/john.maynard.smith/22 .
42 . Medawar, Review of Biology and Language , 339.
43 . Woodger tentatively approached Medawar again in the mid-1960s and they seem to have come to
some understanding at that time. See correspondence in folder “J. H. Woodger 1964–65,” PP/PBM/A.30,
PBM-W.

10. MECHANISM REDUCED TO MOLECULES

1 . Goodfield, “Theories and Hypotheses in Biology,” 427.
2 . Cassirer, Problem of Knowledge , 205.
3 . Bertalanffy, “Outline of General Systems Theory,” 134.
4 . Bertalanffy, “Outline of General Systems Theory,” 165.
5 . E. S. Russell, Interpretation of Development and Heredity [Oxford: Clarendon, 1930].
6 . Nagel, “Mechanistic Explanation,” quoting E. S. Russell, Interpretation of Development . Though
not necessarily germane to Nagel’s critique, it is interesting to note that Needham and Woodger attempted
to redeem Russell’s work. See Needham to Woodger, 12 Oct. 1929 and return, 3 Nov. 1929: “I agree with
what you say about E. S. Russell. I do not myself see that anything can be done by way of ‘psychobiology.’
I understand from Russell himself that he has changed his opinions,” M.97, JN-C. And see Needham,
“Mechanism,” 239.
7 . E. S. Russell, Interpretation of Development , 172. Nagel examined four quotes from this book in
all, though the one quoted drew most of Nagel’s ire.
8 . Nagel, “Mechanistic Explanation,” 32.
9 . Nagel, “Mechanistic Explanation,” 32.
10 . E. S. Russell, Interpretation of Development , 173.
11 . See, for instance, the repetition of this historical misnomer even among otherwise sophisticated and
nuanced contemporary philosophers: Brigandt and Love, “Reductionism in Biology.”
12 . Watson, Double Helix , 126. Watson recounts the less-than-straightforward manner by which they
became aware of Franklin’s “B-form” images on pp. 104–8.
13 . Schrödinger, What Is Life? , 77.
14 . Sloan and Fogel, introduction to Creating a Physical Biology .
15 . McKaughan, “Influence of Niels Bohr,” 524.
16 . Asimov, In Joy Still Felt ; and Asimov, Intelligent Man’s Guide to Science . The publication history
of the Intelligent Man’s Guide —with multiple editions reprinted over multiple decades—seems remarkable
in comparison with our own era, when guides for Dummies are thought to be sufficient reference works.
17 . Asimov, Intelligent Man’s Guide , 2: 389.
18 . Asimov, Intelligent Man’s Guide , 2: 535.
19 . Asimov, Foundation and Empire , 1.
20 . Commoner, “In Defense of Biology,” 1746.

11. THE LYSENKO MORALITY TALE AND THE

EPIGENETIC LANDSCAPE
1 . Needham, History of Embryology (Cambridge: Cambridge University Press, 1934).
2 . Waddington to Needham, 23 Feb. [no year written, possibly 1942], M.95, JN-C. Despite
pronouncements by the news media, epigenetics is not new. The “prehistory” of epigenetics would start
centuries earlier with Aristotle’s painstaking observations of developing chicks leading to his endorsement
of epigenesis—the notion that organisms are not pre-formed from an already existing “germ” but emerge
gradually but distinctly, piece by piece. There was no tiny chicken in the egg, merely expanding in size; the
heart might form distinctly from the eye and, at a discrete later moment, the lung might accompany the
formation of bones in the wing. In the seventeenth century, anatomist William Harvey would clarify
Aristotle’s concept of epigenesis as the process of “budding out from one another,” moving from a
homogenous liquid substrate to the profusion of complex parts witnessed in an adult organism (Harvey
[1651], quoted in Rieppel, “Atomism, Epigenesis, Preformation,” 331). The next generation of life
scientists including Francesco Redi, Jan Swammerdam, and Regnier de Graaf seemed to overturn Harvey’s
theory with their own observations of eggs and sperm in animals ranging from insects to mammals. With
the additional support of Albrecht von Haller, Lazzaro Spallanzani, and, especially, Charles Bonnet’s
parthenogenesis experiments in the eighteenth century, the antithesis of epigenesis, preformationism,
seemed indisputable (Nordenskiöld, History of Biology , 234–48). When Caspar Friedrich Wolff attempted
to defend epigenesis in his Theorie von der Generation (1764), he did so knowing his endorsement of the
theory would likely cost him a professorship in Germany (Ruffenach, “Caspar Friedrich Wolff”). “Darwin’s
Bulldog,” T. H. Huxley, entered the debates about cell theory and protoplasm in order to advocate the
importance of epigenesis in any explanation of development. In part, Wilhelm Roux named his new
approach “Entwicklungsmechanik” to avoid the partisan battles of the previous century connected to
epigenesis. He could not predict that Hans Driesch would reignite the battle nonetheless, amplifying it in
fact, by interweaving the preformation-epigenesis debate with the still more heated mechanism-vitalism
one. That intertwining would come to bear once again after Waddington re-coined epigenetics. See
Richmond, “T. H. Huxley’s Criticism of German Cell Theory.”
3 . Waddington, “Epigenetics and Evolution.”
4 . See 19 Sept. 1962 entry of Waddington’s Diary of a Visit to Moscow and China as a member of a
delegation from the Royal Society of London to the Academia Sinica (self-published, 1962), SZ/TQ/2, SZ-
N.
5 . Hamburger, “Embryology and the Modern Synthesis,” 97.
6 . Chouard, “Vernalization,” 193.
7 . Paul, “War on Two Fronts.”
8 . Quote from Lysenko’s third speech at the Lenin All-Union Academy of Agricultural Sciences, 19–
27 Dec. 1936, in Medvedev, Rise and Fall , 28.
9 . Muller to J. Huxley, 9 Mar. 1937, Series I: Correspondence (LMC1899), Box 23, HJM-I.
10 . Gruenberg, “Men and Mice at Edinburgh,” 371–74.
11 . Crew et al., “Social Biology,” 521–22.
12 . Pringle, Murder of Nikolai Vavilov .
13 . Dobzhansky to Muller, 14 May 1947, Series I: Correspondence (LMC1899), Box 21, Folder
“Dobzhansky 1947,” HJM-I.
14 . Muller to Dobzhansky, 29 Oct. 1947, Series I: Correspondence (LMC1899), Box 21, Folder
“Dobzhansky 1947,” HJM-I.
15 . See the exchange in Series I: Correspondence (LMC1899), Box 21, Folder “Dobzhansky 1948–
50,” HJM-I.
16 . Sean B. Carroll, personal communication with author in Tuscaloosa, AL, USA, 23 Apr. 2015.
17 . Muller, “Gene as the Basis of Life.”
18 . Medvedev, Rise and Fall , 18.
19 . Chemistry professor Sprizer quoted in “Freedom & Lines,” 44.
20 . President Strand quoted in “Freedom & Lines,” 44.
21 . See, for instance, Waddington, “Lysenko and the Scientists, 1,” 566.
22 . Waddington, “Genetic Assimilation.”
23 . Waddington, Evolution of an Evolutionist , 9.
24 . Waddington, Evolution of an Evolutionist , 9–10.
25 . Waddington, Evolution of an Evolutionist , vii.
26 . Waddington, “Evolutionary Adaptation”; see also Waddington, Evolution of an Evolutionist , 36–
59.
27 . Waddington, Nature of Life , 29.
28 . Waddington, Nature of Life , 22.

12. ERNST MAYR, NEO-DARWINISM, AND BEANBAG

GENETICS
 1 . Mayr repeated many of the claims made in Waddington’s 1952 address to the Symposium of the
Society of Experimental Biology. Sarkar, “Haldane,” 73.
2 . Demerec and Wooldridge, foreword to Cold Spring Harbor Symposium on Quantitative Biology 24,
v.
3 . Stebbins, “Synthetic Approach,” 306.
4 . Stebbins, “Synthetic Approach,” 305.
5 . When discussing evidence for “A-rate” evolution of body size, Kurtén insisted that all variability in
body size must arise through point mutations treated as “‘atomistic’ threshold changes” (Kurtén, “Rates of
Evolution,” 209). The relegation of broad morphological change to minute gene changes was a relatively
new interpretation in midcentury European paleontology, demonstrating the rhetorical force of population
genetics.
6 . Mayr, “Where Are We?” (1959).
7 . Mayr, “Where Are We?” (1959), 2.
8 . Mayr, “Where Are We?” (1959), 2 (my italics).
9 . Waddington made this point in The Strategy of the Genes . Mayr quoted Waddington to make the
same point both in his 1959 and 1976 versions of “Where Are We?” See also Smocovitis, Unifying Biology
, 13–17.
10 . Mayr, “Where Are We?” (1976), 309–10.
11 . “It appears possible, if not probable, that the majority of the findings made on experimental [i.e.,
laboratory] populations cannot be automatically applied to natural populations.” Mayr, “Where Are We?”
(1976), 320.
12 . J. B. S. Haldane, “Defense of Beanbag Genetics,” 435–42. Haldane waited to defend FHW
population genetics against Mayr’s attacking “beanbag genetics” caricature until 1964, after the publication
of Mayr’s Animal Species and Evolution (1963), in which Mayr simply repeated more forcefully the
charges made at the 1959 Cold Spring Harbor symposium. In his “Defense,” Haldane did not hide the fact
that “Wright, Fisher, and I all made simplifying assumptions which allowed us to pose problems soluble by
the elementary mathematics at our disposal, and even then did not always fully solve the simple problems
we set ourselves. Our mathematics may impress zoologists but do not greatly impress mathematicians”
(435). His justification—perhaps unsurprising given Haldane’s socialist proclivities—was an unapologetic
mix of science and politics. Because the mutation rate of individual genes could be thought of as the least
frequent of the list of change agents (having a probability of .00001 per generation, at best, according to
Haldane), it should be possible to establish not only a basal rate of evolutionary change but a means by
which to determine the frequency and, thereafter, check the spread of human genetic disease. But he could
only attack this problem using moderately complex mathematics, though Haldane deemed it “elementary,”
“simple,” and unimpressive. He admitted that “As few people have read my papers on the spread or
diminution of autosomal recessives, and still fewer understood them, the ‘balance’ method, which I
invented, is applied to situations where I claim that it leads to false conclusions.” In other words, Haldane
saw himself and his beanbag genetics as caught between prohibitively complex mathematical and biological
problems on the one hand and important social decisions that could not wait for more understandable, if
also less exact, interpretations of his results on the other. Allowing for an oversimplified way out, beanbag
genetics seemed more desirable from a social standpoint, where one could begin to directly address
problems like hemophilia and muscular dystrophy. Later theorists could go back and make the mathematics
support a less beanbag-like view of genetics.
13 . “Beanbag” debates continue in Borges, “Commentary: The Objection Is Sustained”; Crow,
“Beanbag Lives On,” 771; Crow, “Commentary: Haldane”; Ewens, “Commentary: On Haldane’s
‘Defense’”; Morton, “Commentary: Growth”; and Smith, “‘Something Funny Seems to Happen.’”
14 . Waddington, Strategy of the Genes , 2.
15 . Mayr to Waddington, 20 July 1959, MS 3035.3, CHW-E.
 16 . See Lewontin and Gould, “Spandrels of San Marcos.”
17 . Two decades later, in much more explicitly philosophical works, Mayr did throw his weight behind
organicism—albeit in a qualified way and without acknowledging either the Theoretical Biology Club’s
work or even the first generation of organicists. Mayr, Growth of Biological Thought , 66.
18 . Richmond, “1909 Darwin Celebration.”
19 . “To no other man has it been given to effect a revolution in human thought so large, so pervading,
so sudden, and yet so enduring. Darwin taught mankind to see all things in a new light, not only the
operations of nature, great and small, the mysteries of existence and the innumerable objects of research,
but the common things of every-day life.” “The Darwin Centenary at Cambridge,” The Times (London), 22
June 1909, 9, quoted in Richmond, “1909 Darwin Celebration,” 484.
20 . The committee included Robert Redfield, Tax’s former mentor, who died unexpectedly, but
important anthropologists such as Margaret Mead and Gregory Bateson were not invited. The
paleontologists invited were supporters of the Mayr and Tax party line. See Gould, “Irrelevance,
Submission, and Partnership.”
21 . Tax did not invite the aged Ronald A. Fisher to the event, possibly because of the controversy
between him and Sewall Wright. Wright’s attendance, especially after his conversion to pan-selectionism,
was assured by Mayr, Simpson, and Dobzhansky. See Plutynski, “Parsimony and the Fisher-Wright
Debate”; and Smocovitis, “1959 Darwin Centennial Celebration in America.”
22 . Tax, “Celebration,” 274–75.
23 . Smocovitis, “1959 Darwin Centennial Celebration,” 303–5.
24 . Waddington to Mead, 30 Nov. 1959, Box B17, Folder 1, MM-LC.
25 . Tax included only the panel on paleoanthropology in the Issues in Evolution , vol. 3. Simpson et al.,
“Panel Three.”
26 . Simpson et al., “Panel Three,” 147–49 and 171–73. Notably, Waddington complained that the
transcription of his contributions to this panel “required very considerable alteration.” Waddington to Tax,
11 Dec. 1959, MS 3059.10, CHW-E.
27 . Waddington to Mead, 15 Apr. 1959, Box B17, Folder 1, MM-LC.
28 . Mayr, “Emergence of Evolutionary Novelties”; and Dobzhansky, “Evolution and the
Environment,” 422.
29 . Waddington, “Evolutionary Adaptation,” 400.
30 . Waddington, “Evolutionary Adaptation,” 401. A strikingly similar model, and cry for “escape”
from neo-Darwinian oversimplification, would be (re-)issued nearly a half century later by Jablonka and
Lamb, Evolution in Four Dimensions .
31 . Hamburger, “Embryology and the Modern Synthesis,” 97–112.
32 . Schmidt to Waddington, 13 Sep. 1956, MS 3059.10, CHW-E.
33 . Schmidt to Waddington, 12 Apr. 1957, MS 3059.10, CHW-E.
34 . Tax to Waddington, 6 Dec. 1957, MS 3059.10, CHW-E. Waddington visited Mead both before and
after the Chicago Darwin celebration. From his written responses to her unpreserved verbal commentary,
she seemed concerned about the lack of rapprochement between the neo-Darwinians and cultural
anthropologists leading to oversimplifications about humans—biologists making “expert” comments in a
field in which they had no training. Waddington to Mead, 4 June 1958, 15 Apr. 1959, and 30 Nov. 1959,
Box B17, Folder 1, MM-LC; and 18 Sep. 1959 and 17 Jan. 1960, MS 3060.1, CHW-E.
35 . Waddington to Tax, 13 Dec. 1957, MS 3059.10, CHW-E.
36 . J. Huxley to Waddington, 4 Nov. 1959, MS 3059.10, CHW-E.
37 . Grene, “Two Evolutionary Theories, 1,” and “Two Evolutionary Theories, 2.”
38 . Smocovitis, “1959 Darwin Centennial Celebration,” 287–88. Tax’s attitude here contrasts strikingly
with the supposed détente between nature and nurture camps understood to be “canonical” following World
War II in works like Cravens, Triumph of Evolution .
39 . Kluckhohn et al., “Panel Five,” 208–10.
40 . Kluckhohn et al., “Panel Five,” 243.
41 . Julian Huxley headed the London Eugenics Society in the late 1950s. C. D. Darlington preceded
him in this role, according to notes in MS 3044.2, folder 1, CHW-E.

