GENES, RADIATION,

AND SOCIETY
The L'ife and W ork
of H. J. Nluller
I '

!
I
EiofAxeI Carlson

Cornell University Press

H. J. MuHe:-, 1940 (photo by Hans Reichenbach) I thaca and London
 Copyright © 1981 by cornen University Press Dedicated, with love
All rights reserved. Except for brief quotations in a review, this book, or parts and admiration, to
thereof, must not be reproduced in any form without permission in writing from
the publisher. For information address Cornell University Press, 124 Roberts NEDRA CARLSON
Place, Ithaca, New York 14850.

First published 1981 by cornen University Press.

Published in the United Kingdom by cornen University Press Ltd.,
Ely House, 37 Dover Street, London, W1X 4HQ.

International Standard Book Number 0-8014-13°4-4

Library of Congress Catalog Card Number 81-5486
Printed in the United States of America
Librarians: Library of Congress cataloging information appears
on the last page uf the book.

5Tb. 012 y
r
MAK- C K - ! N STiTUT
FOR WISfHH\l8CHAfTaeeGCHICHTE
Bibli oth ek
 Illustrations
Kirkwood Hall, Indiana University
Members of the zoology department, Indiana
,, ' University, 1949 293
Helen Muller and her parents 30 9
Muller receives the Nobel Prize 314
The Drosophila laboratory course, 1957 345
A genetic scheme 38 5 Preface
Thea and H. J. Muller 41 9