13. HISTORY OF SCIENCE IS WRITTEN BY THE

LAUREATES
1 . The following quotations are from Waddington to June Goodfield Toulmin, 9 Apr. 1965, M.89, JN-
C.
2 . The following quotations are from Needham to June Goodfield Toulmin, 26 Apr. 1965, M.89, JN-C.
3 . Goodfield Toulmin confirmed this in her return letter of 23 May 1965, M.89, JN-C.
4 . J. Needham to Medawar, 21 Apr. 1951, M.57, JN-C.
5 . Medawar to J. Needham, 3 May 1951, M.57, JN-C.
6 . J. Needham to Medawar, 10 May 1951, M.57, JN-C.
7 . Medawar, “Biological Retrospect,” 1327.
8 . Medawar, “Biological Retrospect,” 1327.
9 . Medawar, “Biological Retrospect,” 1328.
10 . Medawar, “Biological Retrospect,” 1329.
11 . According to their comments, Needham and Waddington seemed especially dismayed at the lack of
recognition Medawar accorded to Needham’s Order and Life and Waddington’s attempt to synthesize
development and genetics in Organisers and Genes —omissions that may have allowed Medawar to make
comments such as, “The underlying assumption of the theory (though not then so expressed) was that we
should look to the chemical properties of the inductive agent to find out why the amino-acid sequence of
one enzyme . . . should differ from the amino-acid sequence of another.” Waddington underlined Medawar’s
parenthetical phrase “(though not then so expressed)” and mocked it in another portion of the article. His
sarcasm indicates the extent to which he and Needham felt someone who was in a position to know better
was intentionally distorting their work. M.89, JN-C.
12 . Waddington to J. Needham, 15 Nov. 1963, M.95, JN-C.
13 . Goodfield, “Theories and Hypotheses in Biology.”
14 . The following quotations are from Waddington’s report on June Goodfield’s renewal grant
application for 1968. Waddington marked it “first draft, not used” but attached this copy to his 23 July 1968
response to the National Science Foundation. MS 3041.4, CHW-E.
15 . Waddington, “No Vitalism for Crick.”
16 . Francis Crick to Waddington, 9 Nov. 1967, PP/CRI/I/2/6/5, FC-W. Throughout their
correspondence, Waddington and Crick referred to each other in quite familiar, cordial terms. Though
Waddington was not at Cambridge concurrently with Crick, they were well known to each other through
other circles. Crick would attend Waddington’s IUBS theoretical biology meeting at Lake Como, Italy, in
1969.
17 . Waddington to Crick, 27 Dec. 1967, PP/CRI/I/2/6/5, FC-W.
18 . Lord Halsbury’s memorandum of 31 Oct. 1951 and the correspondence from Lord Halsbury to D.
Gabor, 23 Apr. 1952, were retrieved by Anne Brown, Waddington’s secretary in 1976, with indications that
Waddington had first brought this up to the broader scientific community in 1970. MS 3040.6, CHW-E.
19 . Crick to Waddington, 6 June 1974, MS 3040.6, CHW-E.
20 . Waddington to Crick, 8 June 1974, MS 3040.6, CHW-E.
21 . Crick to Waddington, 28 June 1974, MS 3040.6, CHW-E.
22 . Interestingly, just before this exchange, Crick had endorsed the notion of directed panspermia—the
intentional “infecting” of Earth with living microorganisms by intelligent, and undetectable,
extraterrestrials. Though admittedly without evidence and unfalsifiable, Crick begged his audience to take a
“wait and see” approach rather than passing the kind of quasi-Popperian judgment on his speculations that
he was now passing on Waddington’s epigenetics. See Crick and Orgel, “Directed Panspermia.”
23 . Waddington to Crick, 10 July 1974, MS 3040.6, CHW-E.
24 . Waddington to Crick, 10 July 1974, MS 3040.6, CHW-E.
25 . Crick to Waddington, 30 July 1974, MS 3040.6, CHW-E.
26 . Waddington to Crick, 26 Aug. 1974, MS 3040.6, CHW-E.
27 . “Peace! Peace!,” pronounced Crick. Crick to Waddington, 4 Sep. 1974, MS 3040.6, CHW-E.

14. THE 1960s REINCARNATION OF THE DEBATE

Epigraph: Carson, Silent Spring , 189.
1 . Esposito, who offers a different gloss on organicism in the interwar period, and who agrees that
organicism “survived” the midcentury gene-centered redefinition of mechanism, nevertheless repeats the
mistaken impression of earlier historians that “third way” theorists in the mid-twentieth century “had no
relevant impact and stimulated no change within the scientific world” (Romantic Biology , 183). The
present chapter argues that, while the level of interpenetration of organicism into the biological community
in the 1960s did not reach the level of the 1930s, organicism continued to reach a significant number of
scholars. Much like the situation almost a half century earlier, the backlash against mechanism expressed
popularly and within the scientific community drove organicists including Waddington to articulate an
alternative between gene-centric mechanism and a new, strangely vitalistic bioexceptionalism.
2 . Commoner, “In Defense of Biology,” 1746. See also the extensive correspondence between Barry
Commoner and Linus Pauling at Oregon State University Libraries Special Collections, “Linus Pauling
Day-by-Day,” http://osulibrary.oregonstate.edu/specialcollections/coll/pauling/calendar/index.html .
3 . Commoner, “In Defense of Biology,” 1746–47.
4 . Commoner, Science and Survival , 27–28.
5 . Commoner, Science and Survival , 45.
6 . T. H. Huxley, for instance, worried that specialization would fracture the discipline, with detrimental
social consequences. Desmond, Huxley , 628–30.
7 . Bertalanffy, Problems of Life .
8 . Hans Jonas, The Phenomenon of Life: Towards a Philosophical Biology (Evanston, IL: Northwestern
University Press, 1966).
9 . Grene, “Two Evolutionary Theories, 1” and “Two Evolutionary Theories, 2.”
10 . Polyani, “Science and Religion,” 10.
11 . Polyani, “Life’s Irreducible Structure,” 1310.
12 . Sonneborn, “Bearing of Protozoan Studies,” 221.
13 . Joshua Lederberg asserted that the “DNA paradigm” is the chief representative of this “triumph of
mechanistic interpretation.” Lederberg, “What the Double Helix (1953) Has Meant,” 1985.
 14 . Koestler, “Opening Remarks,” 1. Among others, Koestler was friends with game theorist John von
Neumann, philosophers A. J. Ayer, Bertrand Russell, John Dewey, Karl Jaspers, Jean-Paul Sartre, Simone
de Beauvoir, and novelists George Orwell and Dylan Thomas.
15 . He did not, however, believe that Soviet science was much better. Its faults were simply different in
emphasis. See Koestler, “Retrospect.”
16 . Koestler, Sleepwalkers , 545–46.
17 . Koestler, Yogi and the Commissar ; and Ghost in the Machine .
18 . Koestler, Sleepwalkers , 550.
19 . Koestler, Sleepwalkers , 547.
20 . Koestler, Case of the Midwife Toad . Koestler completely ignored the fact that the legacy of
William Bateson was already being deployed by Ernst Mayr, among others, as anti -Darwinian and the chief
obstacle to be overcome in the making of the neo-Darwinian synthesis. See Peterson, “William Bateson
From Balanoglossus to Materials for the Study of Variation .”
21 . Alpbach symposium accounting balance sheet, dated “1968,” MS 3062.10, CHW-E.
22 . Weiss, “Living System,” 9–10.
23 . Koestler was supportive of Waddington’s late attendance and, in fact, altered the schedule of the
symposium in such a way to make certain he could attend almost every talk. Koestler to Waddington, 6 May
1968, and Mischa Fischberg to Waddington, 29 Jan. 1968, MS 3062.10, CHW-E.
24 . Waddington, “Comment on Bertallanfy [sic ],” MS 3062.10, CHW-E.
25 . In an 18 Oct. 1968 letter to Waddington [missing], Bertalanffy agreed with Waddington’s earlier
assessment that Bertalanffy’s paper was deficient in several ways. Waddington took Bertalanffy’s
conciliatory letter as implicit permission to critique the paper more thoroughly; hence the “Comment,”
which Waddington assumed Koestler would publish alongside Bertalanffy’s essay. He did not.
26 . Waddington, “Comment on Bertallanfy [sic ].” Similar sentiments would appear in Waddington,
Tools for Thought . Note that Waddington clearly thought the “third way” had the hallmarks of a Kuhnian
paradigm. Though I argue this is incorrect, it is telling that Waddington believed organicism had already
risen to the level of broad acceptance by the 1960s rather than being discarded to the dustbin of history as
some have lamented.
27 . A few years later, Waddington would reverse his judgment, categorizing Bertalanffy’s general
systems theory as part of the same “organicist anti-reductionis[m]” as he and other TBC members refined in
the 1930s. Waddington, “Evolution of Mind,” 75.
28 . Waddington, preface to Essays .
29 . Interestingly, Koestler (Ghost in the Machine , 67) cited Joseph Needham’s work in the 1930s as an
inspiration for the “holon” concept, which he defined as a “part-whole”: “Different levels represent different
stages of [biological] development, and the holons . . . reflect intermediary structures at these stages.” The
notion sounded much like Waddington’s “chreods,” despite Waddington’s rejection of the association at
Alpbach.
30 . Mayr to Waddington, 10 May 1965, HUG(FP) 74.7, Box 12, Folder 874, EM-H.
31 . Mayr to Waddington, 10 May 1965, HUG(FP) 74.7, Box 12, Folder 874, EM-H.
32 . Waddington, “Basic Ideas of Biology,” 5–12.
33 . Mayr to Waddington, 9 Nov. 1966, HUG(FP) 74.7, Box 14, Folder 919, EM-H.
34 . Mayr to Grene, 20 Dec. 1967, HUG(FP) 74.7, Box 14, Folder 940, EM-H.
35 . Mayr mentioned only that he wanted a conference specifically on “philosophy of biology” rather
than Waddington’s “theoretical biology” (Mayr to Waddington, 7 Dec. 1966, HUG[FP] 74.7, Box 14,
Folder 919, EM-H). Years later, Mayr reflected that it was around that time (1967) that he realized he
needed to champion personally the philosophical views he preferred. “Problems of the philosophy of
biology,” interview of Ernst Mayr by Walter J. Bock (Oct. 1997), Web of Stories (24 Jan. 2008),
http://www.webofstories.com/people/ernst.mayr/134?o=SH .
36 . Waddington, ed., Towards a Theoretical Biology .
37 . Pattee, “Problem of Biological Hierarchy,” 135, 1n.
38 . Pattee, “Problem of Biological Hierarchy,” 134–35 (italics in the original).
39 . Waddington, Behind Appearance . This defaced example was found in the stacks at the University
of Notre Dame’s Theodore Hesburgh Library, somewhere behind the iconic “Touchdown Jesus.”

15. THE CONVENTIONAL WISDOM OF THE DOMINANT

GROUP, OR COWDUNG
1 . “Comment on Bertallanfy” [sic ], attachment, Waddington to Koestler, 21 Sept. 1968, MS 3062.10,
CHW-E.
2 . Waddington, Tools for Thought , 233 (italics in the original).
3 . Waddington, Tools for Thought , 16.
4 . Waddington, Tools for Thought , 16, footnote. By the middle of the twentieth century, philosophers
of science had long been interested in logical empiricism, which likely also has relevance for this story. See
Reisch, How the Cold War Transformed Philosophy of Science , 344–68.
5 . Waddington, Tools for Thought , 18 (italics in the original).
6 . Waddington was certainly correct that he was following the path blazed by Whitehead in particular.
Whitehead spent his Lowell Lectures attacking “scientific materialism” as “entirely unsuited to the
scientific situation at which we have now [1925] arrived” (Science and the Modern World , 18). Fifty years
later, Waddington found himself facing opponents making surprisingly similar claims.
7 . Waddington, Evolution of an Evolutionist , 11.
8 . Waddington, Tools for Thought , 20.
9 . Waddington, Tools for Thought , 23.
10 . Waddington, Tools for Thought , 24–25.
11 . Waddington, Tools for Thought , 22.
12 . Waddington, “Mindless Societies.”
13 . Allen, Beckwith, Beckwith, Chorover, et al., “Against Sociobiology.”
14 . The most accessible reprinting of Waddington’s essay appears in Arthur Caplan, ed., The
Sociobiology Debate: Readings on Ethical and Scientific Issues (New York: HarperCollins, 1978). This
version excises the entire second half of the essay, which reviews Laughlin and d’Aquili and connects their
discussion of structuralism to Waddington’s Whiteheadian philosophy of organism in order to critique some
of the central themes in Wilson’s Sociobiology .
15 . Waddington to Mead, 17 Oct. 1973; and return, 19 Oct. 1973, Box B17, Folder 2, MM-LC; and
Waddington to Bateson, 12 Jan. 1974, MS 3040.3, CHW-E.
16 . Waddington to Mead, 17 Oct. 1973, Box B17, Folder 2, MM-LC.
17 . Waddington, “Age of Conformity.”
18 . Waddington and Cage (eds.), Biology and the History of the Future .
19 . Waddington, “Mindless Societies.”
20 . Robertson, “Conrad Hal Waddington,” 612–14.
21 . D. R. Newth to J. Needham [undated], M.95, JN-C.
 22 . E.g., Waddington, “Canalization of Development and the Inheritance of Acquired Characters”
(1942), “Selection of the Genetic Basis for an Acquired Character” (1952), and “Paradigm for an
Evolutionary Process” (1969).
23 . Waddington, “Practical Consequences,” 81. Grene’s comments are problematic and deserve
consideration. At the 1967 IUBS meeting in Italy, Grene lumped Waddington with other conventional neo-
Darwinians, including John Maynard-Smith. Waddington had attacked, on more than one occasion,
Maynard-Smith’s conception of neo-Darwinism, and both Waddington and Maynard-Smith believed they
had significant theoretical conflicts with one another. Grene, perhaps reading out of Waddington’s
“unaggressive character,” saw the two as fairly closely related. According to Grene, “Almost everyone
used, constantly and as self-evident, the term ‘biological’ as synonymous with ‘functional,’ ‘adaptive,’
‘conducive to survival’—strictly, in evolutionary terms, conducive to leaving descendants—or ‘produced by
Natural Selection.’ . . . even Waddington, though not quite orthodox, appears in his summarizing notes as an
interesting mutant of the same species.” Grene, “Bohm’s Metaphysics and Biology,” 65. It seems possible
that Grene’s critique spurred Waddington to disclose his organicist views somewhat more openly in his final
works.
24 . Robertson to Bateson, 7 May 1976; and return, 19 Aug. 1976, MS 98, Box 36, Folder 1447, GB-
SC.
25 . Waddington, “Fifty Years On,” 21.

CONCLUSION: A THIRD WAY AFTER WADDINGTON?

1 . Grossman, “Review of Gödel, Escher, Bach .”
2 . D. Hofstadter, Gödel, Escher, Bach , 311–36.
3 . D. Hofstadter, Gödel, Escher, Bach , 321 (emphasis in the original).
4 . D. Hofstadter, Gödel, Escher, Bach , 322.
5 . Interestingly, in his summary of the fourth and final IUBS symposium, written in 1972, Waddington
inverted the comparison: Since biology, specifically the genotype-to-phenotype developmental process, was
so difficult to understand but seemed to function like symbolic communication, theoretical biologists should
make use of those tools used in the philosophy of language (e.g., Wittgenstein) to help make sense out of
biology.
6 . D. Hofstadter, Gödel, Escher, Bach , 233.
7 . Junod, “E. O. Wilson,” http://www.esquire.com/news-politics/interviews/a5375/eo-wilson-quotes-
0109/ .
8 . Waddington, Tools for Thought , 16.
9 . See, for example, Pitney, “Fabled Fountain of Youth.”
10 . E. S. Russell, Interpretation of Development , 173.
11 . Ritter, Unity of the Organism , v. 2, 149.
12 . The connection between the second generation of organicists and these new proponents is
straightforward: Waddington was on good terms with both Simpson and Dobzhansky at the end of his
career, counting Dobzhansky among his closest friends. Gould corresponded with both Waddington and
Needham, having been assigned to read their texts by Simpson; Lewontin knew Waddington not only
through Dobzhansky and publications but participated in several of Waddington’s IUBS theoretical biology
gatherings in the late 1960s. See Gould to Waddington, 16 June 1972, MS 3040.3, CHW-E; and Gould to J.
Needham, 4 Apr. 1965, B.132, JN-C.
13 . Simpson, This View of Life , 106–7.
14 . Dobzhansky, Genetic Diversity , 8.
 15 . Eldridge and Gould, “Punctuated Equilibrium.”
16 . Lewontin, Biology as Ideology , 122.
17 . Pearl to Woodger, 1 Aug. 1929, C1/2/P, JHW-L (italics in the original).
18 . Snow, The Two Cultures .
19 . Waddington, “Fifty Years On,” 21.
20 . Zuckerman to Medawar, 28 Jan. 1980, Folder “Personal correspondence, Medawar,” SZ-N.
21 . Massimo Pigliucci, “Neil deGrasse Tyson and the Value of Philosophy,” Huffpost Science (16 May
2014; updated 16 July 2014), http://www.huffingtonpost.com/massimo-pigliucci/neil-degrasse-tyson-and-
the-value-of-philosophy_b_5330216.html ; and Olivia Goldhill, “Why Are So Many Smart People Such
Idiots About Philosophy?,” Quartz (6 Mar. 2016), http://qz.com/627989 .
22 . Kuhn, Structure of Scientific Revolutions , 165.
23 . Kuhn, Structure of Scientific Revolutions , 169.
24 . Duhem, Aim and Structure , 334–35.
25 . Polanyi, Personal Knowledge , 347–64; and Nye, Michael Polanyi .
26 . Waddington, Evolution of an Evolutionist , 10.
27 . Sachs, “Is the NIH Budget Saturated?,” 29–30.
28 . March, “Broader Impacts Review Criterion.” Susan Fitzpatrick, scientist and president of the James
S. McDonnell Foundation, recently made a similar point. See Fitzpatrick, “Opinion—Making Progress by
Slowing Down,” The Scientist (24 Aug. 2015), http://www.the-scientist.com/?
articles.view/articleNo/43820/title/Opinion—Making-Progress-by-Slowing-Down/ .
29 . Waddington, “Sketch,” 1.
30 . Waddington, Strategy of the Genes , 2.
31 . Hein, “Molecular Biology vs. Organicism,” 238. Though Hein does not mention him by name,
Popper does seem to make this sort of claim.
32 . See Quine, “Two Dogmas.”
33 . Hein, “Mechanism and Vitalism.”
34 . Goodfield, “Changing Strategies,” 65.
35 . Goodfield, “Postscript,” 85–86. She clearly did not recall her exchanges with Waddington and
Needham a decade earlier over how much their belief in holism did influence the organizer pursuit and
after.
36 . Loeb to Pearl, 21 Jan. 1913, Box 11, Pearl correspondence folder, JL-LC.
37 . Hein, “Molecular Biology vs. Organicism,” 253, 10n.
38 . Both quotes appear in Elsasser, Reflections , 3.
39 . Suttie to Woodger, 7 Nov. 1919, C1/3/Suttie, JHW-L.

EPILOGUE: IS MODERN EPIGENETICS ORGANIC?

1 . Some of the hype of the early years of twenty-first-century biomedical research into epigenetics is
captured in Joel Wallach, Ma Lan, and Gerhard Schrauzer, Epigenetics: The Death of the Genetic Theory of
Disease Transmission (New York: Select Books, 2014).
2 . Much more information, including the methodology of the larger study, can be found at
http://epigenetics.as.ua.edu .
3 . Ezkurdia et al., “Shrinking Human Protein Coding Complement.”
 4 . Waddington carried his work to the egg level with Okada and Waddington, “Submicroscopic
Structure of the Drosophila Egg.” But his desire for epigenetics was not limited to eggs. The problem, as he
identified it, was that there was no understood connection between genotype and phenotype. Neo-
Darwinians might wave their hands and say that evolution has to do with genes in a population, but that
cannot be the real story, insisted Waddington. Evolution is about phenotypes. By the end of the twentieth
century, we still did not know much about how phenotypes are created from genotypes.
5 . Robin Holliday’s terminology, quoted in Lederberg, “Meaning of Epigenetics,” 6. Other historians
of science and scientists have made similar observations, though Lederberg makes the distinction clearest.
See, for instance, Morange, “Relations Between Genetics and Epigenetics.”
6 . Veenendaal et al., “Transgenerational Effects of Prenatal Exposure to the 1944–45 Dutch Famine.”
7 . Lewontin, Rose, and Kamin, Not in Our Genes , 57.
8 . Sapp, Beyond the Gene , particularly chapter 5.
9 . See Peterson, Grace, Hauger, and Wilson, “The Genesis of Epigenetics,” for a more detailed outline
of our work.
10 . Holliday and Pugh, “DNA Modification Mechanisms and Gene Activity During Development.”
11 . Weinberg, “Coming Full Circle.”
12 . For example, see Yehuda et al., “Holocaust Exposure Induced Intergenerational Effects on FKBP5
Methylation.”
13 . Crew et al., “Social Biology,” 521–22.
 BIBLIOGRAPHY

ARCHIVAL SOURCES
American Philosophical Society, Philadelphia, PA, USA

Theodosius Dobzhansky papers [TD-APS]

Leslie C. Dunn papers [LD-APS]
M. F. Ashley Montagu papers [AM-APS]
George G. Simpson papers [GS-APS]

Cambridge University, Cambridge, England, UK

John Desmond Bernal papers [JB-C]

Joseph Needham papers [JN-C]

Harvard University, Cambridge, MA, USA

Ernst Mayr papers [EM-H]

Hoover Institution, Stanford University, Palo Alto, CA, USA

Karl Popper papers [KP-S]

Indiana University, Bloomington, IN, USA

Hermann J. Muller papers [HJM-I]

Library of Congress, Washington, DC, USA

Jacques Loeb papers [JL-LC]

Margaret Mead papers/Southeast Pacific Ethnography archive [MM-LC]

National Archive, Kew, England, UK [NA-K]

Rockefeller Foundation Archive Center, Sleepy Hollow, NY, USA [RF-NY]

University College London, London, England, UK

Joseph H. Woodger papers [JHW-L]

University of California-Santa Cruz, Santa Cruz, CA, USA

Gregory Bateson papers [GB-SC]

University of Chicago, Chicago, IL, USA

Darwin Centenary Celebration (1959) archive [DCC-C]

University of East Anglia, Norwich, England, UK

Solly Zuckerman papers/Tots & Quots archive [SZ-N]

University of Edinburgh, Edinburgh, Scotland, UK

Conrad H. Waddington papers [CHW-E]

Wellcome Trust, London, England, UK

Francis H. C. Crick papers [FC-W]

Peter Medawar papers [PM-W]

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 INDEX

Abir-Am, Pnina, 262n17

acquired characteristics becoming fixed. See canalization
Agassiz, Louis, 21
“The Age of Conformity: The Autocatalytic Loss of Freedom” (Waddington), 230
Agricultural Research Council (ARC), 139 , 140
AI. See artificial intelligence (AI)
Akademic Olympia, 268n21
All-Union Congress of Genetics, Selection, Plant and Animal Breeding, 174
Alpbach Symposium, 217 –19 , 220 –21 , 224 , 226 , 289n23 , 289n29
American Association for the Advancement of Science, 213
American Philosophical Association (APA), 12 , 31 –33 , 35 , 40 , 61
Analyse der empfindungen [Analysis of Sensations] (Mach), 63 , 268n21
anti-Darwinism, 289n20
anti-Lysenkoism, responses to, 179 –81
anti-mechanism, 11 , 40 –41 , 114 , 236 , 238 , 248 –49 , 259 , 268n21 ; Driesch’s attacks on mechanism, 31
, 211 , 281n2 (chap. 11) ; E.S. Russell and, 38 , 41 ; GST theory as, 160 , 162 –63 ; Hofstadter on, 243 ;
J.S. Haldane and, 30 , 38 , 39 , 64 , 85 , 105 ; Koestler and, 214 –21 ; neo-mechanism as alternative, 53
; Rignano and, 42 –45 , 48 , 50 , 51 –52 ; Woodger on, 61 –62 , 63 , 64 , 66 , 86 , 160 , 268n22
anti-reductionism, 248 , 289n27
anti-vitalism, 32 , 33 , 35 , 85
ants as analogy for human complexity, 235 –37
“Are We ‘Cell Aggregates’?” (Montgomery), 18
Aristapedia experiments, 108
Aristotle, 5 , 62 , 108 ; impact of on new generations, 35 , 39 , 41 , 52 , 87 , 134 , 147 ; precursor of
epigenetics, 172 , 281n2 (chap. 11)
Aristotelian Society, 105
Arnold Gerstenberg Prize in Philosophy, 101
artificial intelligence (AI), 235 –36
Asimov, Isaac, 159 –60 , 168 –70 , 173 , 184 , 191 , 212
Atlantic, Battle of, 128
atomism, 5 , 66
The Axiomatic Method in Biology (Woodger), 123 , 152 –53 , 154 , 156 , 275n29
Ayer, A. J., 114 , 288n14
Ayer, Freddie, 118 , 156
Bacon, Francis, 33
Baer, Karl Ernst von, 39 , 68
Baird, Spencer Fullerton, 21
Ballard Matthews Memorial Lectures, 220
Barnes, E. W., 48
“The Basic Ideas of Biology” (Waddington), 220 , 221
Bateson, Gregory, 216 , 271n23 ; Margaret Mead and, 132 , 133 , 139 , 278n1 , 285n20 ; Waddington and,
100 –101 , 125 , 132 , 133 , 233 , 271n23 , 276n40
Bateson, William, 62 , 100 , 191 , 193 , 216 , 218 , 289n20
Beadle, George, 184
“beanbag genetics,” 174 , 189 –93 , 194 , 195 , 200 , 283n1 , 284n12
Beauvoir, Simone de, 288n14
Beckner, Morton, 7
behavioralism, 264n30
Behind Appearance: A Study of the Relations Between Painting and the Natural Sciences in This Century
(Waddington), 224 –25 , 290n39
Benedict, Ruth, 132
Bergson, Henri, 19 , 31 , 32 , 211 ; concept of élan vital, 4 –5 , 29 , 51 , 131 , 160 ; as a neo-vitalist, 51 , 52 ;
popularity of, 3 –4 , 142 , 261n3 ; as a vitalist, 6 , 12 , 13 , 15 , 35 , 41 , 65 , 108 , 132 , 143 –44
Bernal, John Desmond “Sage,” 12 , 83 –85 , 127 , 129 –30 , 242 , 249 ; change of careers, 123 , 124 , 128 ;
co-writing Science in War (anonymously published), 139 ; crystallography work, 83 , 107 , 112 , 116 ,
124 ; Needham dedicating Order and Life to TBC members, 114 –15 ; politics of, 85 , 111 , 114 , 119 ,
124 , 129 , 146 , 214 , 274n8 ; and SICHST, 81 , 83 , 85 , 129 –30 ; and the Theoretical Biology Club,
104 , 107 , 109 , 114 ; and Tots & Quots, 127 , 128 , 129 , 130
Bernard, Claude, 19 , 30 , 33
Bertalanffy, Ludwig von, 70 , 188 , 192 , 213 , 226 , 241 ; and the Alpbach Symposium, 217 , 218 –19 ,
289n25 ; and the General Systems Theory, 160 –63 , 165 , 218 –19 , 289n27 ; on organic philosophy,
162 –63 ; Woodger and, 112 , 163
Beyond Reductionism (edited by Koestler and Smythies), 217 , 224
biocentric, universe as, 266n54
Biochemical Embryology (Needham), 124 , 146 –47
biochemical warfare, 149 , 279n15
biochemistry, 8 , 78 , 93 , 129 , 154 , 163 , 209 , 212 ; Asimov and, 168 , 170 ; and embryology, 119 , 141 ,
184 , 202 ; Joseph Needham and, 46 , 47 , 49 , 71 , 118 , 119 , 122 , 124 , 146 –47 ; and mechanists, 39 ;
Waddington and, 99 , 122
Biochemistry and Morphogenesis (Needham), 147 , 240
Biogenetic Structuralism (Laughlin and d’Aquili), 230
Biographical Memoirs of the Royal Society (Robertson), 231 –32
Biological Principles (Woodger), 62 , 63 –65 , 68 –69 , 70 , 72 –73 , 75 , 80 , 172 , 269n50
“A Biological Retrospect” (Medawar), 203 –5
Biologische Versuchsanstalt [Biological Research Institute], Vienna, 60 –61 , 70 , 105
biology. See biochemistry; embryology; evolution; genetics; language of biology; philosophy of biology;
philosophy of science; theoretical biology
“Biology and the History of the Future” symposium, 230 –31
Biology as Ideology (Lewontin), 241
Biology Under the Influence (Lewontin and Levins), 241
“biotheoretical gathering.” See Theoretical Biology Club
Black, Max, 114 –15
Blackett, Patrick, 128
Bohm, David, 222 , 246
Bohr, Niels, 12 , 62 , 90
Bonnet, Charles, 281n2 (chap. 11)
Boston Colloquium for the Philosophy of Science, 205
Boyle, Robert, 63 , 81
Brachet, Jean, 50 , 112 , 121 , 147 , 204
The Brain Trust (radio program), 128
Brave New World (Huxley), 177
Bristol University (University College Bristol), 24
Britain, Battle of, 155
British Academy, Fellows of, 45 , 267n6
British Association for the Advancement of Science, 88 –92 , 105 , 157 , 203 –5
British Bombing Survey Group, 157
British Socialist Party, 49
Brown, Anne, 287n18
Brücke, Ernst, 5
Bukharin, Nikolai I., 55 , 82 –83 , 85 , 87 , 88 , 92 , 117 , 129 ; Levy publishing reinterpretation of, 130
Bultmann, Rudolf, 213
Burdon-Sanderson, John Scott, 28 –29
Bush, George W., 251
Butterfield, Herbert, 8