FIGURES
1. Drosophila melanogaster 48
2. X-linked inheritance 56 H . J. Muller is best known as the recipient of a Nobel Prize
3. Crossing over 60 for being the first to show that' mutations can occur in living or-
4. Crossing over and mapping 65 ganisms after exposure to X rays. He was also a controversial fig-
5. The sum rule and linear mapping ure in both the public and the private worlds of science. It was
72
6. Interference and coincidence his mixed blessing to be widely admired, honored, despised, mis-
73
7. Nondisjunction trusted, and misrepresented.
84
Throughout this biography I have been guided by the belief
8. Chief genes and modifiers 95 that Muller's life was symbolic of the human plight in the twen-
9. Balanced lethals 97 tieth century. He placed his faith in rationalism; he fought for
10. The origin of mosaics 100
his ever changing political beliefs; he erred in viewing the Soviet
11. The method for detecting lethals 111 experiment under Stalin as the hope of the future ; he believed
12. The ClB method for detecting lethals 118 in educating the public ab out the dangers of misapplied science;
13. The X-linked visible test 147 and he advocated the use of more science and the conscientious
14. The genetic test for translocations 157 appraisal of values to remedy the failures of contemporary society.
15. Dosage compensation 172 Muller remained throughout life an idealist. He never abandoned
16. The left-right test 201 an inner faith in the capacity of humans to solve their own prob-
17. Bar eyes-a position effect duplication 211 lems. He learned to value individual freedom, extending it from
18. The breakage-fusion-bridge cyde 348 his own advocacy of academic and scientific freedom to encom-
Al. Mendel's principle of segregation pass every aspect of human liberty.
439
Az. Mendel' s principle of independent assortment Muller was frequently buffe ted by personal failure. For most
440
of his life (until1946) he lived on a modest income, unable to save
A3. Mitosis 44 1 or plan financially for his long-range future. He was subject to
A4. Meiosis 44 2 an insecurity that welled up in a competitive, often polemically
A5. The chromosomal basis of Mendelism 444 expressed and self-defeating des ire to prove the priority of his
ideas and experiments in establishing the principles of genetics .
Too often he was victimized by colleagues who freely borrowed
x
xi
 Preface
his ideas or imposed on his time and judgment, and he usually
remained silent to avoid the old charge of being obsessed with
"priorities." He suffered the political consequences of his mistaken
faith in communism, both in the USSR and in the United States.
11 In this book I have quoted frequently from Muller' s letters and
articles, not only to convey his habits of thinking and personal-
ity, but to present his lue as if he were writing his own account
of it. I have not ignored Muller's personal inadequacies and mis-
takes; but as his student I am also aware that I cannot divest
mys elf of the unconscious bias and vie'''Points I absorbed from
hirn . I have deliberately not intended this biography as a critical
review and evaluation of his life and work. Rather, I hope it is
an accurate account of wh at he did, what he believed in, and what
his values were.
I began my research on Muller's life in 1966, after I had com-
pIe ted my book The Gene: A Gritical Histo1'Y, which attempted
to follow the successes and blind alleys of the development of the
,,
modern concept of the gene. Muller agreed to help me, and he
ii asked his dosest friend, Edgar Alten burg, to send me the letters
I he had written to hirn over a span of fifty years. He deferred a
personal interview until he could arrange for his children and
I
grands on to be present for a detailed account of his early years.
,'i.
This meeting never took place because of his progressivei)' deteri-
orating health. I did, however, hold a long series of interviews and
conversations with Edgar Altenburg, Loulin Altenburg Browning,
C. P. Oliver, W . S. Stone, T. S. Painter, A. C. Faberge, A. H .
Sturtevant, Max Delbrück, Linus Pauling, Jack Schultz, Alexander
Weinstein, Charlotte Auerbach, Guido Pontecorvo, J. S. Huxley,
Peo Koller, P. T. Ives, H. H. Plough, Oscar Schotte, Willard
Libby, Raissa Berg, Zhores Medvedev, I. I. Oster, A. P. Schalet,
Fritz Sobels, K. G. Lünning, Bronson Price, Roselee Raffel,
Herman B WeHs, Bentley Glass, Paul Klinge, T. M. Sonneborn,
Oscar Riddle Fernandus Pavne, Salvador Luria, Carl Sagan, and
Edward and Martha Baylor. Most ofthese persons admired Muller,
but some disliked hirn. I have tried to be accurate in paraphrasing
their views, and, of course, Iassume full responsibility for any
errors I have made in representing them.
Alter Muller's death I spent two summers organizing his papers,
xii
Preface
and I took notes on his unpublished manuscripts, research data
books, and the notebooks and writings of his early years. When
his papers were transferred to the LilIy Library at Indiana Uni-
versity, I began reading the thirty thousand letters in the coHec-
tion. These represent, in my estimation, about 80 per cent of his
total correspondence and virtually all of it since 1945. I tried with-
out success to locate many of the letters from his Texas years
(1915-1932). Fortunately, Altenburg and Huxley had a consider-
able number of Muller' s handwritten letters to them from this
period, and these helped supply some of the details of this for-
mative stage in his life. The LilIy Library provided solitude and
direct access to the Muller papers, and over the course of five
years I amassed an immense quantity of notes. I am grateful to
David RandalI, Josiah Q. Bennett, Elfrieda Lang, and the Lilly
Library staff for their help in making these facilities so comfort-
able for scholarly work.
No one has been of more helpin trus project than Thea Muller.
Her numerous recollections and her advice over the past several
years have been invaluable in providing the background to inci-
dents and personalities in her husband' s life that would otherwise
have remained murky or unknown. She read every chapter with
great care, and her corrections were most useful. She organized
the chronological file of correspondence during the semesters
while I was teaching so that I could rapidly and efficiently use
it when I returned each summer. Putting it in order involved a
good deal of work because Muller never retired or took the time
to work on his own papers. He kept virtually everything, but
. he lacked, until late in his lue, the secretarial help to file or other-
wise arrange his many writings.
I appreciate the help of Linda Edmunds, who typed most of
the first draft and who helped me organize my notes. I am grate-
ful to my wife, N edra, for typing the final draft as rapidly as I
brought the chapters to her.
I have benefited from the very useful suggestions of those who
read the manuscript, especiaHy Ruth Cowan, Leonard and Susan
Firestone, A. Peter Gary, and William Provine.
The drawings were prepared (unless it is otherwise indicated
in the legends) by Joyce Schirmer, staff artist for the Division of
xiii
 Preface
Biological Sciences, State University of New York at Stony Brook.
l thank Christina and John Carlson for their help in preparing the
index and my students Richard Levy, Dan Ciccarone, Gregory
Jay, and Tracy Meyers for reading the proofs with me.
l also thank the National Science Foundation for fun ding por-
tions of the work involved in writing this biography, particularly
for making possible a trip to Europe and several trips in the United
G ENES , RADIATION ,
States to interview colleagues and contemporaries of H. J. Muller. AN D SOCIETY
ELOFAxEL CARLSON The Life and Work
Setauket, NewYork of H . J. Muller