Calkins, Mary Whiton, 31

Caltech, 118 , 125 , 139 , 182 , 201
Cambridge Moral Science Club, 130
Cambridge Theoretical Biology Club. See Theoretical Biology Club
Cambridge University, 9 , 84 , 127 ; Darwin celebration in 1909, 193 –94 ; lack of support of research on
experimental embryology at, 117 –22 , 123 , 139 , 146 , 276n1 , 278n35 ; Needham and, 43 , 47 , 49 ,
50 , 111 , 124 , 146 , 151 , 276n1 ; Needham Research Institute, 46 , 149 , 276n1 ; organizer
experiments of Waddington and Needhams, 55 , 93 –94 , 98 –99 , 103 , 112 , 117 , 119 , 143 , 156 , 159
, 200 –202 , 204 , 255 ; Waddington and, 99 , 103 , 125 , 127 , 151 –52 , 232 , 287n16
Cambridge Zoological Society, 70
canalization, 216 ; and acquired characteristics becoming fixed, 180 , 258 ; Waddington on acquired
characteristics and, 134 , 135 –38 , 162 , 181 –82 , 183 , 232
“Canalization of Development and the Inheritance of Acquired Characters” (Waddington), 134 , 137 , 138
Carnap, Rudolf “Rudi,” 118 , 152 , 163
Carson, Rachel, 211 , 224
The Case of the Midwife Toad (Koestler), 216 , 217
Cassirer, Ernst, 160 , 163
“Cause and Effect in Biology” (Mayr), 220
cell theory, 67 , 71 ; cell lineage studies, 209 ; E. B. Wilson on, 64 ; Montgomery’s views on, 16 –17 ;
Scheiden and Schwann on, 21 ; T. S. Huxley on, 67 , 281n2 (chap. 11) ; Weismann on chromatin
controlling properties of cells, 77 –78 ; Whitman on “cell-doctrine,” 22
central dogma of Crick, 222 , 223 , 255 , 261n12
Chemical Embryology (Needham), 47 –48 , 50 –51 , 62 , 71 , 81 , 93 , 117 , 142 , 143 , 147 , 202
chemical weapons, 27 –28 , 47 , 149 , 154 , 249 , 265n34
China, 45 –46 ; Joseph Needham’s interest in Chinese science and technology, 124 , 145 , 147 –50 ; Joseph
Needham’s visit to, 128 , 141 , 145 , 154 , 278n1
chlorine gas, use of in First World War. See chemical weapons
chreods, 218 , 289n29
chromatin, 77 –78 , 79 , 180
chromosomes, 144 , 180 , 253 ; discussed at the Theoretical Biology Club, 110 , 112 ; and DNA, 167 , 254 ;
Woodger on, 60 , 75 , 79 , 110
Churchill, Winston, 130
Church Socialist League, 49
Clifford, William Kingdon, 265n31
Cofman, Victor, 85 , 86 –87
Cold Spring Harbor Symposium on Quantitative Biology, 189 –93
Collins, Randall, 261n7
Columbia University, 3 –4 , 19 , 32 , 125 , 163 , 176 –77
“Comment on Bertallanfy [sic ]” (Waddington), 219 , 289nn25–26
Commoner, Barry, 169 –70 , 212 –13 , 224
communication: biological communication, 75 , 76 –77 , 237 ; new methods of, 161 ; symbolic
communication, 231 , 236 –37 , 292n5 . See also language of biology
Communism, 9 , 82 , 111 , 119 , 121 , 124 , 146 , 180 , 194
Conant, James, 129
“The Concept of Life: Operational Definition” (Cofman), 86 –87
“The ‘Concept of Organism’ and the Relation Between Embryology and Genetics” (Woodger), 74 –78 , 79 ,
161 , 269n50
“conventional wisdom of the dominant group” (COWDUNG). See COWDUNG
Cornford, Frances (Darwin), 108 , 113 , 114 , 276n1
Cornford, Francis Macdonald, 108 , 113 , 114 , 276n1
COWDUNG (“conventional wisdom of the dominant group”), 228 –29 , 231 , 238
Cox, Sidney, 102
Crew, Francis A. E., 140 , 178 , 278n36
Crick, Francis H. C., 6 , 13 , 222 , 227 , 237 , 287n22 ; central dogma of, 222 , 223 , 255 , 261n12 ; on the
epigenetic landscape, 208 –10 ; as a mechanist, 11 , 211 , 218 , 247 , 250 ; and molecular biology, 200 ,
209 , 223 , 237 –38 , 255 ; and Waddington, 200 , 206 –10 , 287n16 ; winning Nobel prize, 173 , 210 ;
work with Watson, 112 , 124 , 167 , 168 , 173 , 183 , 184
Critique of Judgment (Kant), 48
crossveinless (wing abnormality), 183
Crowther, James C., 128
Cuvier, Jean Léopold Nicolas “Georges,” 39 , 40
cytoplasmic inheritance, 214 , 255 , 256

Dale, Henry, 274n8

d’Aquili, Eugene G., 230 , 291n14
Darkness at Noon (Koestler), 82
Darlington, Cyril D., 128 , 271n25 , 286n41
Darwin, and After Darwin (Romanes and Morgan), 24
Darwin, Charles, 13 , 24 , 40 , 89 , 136 , 175 , 229 , 244 , 264n30 ; conferences celebrating, 189 –93 , 194 ,
195 , 196 , 197 , 198 , 286n34 ; Montgomery and, 16 , 17 , 18 , 19 ; Morgan and, 23 –24 , 25 , 37 ; on
value of his contributions, 193 , 285n19
Darwin Centenary at University of Chicago, 193 –99 , 285nn20–21 , 285n26
Darwinism, 29 , 134 , 137 , 156 , 183 , 215 ; struggle with Mendelism, 216 . See also neo-Darwinism
“Darwin’s Bulldog.” See Huxley, T. H.
“Debatable Issues of Genetics and Breeding” (VASKhNIL), 175 –76
De Beer, Gavin, 153 , 155 , 184 , 205 , 278n35
Delbrück, Max, 168 , 246
Demerec, Milislav, 189 –90
Democritean-Cartesianism, 150 , 165 , 166 , 228 , 229 , 231
deoxyribonucleic acid. See DNA
Department of Scientific and Industrial Research (DSIR), 102
Descartes, René, 5 , 24 , 62 , 142 , 228
determination, 205 , 206 ; embryonic determination, 148 ; hereditary determination, 132 –33 ; organic
determination, 76 ; self-determination, 4
development, 22 , 33 , 60 , 102 –3 , 123 , 126 , 172 , 206 , 209 , 255 , 275n20 , 281n2 (chap. 11) , 289n29 ;
and chemicals, 44 , 47 , 108 , 122 , 143 ; developmental change, 183 , 204 ; embryonic development, 42
, 76 , 78 –79 , 121 , 125 , 170 , 182 , 229 ; epigenetics-W interest in, 254 (see also epigenetics); and
evolution, 9 , 21 , 37 , 68 , 170 , 173 , 181 , 213 , 218 , 221 –22 ; and genes, 110 , 118 , 123 , 133 , 137 ,
173 , 184 , 186 –87 , 201 , 261n12 , 276n40 , 287n11 , 292n5 ; mosaic concept of, 21 ; organ
development (ontogenetics), 44 , 68 , 94 , 98 ; organismal development, 37 , 47 , 53 , 67 , 136 , 231 .
See also canalization
DeVries, Hugo, 191
Dewey, John, 288n14
Diocesan College (Bishops), 23
discontinuity phenomena, 221 –22
DNA, 9 , 13 , 167 , 169 , 173 , 211 , 256 ; DNA-centrism, 170 , 173 ; double-helical structure of DNA, 8 ,
167 , 207 ; early prediction of, 207 ; epigenetic-H mechanisms, 254 , 256 ; importance of to biology and
popular consciousness, 13 , 151 , 160 , 167 , 169 –70 , 214 ; Nanney on, 255 –56 ; and RNA, 223 ,
261n12 ; as a term used in research searches, 257 ; triple-helix model for DNA, 167 ; Watson and
Crick’s work on, 167 , 168 , 173 , 183 , 206 , 209
“DNA Modification Mechanisms and Gene Activity During Development” (Holliday and Pugh), 256
Dobzhansky, Theodosius, 118 , 176 , 179 , 180 , 191 –92 , 194 ; at Darwin Centenary at University of
Chicago, 285n21 ; and neo-Darwinism, 125 , 135 , 195 , 240 ; Waddington and, 182 , 193 , 292n12
“The Doctrine of Auta” (Morgan), 24
Driesch, Hans, 4 –6 , 22 , 29 , 43 –44 , 98 , 170 –71 , 237 ; attack on mechanism, 31 , 211 , 281n2 (chap.
11) ; concept of entelechy, 160 ; and Entwicklungsmechanik, 94 , 111 , 281n2 (chap. 11) ; “harmonious
equipotentiality,” 34 ; Montgomery and, 15 , 19 ; as a neo-vitalist, 51 , 52 ; as a vitalist, 13 , 32 –33 , 35
, 36 , 41 , 62 , 108 , 143 –44 , 147 , 237 ; Woodger and, 65 , 71 –72
Drosophila experiments: Caltech’s work with, 118 , 125 , 137 , 182 ; focus on D. pseudoobscura , 192 ;
Waddington’s work with, 108 , 136 , 151 , 171 , 173 , 182 –83 , 209 , 226
Du Bois-Reymond, Emil, 5 , 32 , 33
Duhem, Pierre, 243 , 248

Edman, Irwin, 32 , 33 , 35
eel migration, 36 –37
Einstein, Albert, 11 , 12 , 62 , 90 , 105 , 160 , 229 , 268n21 , 272n2
élan vital, 4 , 5 , 29 , 33 , 51 , 115 , 131 , 144 , 160
Elementary Morphology and Physiology for Medical Students: A Guide for the First Year and a Stepping-
Stone for the Second (Woodger), 57 –58 , 60
Elements of Experimental Embryology (Huxley and De Beer), unfavorable review of, 278n35
Elsasser, Walter, 246 , 249
embryology: attempt to develop research on at Cambridge University funded by Rockefeller Foundation,
117 –22 ; ban on using embryonic stem-cell lines, 251 ; and biochemistry, 119 , 141 , 184 , 202 ;
bridging gap between genetics and, 184 –85 , 209 ; Driesch’s experiments, 4 , 5 –6 ; experimental
embryology, 77 –78 , 94 , 116 , 121 –22 , 125 , 138 –39 , 172 , 200 , 204 , 232 , 278n35 ; manipulations
of causing changes in adult, 182 –83 ; mechanists on embryonic development, 42 ; Needham’s study of
chick development in egg, 47 ; patternedness occurring embryologically, 68 ; Waddington’s views on
division of genetics and embryology, 201
Embryology and Genetics (Morgan), 125 , 126
Emergent Evolution (Morgan), 26 , 90
emergentism and concept of emergence, 20 , 26 , 27 , 30 , 41 , 265n31
Emerson, Alfred, 194
empiricism, 33 , 53 , 62 , 239 ; logical empiricism, 112 , 290n4
Endeavor (journal), 134
Engels, Friedrich, 144
entelechy, 33 , 51 , 52 , 53 , 115 , 140 , 160 , 166 , 239
entropy, 91 , 144 , 161 , 162
Entwicklungsmechanik, 94 –95 , 103 , 111 , 118 , 155 , 170 , 172 , 281n2 (chap. 11)
environment, 172 , 178 , 233 , 245 ; and development, 133 –37 , 180 , 181 , 186 , 187 ; environmentalism
movement, 211 –12 , 232 ; environmental stress and pressures, 30 , 37 , 126 , 135 , 137 , 145 , 182 –83 ,
187 , 215 ; internal environment, 34 , 210 ; organism and environment, 63 , 68 , 77 , 123 , 192 , 204
Ephrussi, Boris, 184 , 255
“Epigenetic Control Systems” (Nanney), 255
epigenetics, 7 , 187 , 271n23 ; Aristotle as precursor of, 172 , 281n2 (chap. 11) ; changes in key terms used
 in research, 257 –58 ; coming from research on “third way” of looking at life, 251 ; Crick on, 200 , 206
–9 ; current research transitioning from epigenetic-W, 256 –59 ; determining which processes are
genetic vs. epigenetic, 255 ; different meanings for, 254 , 256 –59 ; E. O. Wilson ignoring, 231 ;
epigenetic landscape, rendering of, 138 , 147 , 148 , 184 , 185 , 186 , 208 –10 , 223 ; epigenetic path
(sequence of changes), 125 ; epigenetics and organicism as counterbalances to neo-Darwinism, 159 ;
epigenetics-H, 254 –55 , 257 ; epigenetics-W, 254 , 255 , 256 –59 ; foundational work in, 138 ; as a
growing field, 251 –59 ; importance of epigenetics, 200 ; is modern epigenetics organic?, 252 –59 ; at
the IUBS meetings, 222 –23 ; molecular biology as epigenetics-H, 255 ; preformation vs. epigenesis, 63
, 68 , 73 , 75 , 281n2 (chap. 11) ; prehistory of, 281n2 (chap. 11) ; representations of an epigenetic
landscape, 138 , 147 , 148 , 184 , 185 , 186 , 208 –10 , 223 ; research affected by federal funding shifts,
256 –57 ; role of organicism in creation of, 8 , 261n10 ; Theoretical Biology Club on topic of, 13 , 20 ,
172 ; twenty-first century epigenetics seeing life as gene-centered, 259 ; use of term instead of
experimental embryology, 172 ; and Waddington, 172 –73 , 250 , 254 , 255 , 259 , 294n4 ; ways
scientists studied, 183 –84
“The Epigenotype” (Waddington), 138 , 172 –73 , 232
Ernst Mach Society, 61
Escherichia coli , 136
Esposito, Maurizio, 261n10 , 288n1
ethics, 131 , 195 , 198
eugenics, 177 –78 , 199
evocation principle, 111 , 122 , 156 ; masked evocator, 122 , 201 . See also organizer effect
evolution, 272n34 ; “beanbag genetics” and evolutionary theory of Mayr, 189 –93 , 194 , 195 , 200 , 283n1 ,
284n12 ; cosmic process of, 90 ; cultural evolution, 199 ; and development, 9 , 21 , 37 , 68 , 170 , 173 ,
181 , 213 , 218 , 221 –22 ; evo-devo as a subfield, 257 ; gradualistic evolution, 136 ; interactions with
molecular and evolutionary biologists, 159 ; Mayr’s definition of, 221 ; neo-Darwin interpretation of,
173 –74 ; and phenotypes, 221 , 294n4 ; “A-rate” evolution of body size, 283n5 ; synthetic evolutionary
theory, 190 –93 ; Waddington on, 131 –38 , 187 –88
Evolution and Ethics (T. H. Huxley), 131
“Evolutionary Adaptation” (Waddington), 195 –96 , 197 , 285n26 , 285n30
Evolution of an Evolutionist (Waddington), 232

Fabian Essays in Socialism (Noel), 49

Fabians, 9 , 49 , 111
“Fallacy of Misplaced Concreteness,” 143 , 229
Faraday, Michael, 242
Fell, Honor Bridget, 102 , 118 , 121 , 274n15
FHW triumvirate. See Fisher, R. A.; Haldane, John Burdon Sanderson “J.B.S.”; Wright, Sewall G.
Fichte, Johann, 66
“finalist school,” 51 –52
First World War, 56 –57 , 127 , 212 , 249 ; organists during, 138 –39 ; use of chemical weapons, 27 –28 , 47
, 265n34
Fischberg, Mischa, 217
Fisher, R. A., 134 , 271n25 , 272n30 , 285n21 ; as a population geneticist, 190 , 191 , 192 , 208 , 284n12
The Fitness of the Environment (Henderson), 33 , 266n52
Fletcher, Walter, 103
Ford, Edmund B., 271n25
Ford, Gerald, 20
Ford Foundation, 217
Form and Function (Russell), 35 , 38 , 40
Foundation trilogy (Asimov), 168 , 169
Franklin, Rosalind, 124 , 167 , 275n30 , 281n12
“Free Will versus Determinism” (lecture by Bergson), 3 –4
Freud, Sigmund, 58 , 229
fruit flies. See Drosophila