I
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xiv

Human Mutations and
the Radiation Danger
The award of the Nobel Prize in Physiology and Medicine,
coming so soon after the Hiroshima and Nagasaki bombings, gave
Muller the chance to voice his concern about the genetic hazards of
radiation more actively. The public' s major anxiety concerned
atomic warfare; little attention was paid to the genetic damage
from medical uses of radiation, which, perhaps through familiarity,
seemed both necessary and harmless to both the medical practi-
tioner and the patient. M uller objected to this uncritical endorse-
ment of the use of X rays, claiming that "medical men have got to
accept the result, which they have hitherto been so reluctant to
face, that mutations are produced by exposure of the reproductive
organs to X rays, and that they, their technicians, and their pa-
tients must, therefore, be better protected than they ordinarily are
if we would avoid contaminating the biological basis of humanity,
thus doing more harm to future generations than the good we do' to
this generation. "1
Muller did not challenge the need for diagnostic X in
determining illness; nor did he wish radiation therapy to be aban-
doned for treating malignant tissue or serious disorders. He ob-
jected to a large number of practices that seemed less compelling,
such as the failure to shield the gonads while performing diagnostic
X rays; the routine use of X-ray photographs of fetuses to detect
potential difficulties in delivery; the use of radiation to straighten
out bow legs in children by slowing bone growth; the application of
'H. J. Muller to Robert Campbell, March 19, 1947, Lilly Library.
33 6
Human Mutations and the Radiation Danger
radiation to warts and nonmalignant infections that could be treat-
ed by other means; the radiation of the gonads to sterilize, tem-
porarily, an individual whose medical or psychological problems
made pregnancy unwise; or even the reverse, using X rays to
stimulate ovulation in infertile females . This last procedure, de-
vised and employed extensivelyon several hundred women for
over twenty years by Dr. Ira 1. Kaplan, a fertility specialist, in-
volved three weekly applications of X rays to the pituitary gland
and the ovaries. Z Kaplan administered a total dose of 175 roent-
gens, but other radiologists used as much as 300 roentgens . Kaplan
claimed that neither the children nor the grandchildren arising
from such treated ovaries showed any monstrosities or evidence of
mutational damage.
A New York Times article by Waldemar Kaempffert on Kaplan's
work appeared in March 1951, and it caught attention . He
quickly responded to Kaempffert's claim that although had
"proved mutation monstrosities to be produced by X rays in flies"
he "did not think they would be produced by the radiation from
atomic bombing. "3 Kaempffert cited Kaplan' s work as proof that
there was no need for concern about radiation damage from the use
of atomic weapons. pointed out to Kaempffert that mon-
strosities formed only a minor fraction of apercent of mutations,
regardless of the type of radiation used. Yet, "a very significant
amount of mutation al damage will have been produced, that will
plague future generations in an insidious way, despite the impossi-
bility of demonstrating it by ordinary means (i. e., with uncon-
trolled breeding)." If a lack of monstrosities could not be used as
"evidence against the production of mutations by X rays in human
beings in signillcant amounts" them, Muller argued, "you will see
the complete assininity of [the views of] Dr. Ira 1. Kaplan." This
emotional response to Kaplan's views was accompanied by a sharp
criticism of Kaempffert as a science journalist: "Your article has no
doubt given aid and comfort to the numerous groups of people
working with either the medical, the industrial, the commercial, or
'I. 1. Kaplan to F . C. Fraser, April 10, 1951 (note appended to F. C. Fraser to
1. 1. Kaplan, April 6, 1951), Lilly Library.
JH. J. MuHer to Kaempffert, March 14, 1951, Lilly Library.
337
 Genes, Radiation, and Society
the military aspects of radiation, who wish to give the public an
opiate on the subject of radiation dangers. "4
Kaempffert sent MuHer's letter to Kaplan for a reply, which was
then passed on to MuHer. MuHer, of course, had not intended his
letter to Kaempffert to be sent to Kaplan. The tone of MuHer' s
remarks generated an antagonistic response from Kaplan. He
claimed to be astonished at Muller' s vehemence and rejected the
attributions of "assininity" to his conclusions. "Even an ass, when
God opens its mouth (Numbers XXII) speaks words of wisdom ." He
insisted that the lack of damage in the children he examined
proved that the radiation hazard was imaginary. He praised the
caution of the D.S . Atomic Energy Commission, which claimed
that such radiation risks were unproved. He did not feel that re-
sults from flies could be applied to man and he claimed, in an
argument similar to Nuzhdin's many years earlier in Moscow, that
the dose he administered to a woman only involved a small part of
her body, the gonads and the pituitary gland, while Muller irradi-
ated whole flies. Even if mutations were induced, Kaplan thought
it unlikely that they would ever become homozygous but would
remain harmless in the heterozygous condition. After making these
counterclaims, Kaplan accused Muller of"angry sniping" based on
a faHure "to recognize without envy the success of a modest
contemporary. " 5
Kaempffert, in defense ofhis journalistic integrity, agreed with
M uller that Kaplan overlooked the long-term or insidious effects of