Galen of Pergamon, 5
gappiness, concept of, 20
Gardener, Margaret Emilia, 114
“Geneticist’s Manifesto” (H. J. Muller and J. Huxley), 178
“General Systems Theory” (GST), 160 –63 , 165 , 218 –19 , 289n27 ; as anti-mechanistic, 160 , 162 –63
General Theory (Keynes), 234
genes: autocatalytic nature of, 184 ; and development, 118 , 123 , 133 , 137 , 173 , 184 , 186 –87 , 201 ,
261n12 , 276n40 , 287n11 , 292n5 ; and discrete traits, 133 ; every gene from a pre-existing gene,
261n12 ; Mendelian genes and inheritance, 68 , 190 , 196 ; Muller’s gene theory, 180 –81 ; one-
gene=one-enzyme, 184 ; and possibility of response, 135 ; “random walk” of genes and selection, 134
–35 ; role of, 125 –26 ; twenty-first century epigenetics seeing life as gene-centered, 259
“Genetic Assimilation of an Acquired Characteristic” (Waddington), 182
genetics: “beanbag genetics,” 174 , 189 –93 , 194 , 195 , 200 , 283n1 , 284n12 ; bridging gap between
genetics and embryology, 184 –85 , 209 ; determining which processes are genetic vs. epigenetic, 255 ;
genetic assimilation, 217 ; “Geneticists’ Manifesto” (H. J. Muller and J. Huxley), 178 ; population
genetics, 190 , 191 , 208 , 284n12 ; regulatory genetics, 122 ; Waddington’s views on division of
genetics and embryology, 201 ; working in potentialities, 137
Genetics (journal), 103
Genetics and the Origin of Species (Dobzhansky), 125
“Genetics and Twentieth-Century Darwinism” conference, 189 –93
genomes, 125 , 137 , 183 , 254 ; epigenome, 172 , 173 ; Human Genome Project (HGP), 252 –53
genotypes: cells having same not always expressing same phenotype, 255 ; changes in, 137 ; developing
into phenotypes, 9 , 138 , 187 –88 , 261n12 , 292n5 ; epigenotype, 138 ; translating the, 183 ;
Waddington creating epigenetics to study, 255 , 259 ; Waddington not seeing an understood connection
with phenotypes, 294n4
Ghost in the Machine (Koestler), 220 , 289n29
Gieryn, Thomas F., 265n37
Gilbert, Scott F., 13 , 242 , 257 , 263n21
Glisson, Francis, 47
Gödel, Escher, Bach (Hofstadter), 234 –38
Gödel, Kurt, 105
Goldschmidt, Richard, 103
Goldstein, Kurt, 41
Goodfield [Toulmin], June, 200 , 201 –2 , 203 , 293n35 ; and Waddington, 200 –201 , 205 –6 , 248 , 287n14
Goodwin, Brian, 221 , 222 , 242 , 263n21
GOSPLAN, 129
Gould, Stephen Jay, 54 , 240 –41 ; influence of Waddington on, 240 , 292n12
Graaf, Regnier de, 281n2 (chap. 11)
Grammar of Science (Pearson), 63
Gray, James, 103 , 120 , 139 , 274n12 , 278n35
The Great Amphibium (Needham), 50
Gregg, John R., 275n29
Grene, Marjorie, 198 , 214 , 222 , 232 , 291n23
Grüneberg, Hans, 184
GST. See “General Systems Theory” (GST)

Haber, Fritz, 27 , 47 , 249 ; winning Nobel prize, 27

Haldane, John Burdon Sanderson “J.B.S.,” 156 ; and Communism, 180 , 181 ; and evolution, 272n34 ; and
mathematical models, 134 , 192 , 284n12 ; as a population geneticist, 190 , 191 , 192 , 208 , 284n12 ;
and the Theoretical Biology Club, 113 , 114 ; and Waddington, 103 , 272n34
Haldane, John Scott “J.S.,” 19 , 27 –31 , 32 , 34 , 37 , 43 , 156 ; as an anti-mechanist, 30 , 38 , 39 , 64 , 85 ;
death of, 91 ; against D. W. Thompson, 105 ; as a first generation organicist, 52 , 138 , 165 , 229 , 239 ,
240 , 241 , 245 ; and Henderson, 34 , 35 ; and Montgomery, 34 –35 ; on regulatory systems, 20 , 161 ,
166 ; and Russell, 38 , 39 , 40 ; at SICHST, 82 , 85 , 87
Hall, Brian K., 242 , 257 , 263n21
Haller, Albrecht von, 281n2 (chap. 11)
Halsbury, Lord, 207 , 287n18
Hamburger, Viktor, 95 –96 , 97 , 174 , 193 , 241
Hardy, William Bate, 47
Harland, Sydney Cross, 271n25
“harmonious equipotentiality,” 33 , 34 , 98
Harrison, Ross, 184 , 206
Harvey, William, 5 , 47 , 281n2 (chap. 11)
Hayek, Friedrich von, 123
heat shock, use of by Waddington, 182
Heidegger, Martin, 214
Hein, Hilde, 247 , 248 , 262n15
Helmholtz, Hermann von, 5 , 16
Henderson, Lawrence J., 19 , 20 , 31 –35 , 165 , 245 , 266n52 , 266n54
Heraclitus, 228
Herder, Johann Gottfried von, 66
Hill, Archibald V., 129
Hill, James P., 57
History of Embryology (Needham), 172
Hitler, Adolf, 111
Ho, Mae-Wan, 257
Hodgkin, Dorothy Crowfoot, 112 , 116 , 123
Hoernlé, R. F. Alfred, 31
Hofstadter, Douglas, 234 –38 , 243 , 245 , 248 , 256
Hogben, Lancelot T., 82 , 157 , 278n36
holism, 9 , 30 , 90 , 230 , 237 , 273n17 , 293n35
Holism and Evolution (Smuts), 88 , 90 , 92 , 100
Holliday, Robin, 254 , 256
“holon” concept, 220 , 289n29
Holtfreter, Johannes, 95 , 97 , 112 , 122 , 147 , 202
Hopkins, Frederick Gowland “Hoppy,” 49 , 50 , 111 , 120 –21 , 124 , 146 –47 , 204 ; winning Nobel prize,
47
Human Genome Project (HGP), 252 –53
Hume, David, 33
Hunter, John, 69
Hunter, William, 5
Hutchinson, Evelyn, 100
Huxley, Aldous, 153 , 177
Huxley, Francis, 153 , 155
Huxley, Julian, 44 , 205 , 242 , 271n25 ; at Darwin Centenary at University of Chicago, 194 –95 , 198 , 199
; De Beer and, 153 , 155 , 278n35 ; and “Geneticists’ Manifesto,” 178 ; Joseph Needham and, 46 , 119
–20 , 145 –46 , 278n35 ; and London Eugenics Society, 286n41 ; Muller, and, 176 –78 , 179 –80 , 199 ;
synthesis model of neo-Darwinism, 7 , 10 ; and Tots & Quots, 128
Huxley, Thomas H., 17 , 29 , 44 , 131 , 288n6 ; Julian’s grandfather, 44 , 176 ; Lloyd Morgan and, 23 , 37 ;
and protoplasm, 67 , 90 , 281n2 (chap. 11)
hydras, 22 , 95 , 96

“Inbreeding and Linkage” (Haldane and Waddington), 272n34

Individuation Field, 143
induction, 87 , 95 , 156 , 205 ; embryonic induction, 201 ; experiments involving, 95 , 96 ; indirect
induction, 203 ; neural induction, 122 –23 ; vs. self-differentiation, 94
inorganic substances, 15 ; biochemical regulation of nonliving substances, 52 ; understanding difference
between organic and inorganic, 5 , 15 , 34 , 43 , 266n69 ; viewed by Bertalanffy’s GST, 162
Institute for Mathematico-Physico-Chemical Morphology (MPC), 119 , 134
Institute of Animal Genetics at University of Edinburgh, 140 , 151 –52
“Institute of Problems on the War” meeting, 278n1
The Intelligent Man’s Guide to Science (Asimov), 168 , 169 , 170 , 184 , 212 , 281n16
International Congress of Genetics: Sixth conference, 175 ; Seventh conference, 178
International Congress of Science and Technology. See Second International Congress of Science and
Technology (SICHST)
International Scientific Commission, 149
International Union of Biological Sciences (IUBS) symposia, 219 –24 , 227 , 236 –37 , 287n16 , 291n23 ,
292n5 , 292n12
Interpretation of Development and Heredity (Russell), 163 , 281nn6–7
Introduction to Comparative Psychology (Morgan), 25
Introduction to Mathematical Philosophy (Russell), 106
“Is Vitalism Dead?” (Crick’s desired title of a lecture), 206

Jablonka, Eva, 242 , 257

Jacob, François, 122
James, Walter, 12
Jaspers, Karl, 288n14
Jennings, Herbert S., 31 , 32
Jonas, Hans, 213 –14
Journal of Embryology and Experimental Morphology , 232
Journal of Genetics , 100 , 271n25
Journal of Vernalization , 175
Joyce, James, 84
JSTOR, 252 , 253
Justin. See White, Margaret Justin Blanco

Kaiser Wilhelm Institute for Biology, 95 , 103 , 177

Kammerer, Paul, 216
Kant, Immanuel, 4 , 34 , 48 , 53 , 71
Kingsley, Charles, 25
Koestler, Arthur, 82 , 214 –21 , 226 , 288n14 , 289n15 , 289n23 , 289n29
Kuhn, Thomas, 7 , 10 –11 , 242 –44 , 247 , 259 , 262n15 , 289n26
Kurtén, Björn, 190 , 283n5
Kut-al-Amara, siege of, 57

Lamarck, Jean-Baptiste, 134 , 136 , 175 , 180 , 216 , 218

Lamarckism, 9 , 134 –35 , 174 , 189 , 217 , 218 , 250 , 256
Lamb, Marion, 257
La Mettrie, Julian de, 42 , 43 , 81
language of biology, 60 , 86 , 151 , 153 ; applying mathematized, cybernetic terms to, 161 –62 ; logical
language for expressing classifications, 106 ; new language for “third way” biology, 75 , 223 –24 ;
symbolic language, 77 , 79 –80 ; using tools from philosophy of language, 292n5 . See also
communication
Laplace, Pierre-Simon, 89 , 162 , 169
Lascelles, Lass, 100
Laughlin, Charles, 230 , 291n14
Lederberg, Joshua, 253 –54 , 255 , 256 , 257 ; winning Nobel prize, 253
Leibniz, Gottfried Wilhelm von, 5 , 150
Lenin, Vladimir I., 82 , 144
Lenin All-Union Academy of Agricultural Sciences (VASKhNIL), 175 , 177 , 179 , 181
Levy, Hyman, 114 , 128 , 130
Lewes, George Henry, 265n31
Lewis, C. I., on the study of a priori , 72
Lewis, C. S., 155 , 277n14
Lewontin, Richard, 221 , 222 , 240 –41 , 292n12
Lillie, Ralph and Frank, 21
“Lillie’s paradox,” 269n58
living things. See organic life
Lloyd Morgan, Conwy, 19 –20 , 23 –27 , 33 , 37 , 52 , 90 , 229 , 241 ; and emergentism, 26 , 30 , 265n31 ;
and J.S. Haldane, 28 , 30 , 91 ; “Morgan’s canon,” 264n30 ; and Needham, 52 ; and Smuts, 246
Loeb, Jacques, 6 , 15 , 19 , 31 , 38 , 39 , 165 , 248 , 249 , 266n58 ; as a mechanist, 32 , 33 , 36 , 42 , 48 , 53 ,
72 , 81 , 142 , 160 , 237
logical empiricism, 112 , 290n4
London Eugenics Society, 286n41
Look to the Land (James), 12
Lorentz, Konrad, 277n19
Lowell Lectures, 290n6
Lu, Gwei-djen, 124 , 141 , 145 ; marrying Joseph Needham, 276n1
Ludwig, Carl, 5
Ludwig II (king of Bavaria), 16
Lysenko, Trofim Denisovich, 13 , 173 –76 , 177 , 178 , 179 , 180 , 188 , 250 , 263n18 ; use of vernalization,
174 –75 , 176 , 181
Lysenkoism, 174 , 176 , 189 , 250 , 263n18 ; responses to anti-Lysenkoism, 179 –81 ; Waddington agreeing
with tenets of, 181 –82 , 189