radiation damage, but he denied there were groups opposed to
MuHer's views of radiation and he doubted he was "giving aid and
comfort to an unknown number of persons who would minimize
the dangers 0 f exposure to gamma or X rays .
Muller's allegation that "numerous groups" denied the possibil-
itv of radiation damage to the genes and chromosomes was not,
imaginary. He had run into this barrier repeatedly since
1928, when "the radiologists at their meeting in Texas canceHed a
scheduled paper by me, telling them of the genetic effects of radia-
tion, simply because they had their minds made up in advance."
'Ibid .
' 1. 1. Kaplan to W. Kaempffert, March 22, 1951, Lilly Library.
6W. Kaempffert to H. J. Muller, March 26, 1951, Lilly Library.
33 8
Human l'vlutations and the Radiation Danger
Muller also insisted that "there has been a group of influential
people connected with the A. E. C. wh ich has attempted to mini:
mize or ridicule the genetic effect of radiation as applied to man.
None of these critics were geneticists or really understood the field
of mutagenesis, "yet they gave the public the impression that they
did. "7 Chief among them was Robley D. Evans, of the Mas-
sachusetts Institute of Technology. With the support of the D.S.
Navy's Office of Naval Research, Evans prepared a review of the
genetic effects of radiation. He relied chiefly on the work ofDonald
R. Charles at the Dniversity ofRochester and D. G. Catcheside, an
English geneticist. During the la te 1930'S, Catcheside criticized
Muller's theories; he was a strong believer in the target-theory
approach to measuring gene size; he also endorsed the contact
theory of rearrangements rather than believing they originated
from independent breakage, and he believed the lethal effects of
such rearrangements to arise from breakage of the individual genes
at the point of rearrangement. All of these views :VI ulIer considered
to be erroneous. Together with D. E. Lea, Catcheside published a
book on radiation genetics of wh ich was strongly critical.
Muller wrote his detailed objections to Evans's article on February
5, 1949, requesting Evans to accept his frank criticisms without
thinking hirn discourteous. "r can understand how a newcomer in
the subject, no matter how good his background in other lines,
could easilv be misled in this particular matter. It has many pitfalls.
Many of the quantities are only very roughly known even for Dro-
sophila, and we are admittedly extrapolating too far in applying
this to man, but it is all we can do in our present state of ignorance
and we must meanwhile remain on the safe side." Muller men-
tioned the activities of "certain persons working under the auspices
of the Navy, as weH as some under the Atomic Energy Commis-
sion, who have been attempting to minimize, in the public eyes at
least, the genetic dangers of radiation." To overcome their efforts,
MuHer urged Evans "to regard the problem objectively and to
prevent the users of radiation from becoming careless or the public
from taking the matter too lightly. The effects, because of their
insidiousness, are only tao easy to ignore. Nevertheless , every
' H. J. Muller to W. Kaempffert, :\pri130, 1951, Lilly Library.
339
 Genes, Radiation, and Society
death, no matter how far in the future, is a death, and must in all
conscience be guarded against as carefully as if taking place now, in
fact more so since we are setting a precedent which may
continue. "8
Not all medical societies were opposed to Muller' s plea for
restraint in the use ofX rays. The New York Academv ofMedicine
invited hirn to address them on April 1, 1947. He discounted the
legend which conveyed to physicians "the picture ofhizarre mon-
strosities, such as the arrnless-legless man, the microcephalic idiot,
the lion-faced boy, and other circus wonders." Some of these mav
have arisen from mutations, but they formed a small portion of aiI
mutations, and it was false to direct concern to them rather than to
the far more real and serious problem of the recessive lethals and
recessive detrimental mutations. These two categories constituted
almost all of the spontaneous and induced mutations affecting the
somatic tissues of the organism. To illustrate the confusion that the
nongeneticist feIt when attempting to assess genetic damage, M ul-
ler described how a 4,500-roentgen exposure could be evaluated in
a wild-type stock of Drosophila. "When we breed these with un-
treated flies, or with each other, we ordinarily find that, among
the offspring which develop, .not one in a hundred will show a
conspicuous visible abnormality, though, on searching through
thousands, we will eventually find a few abnormal types, of varied
kinds." In contrast, when MuHer used marked stocks that could
reveal lethaI mutations and, by the use of additional tests, det-
rimental mutations, "we find that, instead of there being only 1
mutation among over 100 offspring, as appeared at first sight to be
true, each one of the treated germ cells actually carried, on the
average, between 3 and 4 fresh mutations, if we include visibles,
lethals, and detrimentals. 9
The detrimental mutations were first obtained by Muller's stu-
dent J. J. Kerkis, in Moscow. Kerkis used marked X chromosomes
in population cages and measured the ratios of the two classes of
marked males emerging from the conditions of extreme crowding
and competition as embryos and larvae. Nonlethal X chromo-
"H. J.
Muller to R. D. Evans, February 5, 1949, Lilly Library.
"H. J.
MuHer, "Mutational Prophylaxis," Bulletin ofthe New forkAcademy of
Medicine 24(1947):447-46g; p. 453.
34°
\\
Human Mutations and the Radiation Danger
somes, made heterozygous with an untreated but differentlv
marked X, were used to establish a large number of heterozygou's
fernales. Stocks of these fernales were then tested in cages and
compared to control heterozygotes which carried identical X
chromosomes which had not been X-rayed. Kerkis. found that re-
cessive detrimental mutations were about three to four times more
frequent than recessive lethal mutations. 10
A se co nd feature of the mutation process was the way genetic
changes accumulated in the individual. Since most mutations are
recessive and heterozygous, they are only very gradually elimi-
nated and may persist for many dozens cf hundreds of generations.
Modern medicine has increased the rate of accumulation because
in treating a person for the symptoms of a disease, as when insulin
is injected in cases of childhood diabetes, the gene or genes in-
volved in the defect survive and can be transmitted to future gen-
erations. ''The more success we have, and wuh the more ailments,
the more does mutation take advantage of this to encumber the
population with these ailments, for which more <lnd more indi-
viduals have to be treated anew in each generation." .\-1 uller ac-
IUlOwledged. that this "does not menn that we should stop trying to
find the causes and eures of ailments and impairments" but that
"we should try to develop means to discourage the <lccumulation of
these hereditary weaknesses through indiscriminate reproduc-
tion." .\-luller realized, furthermore, that because genetic knowl-
edge was inadequate to identify defective genes and "who carries
the most and the worst," medical personnel could not apply such
"mutational prophylaxis"; he did feel, however, that physicians and
others in the health sciences could "take more effective steps to
protect the germ plasm of our population from having additional
mutations induced in it by penetrating radiation of artificial origin
and other mutagenic agents. "ll
A recessive mutation is rarely without some expression in the
10J. J. Kerkis, "Study of the Frequency of Lethal and Detrimental :\1 utations in
Drosophila," Izvestia Akademii Nauk SSSR, Otdelenie i Estest-
vennyh Nauk, Serija Biologicheskaja [Bulletin' of the Academv of Sciences of the
USSR, Seetion of :\1athematical and Natural Sciences, Biological Series], 1938,
pp. 75-96 .
11 :\1uller, ":\1utational Prophylaxis, " pp. 458, 459, 461.
34 1
 Genes , Radiation , and Society
heterozygous condition, although it mal' require tests other than
inspection of the phenotype to reveal its influence. The partial or
subtle expression of a recessive in the heterozygous carrier is called
partial dominance. In 1948, Curt Stern and Edward Novitski de-
signed experiments to test the partial dominance of spontaneous
and induced recessive lethai mutations. By comparing ratios of
survival between females carrying alethal in one of their marked X
chromosomes and females not carrying the lethaI, they found the
average dominance of the heterozygous lethai to be 5 per cent. 12
This conclusion \...·as consistent with earlier observations of visible
mutations in flies . Flies heterozygous for miniature wings have a
measurably smaller whig than those which are homozygous for
normal wings . Similarly, flies heterozygous for white eyes have less
red pigment than homozygous normal flies. In both cases, the
differences are not apparent to casual observation. 13 Levit, in 1935,
had also predieted the partial dominance of recessive mutations in
man based on his studies of several human disorders.
The effective partial dominance of these mutations suggested
two things to Muller. First, such a mutation would be eliminated
much more rapidly than if it were completely recessive; only a few
dozen generations would be required to eliminate the lethai,
whereas hundreds of generations might pass before a random en-
counter of the defective gene produced a homozygote. The second
implication was a subtle effect on an individual' s accumulated
"load" of mutations. These slightly dominant effects would impair
the vital organs in different degrees , depending on the functions
affected by the heterozygous mutations. In individuals with high .
genetic loads , they would, for example, lead to preeocious death
from heart attacks or stroke; they would increase the likelihood of
major surgery; they would make an individual more siekly, more
suseeptible to infections, and more irritable or diminished in natu-
ral vitality.
Muller's arguments, whieh he presented as his presidential
12Stern and Novitski, "The Viability ofIndividuals Heterozygous for Recessive
Lethals , .. Science 108( 1948):538-539.
i3 H . J. Muller, "Evide'nce of the Precision of Genetic Adaptation, " Haroey
Lectures 43(1950):165-229. The heterozygous w/w '" female has about 95 per cent
of the red eye color seen in homozygous w "' lw + females.
34 2
Human Mutations and the Radiation Danger
address to the American Societv of Human Genetics on December
29, 1949, showed how intensively he had explored the problem of
human mutations. In his paper, "Our Load ofMutations," he com-
bined the growing mass of experimental data providing the fre-
quencies of reeessive and dominant mutations with the newer find-
ings on the partial dominance of recessive genes. He relied on the
equations of C. H. Danforth, whose address he had heard at the
5eeond International Congress of Eugenies in 1921.14 Danforth
showed how the frequency of a genetic disorder in a population
remains in astate of equilibrium. The time required for elimina-
tion depends on the severity of the defect; moreover, less drastie
genetic defects take longer to eliminate. MuHer modined Dan-