Mach, Ernst W. J. W., 63

Macy Cybernetics meetings, 161
Man a Machine (La Mettrie), 42 , 43
Man a Machine (Needham), 43 , 45 , 50 , 62 , 70
Mangold, Christian, 97 –98
Mangold, Hilde (Pröscholdt), 94 , 95 –98 , 111 , 112 , 174
Mangold, Otto, 93 , 95 , 97 , 103 , 111 , 137
Man Not a Machine (Rignano), 42 , 43 , 48 , 51
The Man Who Loved China (Winchester), 46
Marvin, Walter T., 31 , 32 –33 , 61 , 127
Marx, Karl, 144
The Mathematical Theory of Communication (Shannon and Weaver), 161
Matthiola incana , 100 , 271n25
Maxwell, James, 169
Maynard-Smith, John, 222 , 280n41 , 291n23
Mayr, Ernst, 173 –74 , 187 , 189 –93 , 284n11 ; accepting organicism, 193 , 285n17 ; and anti-Darwinism,
289n20 ; attack on population genetics, 190 , 191 –92 , 284n12 ; “beanbag genetics” and evolutionary
theory of, 190 –93 , 194 , 195 , 200 , 283n1 , 284n12 ; conference on modern synthesis, 222 , 290n35 ;
at Darwin Centenary at University of Chicago, 194 , 196 , 198 , 285nn20–21 ; and Waddington, 189 ,
192 –93 , 198 , 220 –22 , 284n9
McDougall, Walter, 70
Mead, Margaret, 132 , 133 , 139 , 195 , 198 , 278n1 , 286n34 ; Gregory Bateson and, 132 , 133 , 139 ,
278n1 , 285n20 ; not invited to Darwin Centenary at University of Chicago, 198 , 285n20 ; sponsoring
a symposium, 230 –31 ; and Waddington, 198 , 286n34
mechanism, 5 , 11 , 35 , 61 ; alternatives to, 53 , 66 , 240 , 245 ; anticipating simple answers for all organic
process, 239 ; APA panel debating concepts of vitalism and mechanism, 31 –33 , 35 ; benefits from, 212
; and biochemistry, 39 ; change of term to reductionism, 166 , 169 , 170 (see also reductionism); Crick
as a mechanist, 11 , 211 , 218 , 247 , 250 ; as gene-centric, 213 –14 , 222 , 227 , 247 , 259 , 288n1 ;
growth of after World War II, 170 –71 ; Joseph Needham on, 42 –55 , 142 –44 , 160 , 245 ; Koestler’s
views on, 214 –21 ; La Mettrie as a mechanist, 42 ; Loeb as a mechanist, 32 , 33 , 36 , 42 , 48 , 53 , 72 ,
81 , 142 , 160 , 237 ; as a major meta-theoretical commitment, 247 –48 , 259 ; mechanistic materialism,
25 ; metaphysical mechanism, 64 , 245 ; methodological mechanism, 64 , 65 , 71 , 245 ; Montgomery
and, 18 , 41 ; neo-mechanism, 53 –54 , 70 , 71 –72 , 117 ; organicism as alternative to mechanism and
vitalism, 6 –7 , 15 , 245 ; organic philosophy as a rejection of, 7 ; as part of a worldview, 44 ; portrayed
by Gödel, Escher, Bach (Hofstadter), 234 –38 , 243 ; re-entrenchment of after World War II, 13 ;
Rothschild’s representation of, 132 –33 ; Russell on vitalism-mechanism debate, 36 , 38 ; on simplicity
and randomness of species, 236 ; vs. vitalism, 6 , 51 , 64 , 142 , 160 , 165 , 245 –47 , 248 , 266n58 ;
Waddington on, 245 ; Woodger on, 64 –66 . See also anti-vitalism
Mechanism, Life, and Personality (Haldane), 30
“Mechanism” (Needham), 50
The Mechanistic Conception of Life (Loeb) [also a lecture], 19 , 266n58
“Mechanistic Explanation and Organismic Biology” (Nagel), 166
Medawar, Peter, 13 , 153 –56 , 241 , 242 , 280n41 ; and Crick, 210 , 211 , 250 ; finances of, 154 , 280n28 ;
and Goodfield, 200 , 202 , 248 ; immunosuppression and tissue grafting work of, 155 , 156 ; and Joseph
Needham, 154 , 156 , 158 , 202 –3 , 204 –5 , 287n11 ; on molecular biology, 203 , 204 ; not interested
in theory, 250 ; and Waddington, 60 , 156 , 158 , 200 , 204 –5 , 287n11 ; winning Nobel prize, 153 , 203
, 210 ; and Woodger, 153 , 154 , 155 –56 , 157 , 159 , 170 , 203 , 280n43
Medical Research Council Laboratory of Molecular Biology, 209
Mendelism, 10 , 134 , 217 ; and Lysenkoism, 175 , 176 , 177 , 263n18 ; Mendelian genes and inheritance,
68 , 190 , 196 ; Mendelian ratios, 271n25 ; Mendelians, 62 , 191 , 193 , 216
Menger, Karl, 105
metaphysics, 66 , 71 , 115 , 129 , 130 , 150 , 192 , 216 ; metaphysical epistemology, 61 , 63 ; metaphysical
mechanism, 64 , 245 ; metaphysical vitalism, 64
meta-theoretical commitments, 247 –48 , 249 , 259 , 262n15
methodological mechanism, 64 , 65 , 71 , 245 . See also neo-mechanism
methodological vitalists, 268n21
methylation, 9 , 254 , 257 , 258
Meyer-Abich, Adolf, 70
“Microcosm” (Bernal), 83 –84
Mind (journal), 19 , 72 –73 , 269n40
“Mindless Societies” (Waddington), 230 , 291n14
Ministry of Agriculture and Fisheries (MAF), 37
Misuse of Mind (Stephen), 131
Mitchison, N. Avrion, 155 , 156
Modern Synthesis, 118
Modern Theories of Development (Bertalanffy and Woodger), 112 , 160 , 163
Moffitt, John, 276n1
Molden (publisher), 217
molecular biology, 138 , 209 , 211 , 212 , 214 , 217 , 247 , 256 , 262n15 ; birth and growth of, 8 , 13 , 159 ,
166 , 171 , 173 , 225 , 228 , 274n2 ; Crick and, 6 , 200 , 209 , 223 , 237 –38 , 255 ; as epigenetics-H,
255 , 256 ; Medawar on, 203 , 204 ; and molecular biology, 6 ; Waddington on, 138 , 183 , 201 –2 , 205
–6 , 218 , 225 , 228
molecules, concept of applied to biology, 166 , 167 , 168 –70 , 182 ; master molecule, 166 , 167 , 227 , 256
monism, concept of, 24 , 26 , 27 , 30 , 265n31 ; analytic monism, 25 ; materialist monism, 63 , 64 ; three-
aspect monism, 265n31
Monist (journal), 19 , 24
Monod, Jacques, 122
Montgomery, Edmund Duncan, 15 –19 , 21 , 24 , 28 , 34 –35 , 41 , 90 , 161
Moore, George E., 72 , 73 , 269n40
Morgan, Thomas Hunt “T.H.,” 118 , 125 , 127 , 137 , 182 , 184 , 191 , 201 . See also Drosophila
experiments, Caltech’s work with
Morse, Edward S., 21
Mortonhall. See University of Edinburgh
mosaic concept of development, 21
Motz, Hans, 155 , 156
Moyle, Dorothy. See Needham, Dorothy Moyle
MPC. See Institute for Mathematico-Physico-Chemical Morphology
mu (Zen doctrine), 237 , 243 , 248
Müller, Gerd, 263n21
Muller, Hermann J., 140 , 184 , 187 , 258 ; and anti-Lysenkoism, 176 –81 ; at Darwin Centenary at
University of Chicago, 198 –99 ; gene theory of, 180 –81 , 183 –84 , 187 , 261n12 ; and “Geneticists’
Manifesto,” 178 ; as a Mendelian, 191 ; and Waddington, 173 , 182 ; winning Nobel prize, 140 , 183
–84
Munson, J. Ronald, 8
mustard gas, antidote for. See chemical weapons

Nagel, Ernest, 159 –60 , 166 –67 , 220 ; dismissal of organicism, 163 –66 , 170 , 173 ; and Russell, 40 , 163
–64 , 165 –66 , 281nn6–7
Nanney, David, 255 , 256
National Institute for Medical Research, 157 , 203
National Science Foundation (NSF) (U.S.), 205 , 244
Nature (journal), 23 , 103 , 112 , 151 , 168 ; and Medawar’s article, 203 , 204 –5 ; Waddington published in,
131 , 134 , 205 , 206
Naven (Bateson), 132
Needham, Dorothy Moyle, 45 , 49 –50 , 54 , 109 , 128 , 137 ; biochemical research of, 116 ; in China, 145 ;
continuing in biochemistry after Joseph left field, 141 ; death of, 276n1 ; as a Fellow of the Royal
Society, 147 ; Needham dedicating Order and Life to TBC members, 114 –15 ; organizer experiments
of Waddington and Needhams, 55 , 93 –94 , 98 –99 , 103 , 112 , 117 , 119 , 143 , 147 , 156 , 159 , 200
–202 , 204 , 255 ; politics of, 111 , 274n8 ; and the Theoretical Biology Club, 104 , 151
Needham, John Turberville, 5
Needham, Joseph, 163 , 232 , 233 , 240 , 241 , 272n2 , 272n34 ; on allegations of U.S. engaging in
biochemical warfare, 149 , 279n15 ; attempt to develop a Rockefeller Foundation research project at
Cambridge, 117 –22 ; and Bernal, 83 –85 ; and biochemistry, 46 , 47 , 49 , 71 , 116 , 118 , 119 , 122 ,
124 , 146 –47 ; and Bukharin, 82 –83 , 129 ; in China, 128 , 141 , 145 , 154 , 278n1 ; on Chinese
science and technology, 124 , 145 , 147 –50 , 152 ; co-writing Science in War (anonymously published),
128 ; dedicating Order and Life to TBC members, 114 –15 ; dropping work in field of biology, 118 ,
124 ; and E. S. Russell, 281n6 ; as a Fellow of the Royal Society, 147 ; and Goodfield, 202 , 248 ,
293n35 ; and Gould, 292n12 ; and Henderson, 266n52 ; and the “holon” concept, 289n29 ; and Julian
Huxley, 119 –20 , 145 –46 ; on lack of recognition of work, 205 , 287n11 ; leaving field of
biochemistry, 141 ; letter to Nature on behalf of De Beer and Huxley, 278n35 ; marrying Lu Gwei-djen,
276n1 ; on mechanism, 42 –55 , 142 –44 , 160 , 245 ; and Medawar, 154 , 156 , 158 , 202 –3 , 204 –5 ,
287n11 ; and Morgan, 52 ; and organicism, 52 –53 , 81 , 88 , 144 –45 , 150 , 229 , 239 ; on organic
philosophy, 124 , 141 –42 , 143 –44 , 150 , 276n1 ; organizer experiments of Waddington and
Needhams, 55 , 93 –94 , 98 –99 , 103 , 112 , 117 , 119 , 143 , 147 , 156 , 159 , 200 –202 , 204 , 255 ;
and the philosophy of biology, 72 , 73 , 144 , 278n35 ; politics of, 111 , 119 , 124 , 146 , 149 , 214 ,
274n8 ; questioning Spemann’s holism, 273n17 ; and reductionism, 254 ; and Rignano, 43 , 44 –45 , 48
, 50 , 51 –52 , 70 ; and Smuts, 90 ; on theoretical biology, 51 , 124 ; and the Theoretical Biology Club,
12 , 104 , 107 , 108 , 109 , 114 –15 , 128 , 151 , 249 ; and Tots & Quots group, 127 ; use of Whitehead
philosophy, 93 , 100 , 117 , 142 –43 , 154 ; on vitalism, 142 –44 , 160 ; and Waddington, 81 , 117 –18 ,
121 , 125 , 137 , 172 , 200 ; and Woodger, 53 , 56 , 60 , 61 –62 , 69 –75 , 80 , 87 , 118 , 250
“Needham Question,” 45 –46
Needham Research Institute, 46 , 149 , 276n1
negentropy, 162
neo-Darwinism, 189 ; as an alternative to mechanism and vitalism, 240 ; anti-Darwinism as obstacle to,
289n20 ; “classical” neo-Darwinism, 272n34 ; and cultural anthropologists, 286n34 ; disregarding
genes in a population, 294n4 ; Dobzhansky and, 125 , 135 , 195 , 240 ; epigenetics and organicism as
counterbalances to, 159 ; as gene-centric, 214 , 232 ; Huxley’s synthesis model of neo-Darwinism, 7 ,
10 ; interpretation of evolution, 173 –74 ; Maynard-Smith defending, 222 ; and Mayr, 189 –93 ; and
Mendelism, 10 ; neo-Darwinism disregarding genes in a population, 294n4 ; neo-Darwinism synthesis,
6 , 8 , 134 –35 , 159 , 173 –74 , 180 , 183 , 193 , 214 , 217 , 222 , 233 , 240 ; organismal change due to
mutations, 183 , 187 , 240 ; oversimplification of, 285n30 ; “random walk” of evolution, 215 ;
Waddington and, 137 , 196 , 198 , 221 , 291n23 , 294n4 . See also Darwin Centenary at University of
Chicago; Darwinism
neo-Kantianism, 32
neo-Lamarckism, 134 , 136 , 215 , 216
neo-mechanism, 53 –54 , 70 , 71 –72 , 117 . See also methodological mechanism
neo-vitalism, 51 , 52 ; Bergson as a neo-vitalist, 51 , 52 ; Driesch as a neo-vitalist, 51 , 52
Neumann, John von, 288n14
“new realism” movement, 32 –33
Newth, David R., 232
New York Review of Books , 229 –30
Ney, Elisabet, 15 , 16
Nineteenth Century (journal), 29
Nixon, Richard, 256
Nobel laureates, 6
Noel, Conrad R., 48 –49 , 83 , 124
nonliving substances. See inorganic substances
non-overlapping magisteria (NOMA), 54
Not in Our Genes (Lewontin, Rose, and Kamin), 241
nuclear weapons radiation, 275n20
Nye, Bill, 242

Odessa Breeding and Genetics Institute, 175

Of Molecules and Men (Crick), 11 , 237
“Of Molecules and Men” (Crick lecture), 206 , 207
On Growth and Form (Thompson), 107 , 153
“On the Molecular Structure of Chromosomes” (Wrinch), 112
“On the Nature of the Amphibian Organization Centre, Evocation By Some Further Chemical Compounds”
(Waddington), 122
On the Origins of Love and Hate (Suttie), 58 –59
On the Origin of Species (Darwin), 16 , 17 , 136 , 189 , 298 ; Darwin Centenary at University of Chicago
celebrating, 193 –99 , 285nn20–21 , 285n26
ontogenetics (organ development), 44 , 68 , 94 , 98
The Open Society and Its Enemies (Popper), 156
Order and Life (Needham), 55 , 114 –15 , 147 , 161 , 163 , 173 , 186 , 287n11
The Order of Nature (Henderson), 34
Oregon State University, 181
“organic homeostasis,” 33 –34
organicism, 12 ; as alternative to mechanism and vitalism, 6 –7 , 15 , 245 ; claim that an organism is
irreducible to the sum of its parts, 164 –65 ; clashing with molecular biology, 247 ; as a counterbalance
to neo-Darwinism, 159 ; cybernetic organicism, 163 (see also “General Systems Theory” [GST]);
difficulty of explaining, 11 –12 ; and empiricism, 226 ; first generation of organicists, 15 –41 ;
interpretation of in the 1960s, 211 –25 , 226 , 288n1 , 289n26 ; is modern epigenetics organic?, 252 –59
; Lewontin and Gould speaking for, 241 ; Mayr accepting organicism, 193 , 285n17 ; and molecular
biology, 262n15 ; Nagel’ dismissal of organicism, 163 –66 , 170 , 173 ; and Needham, 52 –53 , 81 , 88 ,
144 –45 , 150 , 229 , 239 ; organicism before the Theoretical Biology Club, 19 ; organicist anti-
reductionism, 289n27 ; origination of, 234 , 239 , 244 ; Polanyi’s organismic principles, 214 ; reasons
for decline in interest in, 234 –38 , 244 ; role of in creation of epigenetics, 8 , 261n10 ; seen by some as
a kind of vitalism, 166 , 262n15 , 281n11 ; tracing connections between older and newer proponents of,
240 , 292n12 ; Waddington’s Tools for Thought as an organicist’s manifesto, 227 –33 ; ways to combat
social problems, 230 –31 ; Woodger’s version of, 61 . See also organic philosophy
organic life: purposiveness of living things, 40 , 43 , 266n69 ; serial organization of, 67 ; Smuts on place of
the organism in science and philosophy, 90 ; understanding difference between organic and inorganic, 5
, 15 , 34 , 43 , 266n69 ; unity of the organism, 35 ; viewed by Bertalanffy’s GST, 162
organic philosophy, 7 –9 , 108 , 117 , 123 , 248 ; advocates of, 35 , 46 , 53 , 104 , 124 , 143 , 144 , 245 , 246
; attacks on, 157 , 163 –64 , 166 , 193 , 233 , 246 ; Bertalanffy on, 162 –63 ; epigenetics taking root
from, 174 (see also epigenetics); growth and development as, 102 ; Joseph Needham on, 124 , 141 –42
 , 143 –44 , 150 , 276n1 ; as a rejection of mechanism, 7 ; rise of, 13 , 244 ; Smuts on, 100 ; Theoretical
Biology Club on topic of, 111 , 118 , 123 , 141 , 241 , 262n17 ; Waddington on, 151 , 174 , 184 , 229 ,
232 ; Woodger on, 61 , 87 , 151 , 250 . See also organicism
Organisers and Genes (Waddington), 125 , 126 , 138 , 173 , 201 , 276n40 , 287n11 ; Piper’s frontispiece
for, 137 –38 , 147 , 184 , 185 , 223
Organism and Environment as Illustrated by the Physiology of Breathing (Haldane), 30 , 34
organizer effect, 93 –98 , 102 , 273n17 ; attempt to organize RF sponsored research project on at Cambridge
University, 117 –22 ; and belief in holism, 293n35 ; masked evocator as trigger, 201 ; organizer
experiments of Waddington and Needhams, 55 , 93 –94 , 98 –99 , 103 , 112 , 117 , 119 , 143 , 156 , 159
, 200 –202 , 204 , 255 ; as a proving ground for organic philosophy, 123 ; reception of by life scientists,
206 ; Theoretical Biology Club’s interest in, 111 –12 . See also Spemann, Hans
Orwell, George, 288n14
ostriches, callouses on, 135 , 136 –37
“Outline of General System Theory” (Bertalanffy), 160
Out of the Night (Muller), 177
Owen, Richard, 17
Oyama, Susan, 263n21

pangenesis, 17
panspermia, 287n22
paradigms: Kuhn’s model of paradigm shifts, 10 , 41 , 247 , 259 , 262n14 , 289n26 ; new paradigm, 10 , 219
, 226 , 247 , 248 –49
Pasteur, Louis, 13
Pattee, Howard H., 223 –24
Patterns of Culture (Benedict), 132
Pauling, Linus, 124 , 167 , 212
Pearl, Raymond, 62 , 74 –75 , 241 –42 , 248
Pearson, Karl, 63
Peters, Rudolph, 154
phenomenalism, 63
The Phenomenon of Life (Jonas), 214
phenotypes: cells having same not always expressing same genotype, 255 ; constancy of, 136 ; Crick on
phenotypes and DNA, 255 ; epigenotype, 138 ; and evolution, 221 , 294n4 ; genotypes developing into,
9 , 138 , 187 –88 , 261n12 , 292n5 ; impact on an organism’s phenotype and mutations, 254 ;
“punctuations” in, 240 ; ranges of possible phenotypes, 183 ; Waddington creating epigenetics to study,
255 , 259 ; Waddington not seeing an understood connection with genotypes, 294n4
Philosophical Problems in Light of Vital Organization (Montgomery), 19
philosophy of biology, 10 , 27 , 163 , 175 , 221 , 242 , 244 ; and Joseph Needham, 72 , 73 , 144 , 278n35 ;
and Medawar, 155 –56 , 157 , 203 ; vs. theoretical biology, 290n35 ; and Woodger, 72 , 73 , 75 , 203
philosophy of organism. See organicism
philosophy of science, 13 , 23 , 50 –51 , 59 , 115 , 158 , 159 , 166 , 170 , 234 , 241 –42 , 243
phyletic change, 241
“The Physical Basis of Life” (Huxley), 23
physics, 243 ; impact of on biology, 167 ; physicists interested in biology, 246
Piper, John, 137 –38 , 147 , 184 , 185 , 223
Polanyi, Michael, 214 , 243 , 246
Popper, Karl, 7 , 60 , 118 , 123 , 155 , 262n15 , 273n26 , 293n31 ; criticisms of, 208 , 247 , 287n22 ;
Popperian conjecture-refutation model, 10 ; and the Theoretical Biology Clubs, 112 –14 , 152 , 156 ;
and Woodger, 152 –53 , 249
population genetics, 190 , 191 –92 , 208 , 283n5 , 284n12
Poverty of Historicism (Popper), 114 , 273n26
Pravda (newspaper), 82 , 174
preformation, 79 , 177 , 183 , 204 , 207 ; vs. epigenesis, 63 , 68 , 73 , 75 , 281n2 (chap. 11)
Prezent, Isaak, 175 , 176 , 177 , 178
Priestley, Joseph, 242
Principia Biologæ (Woodger’s efforts to create), 56 –80 , 156
Principia Mathematica (Whitehead and Russell), 79 –80 , 105 , 123 , 151 , 153
The Problem of Knowledge: Philosophy, Science, and History Since Hegel (Cassirer), 160 , 163
Problems of Life and Mind (Lewes), 265n31
Proceedings of the Royal Society , 154
Pröscholdt, Hilde. See Mangold, Hilde (Pröscholdt)
protoplasm, 67 , 90 , 281n2 (chap. 11)
Przibram, Hans, 61 , 70 , 105
psychism, 25
psychobiology, 65 , 66 , 281n6
psychohistory, 169
Pugh, John E., 256
Pulitzer Prize, 234 , 237
punctuated equilibrium, 240
Punnett, Reginald C., 100 , 271n25

The Quarterly Journal of Microscopical Science , 17

Quarterly Review of Biology (journal), 51 , 74
Quine, Willard V. O., 118 , 152 , 247

Royal Air Force “RAF,” 128 , 139 , 157

Rashevsky, Nicolas, 118
“Recent Developments in the Philosophy of Biology” (Needham), 51 , 53 , 55 , 56 , 61 , 70
Redfield, Robert, 285n20
Redi, Francesco, 281n2 (chap. 11)
reductionism, 166 , 218 –19 , 226 , 248 , 254 ; American reductionism, 219 ; pre-reductionist Whiteheadian
thought, 230 ; reductionistic mechanism, 169 , 170 , 231 ; reductive mechanism inspired by Carson, 220
. See also mechanism
“Relations of Science, Technology, and Economics Under Capitalism” (Rubenstein), 129
Research Institute on General Embryology at Cambridge University, effort to found, 117 –22
Respiration (Haldane), 31
Richards, O.W., 135
Rignano, Eugenio, 42 –45 , 48 , 50 , 51 –52 , 70
Rise and Fall of the Third Reich (Shirer), 234
Ritter, William E., 35 , 42 , 69 , 229 , 239
river delta analogy describing an epigenetic landscape. See epigenetics: epigenetic landscape, rendering of
RNA, 214 , 223 , 261n12 ; epigenetic-H mechanisms, 254
Robertson, Alan, 231 –32
Robson, Guy C., 104 , 135 , 272n1
Rockefeller Foundation (RF), 110 , 177 , 219 –20 , 274n2 , 274n12 ; grant to Strangeways Laboratory for
Waddington research, 274n15 , 275n21 ; and Hopkins, 121 ; interest in Spemann’s organizer, 117 –22 ;
pulling back from working with Needham, 124 , 146 ; Waddington asking for personal financial help,
120
Romanes, George, 23 –24 , 29 , 37
Rothschild, Miriam, 132 –33 , 277n19
Rothschild, N. M. Victor, 129 , 132
Roughton, F. J. W., 51 –52
Roux, Wilhelm, 4 , 5 , 6 , 21 , 98 , 172 ; and Entwicklungsmechanik, 94 , 111 , 170 , 281n2 (chap. 11)
Royal Commissioners of the Exhibition, 120
Royal Society of London, 16 , 18 , 147 , 274n8 ; Fellows of, 45 , 157 , 267n6 ; financial assistance to
Joseph Needham, 121
Rubenstein, M., 129
Ruse, Michael, 275n29
Russell, Bertrand, 72 , 79 –80 , 105 –6 , 123 , 153 , 233 , 269n40 , 288n14 ; theory of types, 105 , 110 , 163
Russell, Edward Stuart, 19 –20 , 35 –40 , 65 , 70 , 266n58 , 266n69 ; as anti-mechanistic, 38 , 85 , 239 ; as a
first generation organicist, 41 , 52 , 82 , 91 , 229 , 241 , 245 ; Nagel’s critique of, 163 –64 , 165 ,
281nn6–7 ; and the Theoretical Biology Club, 104
Rutherford, Ernest, 90

Sarkar, Sahotra, 13 –14

Sartre, Jean-Paul, 288n14
Saunders, Edith R., 100 , 271n25
The Sceptical Biologist (Needham), 50 , 81
Sceptical Chymist (Boyle), 63
Schleiden, Mattias, 16 , 21
Schmalhausen, Ivan, 153
Schmidt, Karl, 196 –97 , 198
Schrödinger, Erwin, 167 –68 , 169 , 242
Schrödinger’s box, 144
Schwann, Theodor, 16 , 21
Science (journal) and anti-genetic thinking, 179
Science and Civilization in China (series by Needham), 46 , 147 –49 , 202 , 224
Science and Ethics (Waddington), 131 , 134 , 273n26 , 277n14
Science and Philosophy of the Organism (Driesch), 19
Science and Society (journal), 278n1
Science and the Modern World (Whitehead), 62
Science at the Cross Roads (International Congress of Science and Technology), 82 , 83
Science: Curse or Blessing? (Levy), 130
Science for the People (SftP), 229 , 231
Science in War (anonymously published), 128 , 130 , 139 , 140 , 278n1
The Scientific Attitude (Waddington), 130 –31
scientific materialism, 290n6
scientific method, 26 , 36 , 156 , 239 ; process of developing theories, 10 –11 ; use of rejection to define a
position, 7 , 261n7
Scientific Thought in the Twentieth Century, An Authoritative Account of Fifty Years (Medawar), 202
Scriven, Michael, 220
sea urchin experiements, 4 , 22 , 33 , 43 –44 , 170 –71
Second International Congress of Science and Technology (SICHST), 55 , 81 –83 , 85 –87 , 105 , 129
Second World War, 258 ; end of, 141 ; scientists during, 138 –40 , 161 , 167 , 184
self-differentiation, 94 , 95 , 98
Sex and Temperament (Mead), 132
SftP. See Science for the People (SftP)
Shannon, Claude E., 161
“The Shape of the Human Being as a Function of Time” (Medawar), 154
Shapiro, James A., 263n21
Shen, Shizhang, 124
SICHST. See Second International Congress of Science and Technology (SICHST)
Silent Spring (Carson), 211
Simath, Crescentia “Cencie,” 18
Simpson, George G. “G.G.,” 194 , 195 , 240 , 285n21 , 292n12
Skinner, B. F., 264n30
Slater, William K., 128
Smuts, Jan Christian, 12 , 88 –92 , 100 , 101 , 117 , 123 , 161 , 203 , 246
Snow, C. P., 225 , 242
Society for Experimental Biology, 173
Society for the Study of Evolution (SSE), 194
Sociobiology (Wilson), 229 –30 , 231
Socrates, 249 , 250
“Socrates.” See Woodger, Joseph Henry “Socrates”
Sonneborn, Tracy, 214 , 255
The Soviet Journal of Genetics , 177
Soviet science, state of, 289n15
Soviet State Planning Commission (GOSPLAN), 129
Spallanzani, Lazzaro, 5 , 281n2 (chap. 11)
“The Spandrels of San Marcos and the Panglossian Paradigm” (Gould and Lewontin), 240
The Spark (student publication), 177
Spemann, Hans, 93 , 94 –96 , 97 –98 , 111 , 157 , 204 , 206 ; winning Nobel prize, 94 , 97 , 98 , 111 . See
also organizer effect
Spencer, Herbert, 34
Spinoza, Baruch, 66
The Springs of Conduct (Morgan), 24
squid axons and synapses, 153
SSE. See Society for the Study of Evolution (SSE)
Stahl, Georg Ernst, 5
Stalin, Josef, 82 , 85 , 88 , 146 , 174 , 175 , 177 –79 , 214
Stark, Harold R., 129
Stebbing, Susan, 114
Stebbins, G. Ledyard, 190
stem-cells, 251 , 252 , 255
Stent, Gunther, 248
Stephen, Karin, 131 –32 , 133
Stralendorff, Carl Vicco Otto Friedrich Constantin von, 17 –18
Stralendorff, Margaret von, 18
Strand, August L., 181
Strangeways Laboratory, 99 , 102 –3 , 118 , 121 –22 , 139 ; Rockefeller Foundation grant for Waddington’s
research at, 121 , 274n15 ; Waddington’s work on effects of nuclear weapon radiation, 275n20
The Strategy of the Genes (Waddington), 147 , 184 –85 , 187 , 189 , 192 , 200 , 240
The Structure of Scientific Revolutions (Khun), 242 , 243
structure vs. function, 63 , 68
The Study of Living Things: Prolegomena to a Functional Biology (Russell), 38 , 39
Sturtevant, Alfred, 125 , 182
“Submicroscopic Structure of the Drosophila Egg” (Okada and Waddington), 294n4
superego as the basis of ethics, 131
Suttie, Ian Dishart, 60 , 72 , 73 ; Woodger and, 57 –59 , 62 –63 , 68 , 75 , 157 , 224 , 249
Swammerdam, Jan, 281n2 (chap. 11)
symbolic language, 77 , 79 –80

tadpoles, 94
Tarski, Alfred, 118 , 123 , 152 , 153
Tavistock Institute of Medical Psychology, 58
Tax, Sol, 194 –98 , 285nn20–21 , 285n25 , 286n38
TBC. See Theoretical Biology Club
Thaxted movement, 49 , 83
theoretical biology, 9 , 11 , 127 , 128 , 152 , 159 , 212 , 290n35 ; IUBS looking at, 219 , 220 , 223 –24 , 241
, 243 , 287n16 ; Joseph Needham on, 51 , 124 ; Waddington on, 134 , 140 , 141 , 222 , 250 , 290n35 ;
Woodger on, 60 , 112 , 128
Theoretical Biology Club (Second) at Oxford, 152 , 155 –56 , 157
Theoretical Biology Club (TBC), 8 , 35 , 104 –16 , 153 , 188 , 221 , 278n36 , 285n17 ; break up of, 117 –18
, 120 , 123 , 127 ; convening of, 104 , 128 , 249 ; Medawar corresponding with, 154 ; meeting place,
108 , 113 , 114 , 276n1 ; members of, 12 –13 , 104 –5 (see also Tots & Quots group); naming of, 12 ,
104 , 262n17 ; Needham dedicating Order and Life to TBC members, 114 –15 ; organicism before
TBC, 19 ; and Popper, 112 –14 , 152 , 156 ; relaunched after World War II by Woodger, 123 ; and the
“third way,” 15 , 20 , 60 , 251 ; on topic of epigenetics, 13 , 20 , 172 ; on topic of monism, 24 ; on topic
of new language of biology, 224 ; on topic of organic philosophy, 111 , 118 , 123 , 141 , 241 , 262n17 ;
on topic of Spemann’s organizer, 93 , 111 –12 ; Waddington only member continuing to pursue
theoretical biology, 141 , 151
Theorie von der Generation (Wolff), 281n2 (chap. 11)
“Theory and Practice from the Standpoint of Dialectical Materialism” (Bukharin), 82
theory of types, 105 –6 , 110 , 163
“third way” of looking at life, 6 –7 , 8 , 9 , 11 –12 , 13 –14 , 35 , 226 –27 ; in the 1930s, 226 ; after
Waddington, 234 –50 ; as an alternative to mechanism and vitalism, 240 ; early scientists preparing way
for, 15 –41 ; epigenetics coming from, 251 ; finding a “third way” around the mechanism-vitalism
debate, 161 ; interpretation of in the 1960s, 211 –25 , 226 , 288n1 , 289n26 ; new language for “third
way” biology, 75 , 223 –24 ; Russell’s contribution to, 36 –37 ; and the Theoretical Biology Club, 15 ,
20 , 60 , 251 ; Waddington on, 133 , 151 , 226 –27 . See also emergentism; epigenetics; monism,
concept of; organicism
Thom, René, 221 , 222 , 223
Thomas, Dylan, 288n14
Thompson, D’Arcy W., 105 , 107 , 128 , 153 , 154
Thomson, J. J., 90
Thouless, Robert, 130
“Three Aspects of Monism” (Morgan), 24
“Three Man Paper” (3MP), 168
Through the Looking Glass (Carroll), 234 –35
Time (magazine), 234
Times , 151
Timofeev-Resovskii, Nikolai Vladimirovich, 168 , 177
Tinbergen, Niko, 132 ; winning Nobel prize, 277n19
Tisdale, W. E., 117 , 119 , 120 , 121 , 124
Toivonen, Sulo, 122
Tolkien, J. R. R., 155
Tools for Thought (Waddington), 227 , 289n26
topological models of embryological differentiation rates, 108 , 273n9
Tots & Quots group, 127 –30 , 139 , 140 , 179 , 242 ; members also belonging to TBC, 127 , 128 , 153 , 157
, 278n36 . See also Theoretical Biology Club
traffic jam in New York City, 3 –4 , 261n3
Transactions of the Pathological Society of London , 17
Trembley, Abraham, 96 , 97
Triturus cristatus and Triturus taeniatus , 96 –97
Trotter, Louisa, 28
Truman, Harry S, 146
“Two Cultures” (sciences and humanties with divergent goals), 242 , 248
Two Cultures (Snow), 225
“Two Evolutionary Theories” (Grene), 214