forth' s basic theorem by taking into account the newer findings of
partial dominance and the concept of genetie load. In Muller' s
view, adetrimental effect of 5 per cent for a heterozygous mutation
would lead to the elimination of one such gene eVerj tvlcnty gen-
erations. Almost every newly arising mutation leads sooner or later
to the extinction of an individual. This "genetie death" can oecur as
a result ofhomozygosity or as the consequenee of a gradual buildup
in the genetie load whieh prematurely kills or sterilizes an indi-
vidual. U sing Drosophila data to supplement the meager informa-
tion on humans, estimated the human mutation rate to be
surprisingly high: "one newly arisen mutant gene in 10 germ eells,
on the average. "15 About 20 per eent of eaeh generation suffers
genetie elimination where eonditions of natural seleetion are still
severe, as in the underdeveloped countries or in the Western
world prior to the nineteenth century, when sanitation and mod-
ern medicine began to cut down the high incidenee of infant and
ehildhood mortality. 16
" C. H. Danforth, ''The Frequency of Ytutation and the Incidence of Heredi-
tary Traits in Man, " Eugenics, Genetics, and the Family 1(1923):120-128.
"H. J. MuHer, "Our Load of Ytutations," American Journal of Human Genet-
ics 2(1950h11-1'76; p. 169. This is not surprising: if the spontaneous mutation
rate for an average gene is 1 in 100.000 and if there are , conservatively. 10,000
genes in a sperm or an egg, then the rate would be 1 in 10. Neither the exact
number of genes in a germ cell nor the average mutation rate per gene in humans
is known, although reasonable figures exist.
l0E. A. Carlson, "Eugenies Revisited: The Case for Germinal Choice," Stadler
Genetics Symposia S(19i 3): 13-34.
34 3
 Genes, Radiation, and Society Human Mutations and the Radiation Danger
Since spontaneous mutations are so frequent and the mecha-
nisms governing them are unlikely ever to be controlled by human-
ity, MuHer advocated the avoidance of further genetic deteriora-
tion through the combination of "ameliorative techniques, such as
medicine, with a rationally directed guidance of reproduction,"
rather than through areturn to conditions that existed before the
advent of modern civilization. Muller' s arguments were not easy
for the layman to follow; nor were they clear to the physician or the
physicist, whose need to be aware of the dangers of ionizing radia-
tion was MuHer' s primary concern. U nlike relativity theory or
quantum theory, whose principles were acknowledged to be in-
comprehensible by the layman, genetics was a subject about which
most individuals had some strong opinions, based partlyon family
history and partlyon folklore. The monstrosity legend-the belief
that genetic damage, if it existed, would be clearly visible-was
one of the most ingrained of popular fallacies, and the absence of
the expected monsters was cited repeatedly by professional people The Drosophila laboratory course in Jordan Hall, 1957. The large fly was purchased
and organizations critical of MuHer' s views. trom a Gennan supplier and it included mutant eyes, wings, bristles, and legs that
Muller' s domain was expanding in· the basement of Kirkwood could be removed or inserted far display.
HalL In his own laboratory he designed the elaborate genetic
stocks for his experiments. Adjoining it was his secretary' s room looked for a research associate to take over many of these tasks and
with his large collection of reprints on genetics. Two large labo- engaged Irwin Herskowitz, a former student of Dobzhansky' s who
ratories were equipped for his technicians to carry out the had taken a strong interest in mutation research, especially chem-
thousands of tests required for mutation studies. There were also ical mutagenesis.
two additional laboratories for the graduate students and a kitchen uller wanted Herskowitz to relieve hirn of most of his day-to-
in which four workers prepared the food for the flies and cleaned day contacts with the technicians and to see to it that the research
the vials of their old food. M uHer purchased an ultraviolet micro- projects for the grants were carried out. He did not discuss Hers-
scope to improve the resolution of banding in the chromosomes. kowitz' s responsibilities with his students and staff, some of whom
To supervise his large-scale projects he hired Helen Meyer, a were resentful of an outsider stepping in and inquiring about their
refugee from Germany, who had received her doctorate there in own research activities. Nor did he supervise Herskowitz very
cytogenetics. His graduate students increased in number, with closely. To Yluller's surprise, a laboratory revolt took place, with
John McQuate, Margaret Edmondson, James Telfer, Seymour an anti-Herskowitz faction led by Juan and Ruby Valencia and
Abrahamson, Shanta Iyengar, Abraham Schalet, and Irwin Oster Oster circulating a petition for Herskowitz' s removal. MuHer caHed
carrying out their own projects or assisting in Muller's experi- the laboratory staff together and backed Herskowitz as a necessary
ments. By 1951 MuHer feIt swamped with the administrative de- addition to his research plans. The Valencias left in protest, and
tails of filling out grant applications, ordering materials and equip- Muller remained, from then on, without a salivary-gland cytolo-
ment, supervising research, preparing his own stocks, writing in- gist. It was a frustrating dispute, but MuHer believed his research
vited lectures, and getting ready for classroom meetings. He would suffer from an overextension without Herskowitz's help and
344 345