“Über Induction von Embryonalanlagen durch Implantation artfremder Organisatoren” [“Induction of

Embryonic Primordia by Implantation of Organizers from a Different Species”] (Spemann and
Mangold), 97
Uexküll, Jakob Johann Von, 41 , 160
umwelt , 160
UNESCO. See United Nations Educational, Scientific and Cultural Organization (UNESCO)
unified field theory, 105 , 272n2
The Unitary Principle in Physics and Biology (Whyte), 272n2
United Nations, 88 , 145 –46
United Nations Educational, Scientific and Cultural Organization (UNESCO), 46 , 145 –46 , 149 , 279n15
“The Unity of the Organic Individual” (Montgomery), 18 , 161
The Unity of the Organism; or, the Organismal Conception of Life (Ritter), 35 , 69
University of Chicago, 21 ; Darwin Centenary, 193 –99 , 285nn20–21 , 285n26
University of Edinburgh, 140 , 150 –52 , 182 , 231 ; Mortonhall, 151 –52 , 173
“The Utilization of Scientists” (Needham), 278n1

Vandenberg, Hoyt, 146

The Variation of Animals and Plants Under Domestication (Darwin), 17
The Variation of Animals in Nature (Robson and Richards), 135
Vavilov, Nikolai Ivanovich, 173 , 175 –77 , 178 –80
vernalization, 174 –75 , 176 , 181
Villa Serbelloni (site of International Union of Biological Sciences meetings), 220 –22
Virchow, Rudolf, 16 , 108 , 176
vitalism, 4 –5 , 27 ; alternatives to, 240 , 245 ; anti-mechanistic positions, 66 , 217 –20 ; APA panel
debating concepts of vitalism and mechanism, 31 –33 , 35 ; Bergson as a vitalist, 6 , 12 , 13 , 15 , 35 ,
41 , 65 , 108 , 132 , 143 –44 ; concept of élan vital, 4 –5 , 29 , 51 , 131 , 160 ; Crick’s arguments against
vitalism and epigenetics, 206 –9 ; Driesch as a vitalist, 13 , 32 –33 , 35 , 36 , 41 , 62 , 108 , 143 –44 ,
147 , 237 ; “finalist school,” 51 –52 ; Haldane questioning concept of, 29 –30 ; having untested
concepts, 160 ; Joseph Needham on, 142 –44 , 160 ; as a major meta-theoretical commitment, 247 –48 ,
259 ; vs. mechanism, 6 , 51 , 64 , 142 , 160 , 165 , 245 –47 , 248 , 266n58 ; metaphysical vitalism, 64 ;
methodological vitalists, 268n21 ; mysteries of development supporting, 44 ; neo-Darwinism as an
alternative to mechanism and vitalism, 240 ; neo-vitalism, 51 , 52 ; “new physical laws” group of, 66 ;
organicism as alternative to mechanism and vitalism, 6 –7 , 15 , 245 ; organicism seen by some as a
kind of vitalism, 166 , 262n15 , 281n11 ; psychobiologists, 66 ; quasi-vitalistic rejection of mechanism,
212 ; Rignano as a vitalist, 42 –45 , 48 , 50 , 51 –52 ; Russell on vitalism-mechanism debate, 36 , 38 ;
some seeing organicism as a kind of, 166 , 281n11 ; Stephen’s representation of, 131 –32 ; Woodger’s
dismissdal of, 65 –66 . See also anti-mechanism
“Vitalism” (Haldane), 29 , 30 , 37
“Vitalism” (Russell), 35 –36 , 37
“Vivarium.” See Biologische Versuchsanstalt [Biological Research Institute], Vienna
Vogt, Karl, 16

Waddington, Conrad H. “Wad,” 99 –103 , 109 , 163 , 271n25 , 273n26 , 281n2 (chap. 11) ; at the
Agricultural Research Council, 139 , 140 ; and the Alpbach Symposium, 217 –20 , 226 , 289n23 ,
289nn25–27 ; assembling of his most definitive works as a legacy, 232 ; and Bateson, 100 –101 , 125 ,
132 , 133 , 233 , 271n23 , 276n40 ; on biochemistry, 99 , 122 ; bridging gap between genetics and
embryology, 184 –85 , 209 ; at Caltech, 139 , 182 , 201 ; and Cambridge University, 118 , 120 –21 , 127
; on canalization and acquired characteristics, 134 , 135 –38 , 162 , 181 –82 , 183 , 232 ; COWDUNG
concept applied to science, 228 –29 ; co-writing Science in War (anonymously published), 128 , 130 ,
139 , 140 , 278n1 ; and Crick, 200 , 206 –10 , 287n16 ; C.S. Lewis caricaturing, 277n14 ; at Darwin
Centenary at University of Chicago, 194 –95 , 285n26 , 285n30 , 286n34 ; death of, 193 ; on
development, 102 –3 , 108 , 123 , 125 , 133 , 136 –37 , 138 , 172 –73 , 182 , 185 –87 , 201 , 209 , 221
–22 , 231 , 275n20 , 276n11 , 292n5 ; on division of genetics and embryology, 201 ; and Dobzhansky,
182 , 193 , 292n12 ; and the epigenetic landscape, 138 , 147 , 148 , 184 , 185 , 186 , 208 –10 , 223 ; and
epigenetics, 172 –73 , 200 , 250 , 254 , 255 , 259 , 294n4 ; on evolution, 131 –38 , 187 –88 ;
experiments on Drosophila , 171 , 173 , 182 –83 , 226 , 294n4 ; and Goodfield, 200 –201 , 205 –6 , 248
, 287n14 , 293n35 ; and Gray, 120 , 274n12 ; health problems of, 224 , 230 ; on the importance of
human arts, 225 ; incorporating science, philosophy, and ethics, 130 –34 ; influence on Gould, 240 ,
292n12 ; interacting with molecular and evolutionary biologists, 159 ; isolation from other scientists
and colleagues, 229 ; and the IUBS, 219 –20 , 236 , 241 , 243 , 292n5 ; and J.B.S. Haldane, 103 ,
272n34 ; joining Tots & Quots group, 127 ; and Joseph Needham, 81 , 117 –18 , 121 , 125 , 137 , 172 ,
200 ; Joseph Needham dedicating Order and Life to TBC members, 114 –15 ; lack of recognition of
work of, 205 , 231 –32 , 287n11 ; Lysenkoism, Waddington agreeing with tenets of, 181 –82 , 189 ;
marriage to Margaret Justin Blanco White, 137 , 140 , 151 ; and Mayr, 189 , 192 –93 , 198 , 220 –22 ;
and Mead, 198 , 286n34 ; on mechanism, 245 ; and Medawar, 200 , 204 –5 ; on molecular biology, 138
, 183 , 201 –2 , 205 –6 , 218 , 225 , 228 ; and neo-Darwinism, 137 , 196 , 198 , 221 , 272n34 , 291n23 ,
294n4 ; on nuclear weapon radiation effects, 275n20 ; on organic philosophy, 151 , 174 , 184 , 229 , 232
; organizer experiments of Waddington and Needhams, 55 , 93 –94 , 98 –99 , 103 , 112 , 117 , 119 , 143
, 156 , 159 , 200 –202 , 204 , 255 ; on philosophy of language, 292n5 ; politics of, 111 ; questioning
Spemann’s vitalism, 273n17 ; on the “quick (scientific) buck,” 244 ; receiving honorary degree from
University of Geneva, 217 , 218 ; and reductionism, 254 ; reviews by in New York Review of Books ,
229 –30 , 231 , 291n14 ; and the Rockefeller Foundation, 117 , 120 , 274n15 , 275n21 ; and Simpson,
292n12 ; sociopolitical predicaments, study of, 227 ; on Spemann’s organizer, 112 , 119 ; and
Strangeways Laboratory, 121 , 274n15 , 275n20 ; and theoretical biology, 134 , 140 , 141 , 222 , 250 ,
290n35 ; and the Theoretical Biology Club, 12 –13 , 104 , 107 , 110 , 114 , 128 , 251 ; on the “third
way,” 133 , 151 , 226 –27 ; “third way” after Waddington, 234 –50 ; topological models of
embryological differentiation rates, 108 ; at the University of Edinburgh, 140 , 150 –52 , 173 , 182 ; use
of Whitehead philosophy, 24 , 60 , 101 –2 , 120 , 128 , 137 , 184 , 192 , 220 , 225 , 228 , 233 , 242 ,
290n6 , 291n14 ; on vitalism and mechanism, 245 ; wanting to reform mechanism, 245 ; work in 1938–
1939 in the U.S., 124 –26 ; work with the RAF, 139 , 140 , 157
Wang, Yinglai, 124
Warren, Howard C., 31 , 32
Washington Anthropological Society, 198
Waterbabies (Kingsley), 25
Watson, James “Jim,” 207 , 249 , 281n12 ; winning Nobel prize, 173 ; work with Crick, 112 , 124 , 167 ,
168 , 173 , 183 , 184
Weaver, Warren, 110 , 119 , 121 , 122 , 129 , 274n2 , 274n8 ; co-author of The Mathematical Theory of
Communication , 161
Weinberg, Robert A., 257
Weismann, August, 77 –78 , 222
Weiss, Paul, 206 , 217 , 248
Weldon, T. D. “Harry,” 153
Wells, H. G., 129
West-Eberhard, Mary-Jane, 242 , 263n21
WFSW. See World Federation of Scientific Workers (WFSW)
What Is Life? (Schrödinger), 167
What Mad Pursuit (Crick), 227
“Where Are We?” (Mayr), 190 –93
Whewell, William, 242 , 244
White, Hayden, 13 , 263n19
White, Margaret Justin Blanco, 137 , 140 , 151
White, R. G., 140
Whitehead, Alfred North, 12 , 24 , 60 , 81 , 100 , 105 , 161 , 166 , 188 , 229 ; co-author of Principia
Mathematica , 79 , 80 , 123 , 153 ; concept of canalization, 137 , 181 –82 ; impact of on Koestler, 217 ,
220 ; Needham’s use of Whitehead philosophy, 93 , 117 , 142 –43 , 154 ; process philosophy of, 151 ;
on replicability, 246 ; stating that physics was organic, 165 ; Waddington’s use of Whitehead
philosophy, 24 , 60 , 101 –2 , 120 , 137 , 192 , 220 , 228 , 233 , 242 , 290n6 , 291n14 ; Woodger’s use of
Whitehead philosophy, 62 , 72 , 87 , 123 , 153 , 249
Whitman, Charles Otis, 19 , 20 –22 , 28 , 41 , 52
Whyte, Lancelot L. “Lana,” 85 , 105 , 116 , 249 –50 , 272n2 ; joining Tots & Quots, 127 ; and the
Theoretical Biology Club, 105 , 123 , 249 –50
Wiesner, B. P. (Needham dedicating Order and Life to TBC members), 114 –15
Wilkins, Maurice, 124
William Marsh Rice Institute, 176
Wilson, E. B. (Edmund Beecher), 34 , 64 –65 , 231
Wilson, E. O. (Edward Osborne), 229 –30 , 235 –37 , 238 , 242 , 243 , 291n14
Winant, John Gilbert, 129
Winchester, Simon, 46
Winge, Øjvind, 271n25
Wittgenstein, Ludwig, 72 –73 , 130 –31 , 156 , 233 , 269n40 , 292n5
Wolff, Caspar Friedrich, 281n2 (chap. 11)
Woodger, Joseph Henry “Socrates,” 55 , 56 –80 , 99 , 156 , 163 , 233 , 269n50 , 272n2 ; after TBC working
more with philosophy, 118 , 123 , 152 , 153 ; on anti-mechanism, 61 –62 , 63 , 64 –66 , 86 , 160 ,
268n22 ; biologists opinions on work of, 123 , 275n29 ; and cell lineage studies, 209 ; on development,
60 , 67 –68 , 78 –79 ; and E. S. Russell, 281n6 ; impact of World War I on, 127 ; introducing Bauplan
notion, 128 ; and logical language for expressing classifications, 106 ; on mechanism, 64 –66 ; and
Medawar, 153 , 154 , 155 –56 , 157 , 159 , 170 , 203 , 280n43 ; and Needham, 53 , 56 , 60 –62 , 69 –74
, 75 , 80 , 87 , 118 , 250 ; Needham dedicating Order and Life to TBC members, 114 –15 ; and
organicism, 229 ; on organic philosophy, 61 , 87 , 151 , 250 ; and Pearl, 241 –42 ; politics of, 111 ;
Popper writing obituary for, 152 –53 ; and reductionism, 254 ; and second iteration of Theoretical
 Biology Club, 155 –56 ; at SICHST, 85 –87 ; and Suttie, 57 –59 , 62 , 63 , 68 , 75 , 157 , 224 , 249 ; and
the Theoretical Biology Club, 13 , 104 –5 , 107 , 108 , 109 , 114 , 151 , 186 , 249 –50 ; on theoretical
sloppiness, 246 ; use of Whitehead philosophy, 62 , 87 , 100 , 123 , 153 , 249 ; on vitalism, 65 –66
“Woodgery.” See Theoretical Biology Club
Woods Hole Oceanographic Institution (Marine Biological Laboratory), 20 , 21 , 124
The World, the Flesh, and the Devil (Bernal), 85
World Federation of Scientific Workers (WFSW), 146
World War I. See First World War
World War II. See Second World War
Wright, Sewall G., 6 , 135 , 285n21 ; as a population geneticist, 190 , 191 , 192 , 284n12
Wrinch, Dorothy Maud “Dot,” 12 , 105 –6 , 111 , 116 , 123 , 128 , 167 ; and Franklin, 275n30 ; moving to
the U.S., 118 , 124 , 152 ; Needham dedicating Order and Life to TBC members, 114 –15 ; and the
Theoretical Biology Club, 108 , 109 , 112 , 186 , 221 , 249 –50
Wrinch, John, 105

Young, John Zachary, 128 , 153 , 154 , 155 , 156 , 157

Ypres, First Battle of, 27 , 265n34

Zavadovsky, Boris, 85 –86

Zeeman, Erik Christopher, 222
Zen Buddhism and Hofstadter, 237 –38 , 248
Zimmer, Karl, 168
Zuckerman, Solly, 128 –29 , 157 , 241 , 242 ; co-writing Science in War (anonymously published), 130 ,
139 , 278n1