1\
 Genes, Radiation, and Society
he tried to restore confidence in his staff by holding weekly confer-
ences with each of his students and research assistants. I.
Muller was disappointed by the resignation of the Valencias.
He wrote Kenneth Cooper, a cytologist who worked on many
problems of interest to Muller, of the change in his laboratorv:
"Valencia is already on his way back to South America and Ruby
expects to join hirn next September. He was unhappy here because
he no longer held the chief position in our group even though I had
offered hirn the job Herskowitz has before Herskowitz came, and
he preferred not to accept it. "18 It was a particularly severe loss to
Muller because he valued Juan Valencia's excellent cytological
skills. The Valencias had also carried out two projects which MuHer
feit were essential for his mutation studies. One was a studv of
spontaneous mutations at nine specially selected loci on X to assorted journals. 21
chromosome. Determining the mutation frequency, which was
about 1 in 100,000 at each locus involved an immense amount of
work, and only an abstract had been 'written because of the revolt.
The second study, some 200 visible mutations at nine loci
induced by high dos es of X rays to immature eggs, was also left in
the abstract stage of publication. 19
For the first time since he had left Texas in 1932 on a Gug-
gen heim fellowship, Muller applied for a sabbatical leave of ab-
sence and accepted an invitation from friends in Hawaii to enjoy the
relaxing atmosphere in Honolulu. There were several advantages
to this plan: it would give hirn a chance to prepare two comprehen-
sive chapters on radiation genetics for a two-volume series on
radiation biology edited by Alexander Hollaender of the AEC' s
biology laboratory at Oak Ridge, Tennessee; it would give Hers-
kowitz a chance to ron the laboratory in his absence and thus mend
his fences with the students and staff; it would also give MuHer and
his family a much needed vacation.
I'The two camps existed, to some extent, when I was a graduate student there
(1953-1958), but newcomers Iike myselffelt at ease with all parties to the dispute.
ISH. J. Muller to Kenneth Cooper, May 12, 1953, Lilly Library.
lOH. J. Muller, J. 1. Valencia, and R. M. Valencia, "The Frequency of Spon-
taneous Mutations at Individual Loci in Drosophila," abstract, Genetics
35(1949):125-126; H. J. Muller, R. M. Valencia, and J. 1. Valencia, 'lhe Produc-
tion of at Individual Loci in Drosophila by irradiation of Oocytes and
Oogonia," ibid ., p. 126.
Human Mutations and the Radiation Danger
The Mullers sailed on the S.S. Lurline for Hawaii on June 19,
1953, arriving June 24. The delightful beaches, tropical flowers,
and volcanic mountains provided a varied terrain for the MuHers to
enjoy on frequent hikes and outings. MuHer also completed the
two long chapters: ''The Nature of the Genetic Effeets Produeed by
Radiation" was a 123-page primer on radiation genetics, and "The
Manner of Production of Mutations by Radiation" was a lSl-page
evaluation of the complexities of radiation genetics. 20 Together they
were large enough to form a textbook in radiation genetics, but
throughout his career Muller feared eommitting hirnself to writing
a text, which would be too time-consuming, only to find hirnself
doing an equivalent amount of work and dispersing the "ehapters"
Radiation damage in humans was not limited to changes in the
individual genes. Chromosome breakage was also involved, and
while preparing his ehapters on radiation genetics Muller applied
some earlier findings to interpret ceH death from ionizing radia-
tion. He suspeeted, from earlier work with Lamv and Ponteeorvo
that chromosome loss and abnormal formation
after radiation exposure. The effects were expressed only in divid-
ing eeHs. Barbara .vlcClintock, at the University of Missouri, had
also followed the eyde of chromosome movements in some abnor-
mally dividing cells of maize in 1939 and worked out a mechanism
of chromosome breakage, followed by replieation of the
fragments. 22 The fragments fused at their replicated broken tips
and then, during the separation of the replicated parts, remained
connected at their point of fusion, forming a chromosome bridge.
This "breakage-fusion-bridge" cyde eventually killed the cell by
preventing eeH division or by piling up abnormal quantities of
lOChapters 7 and 8 in Radiation Biology, eu. Alexander HoIlaender (New York:
McGraw-Hill, 1954), 1:251-473, 475-626.
uThis is probably why :\-Iuller's views were less influential than they could
have been. :\-Iorgan, of course, wrote a dozen or more books and made his posi-
tions internationally known. :\-lulIer knew the strain of having to meet a deadline
for submitting manuscripts; he knew the demands of a book would cause all his
other activities to be scuttled and deemed it too high a price to pay.
"Barbara :\-lcClintock, The Fusion of Broken Ends of Sister Half-chromatids
Following Chromatid Breakage at }Ieiotic Anaphases, University of :\lissouri,
College of Agriculture, Research Bulletin 290 (Columbia, 1938), pp. 1-.48.
347
 Genes, Radiation, and Society
Human Mutations and the Radiation Danger

ing (10), producing a giant cell with nearly double its chromosome number (11).
If the bridge breaks (12), the nudeus does divide (13) but each cell retains a
single break to perpetuate the cyde (14). Note that the acentric fragment is
often left out and is digested by the cell cytoplasm.
7
applied the breakage-fusion-bridge cyde to radiation damage and the
6,Acenlric 5 ,/
symptoms of radiation sickness, which affect actively dividing tissues. A chromo-

,feen,;;: d; some break does littIe damage to a nondividing cel!.

genes in the chromosome fragments involved in the cycle (see

Figure 18). The victims of Hiroshima and Nagasaki who died of
radiation sicklless were killed, in view, primarily by the
breakage-fusion-bridge cycle, which prevented rapidly dividing
cells in the blood and in the lining of the capillaries and intestines
from replenishing the cells lost by chromosome breaks. The symp-
1 toms of radiation sickness- . -anemia, susceptibility to infection,
10 1 hemorrhagic rash, and peritonitis-were explainable by the fact
that the organs governing them contained numbers of
r
12
dividing cells. Tissues like muscle , nerve, and bone , which da not
divide much, if at all, in the adult, were not affected by radiation.
The breakage-fusion-bridge cycle also explained radiation-induced
sterility and the high susceptibility to radiation damage of fetuses
/ in early pregnancy. !J
11 t Muller's pleasure in Hawaii was based on more than the para-
dise it offered weary tourists. Hawaii was one of the most racially
integrated regions of the world. admired the vigorous and
handsome children, part Hawaiian, Caucasian, and Oriental, who
accepted their mixed heritage with pride. Unlike the early partici-
pants in the American eugenics movement, who looked down on
miscegenation, uller favored the process, which led gradually to
FIGURE 18. The breakage-fusion-bridge cycle: This cyde of single breaks
leading to chromosome loss was analyzed by Barbara McClintock using maize. new combinations of traits and which gave to many of the offspring
Muller and Pontecorvo realized that this model could be applied to similar the hybrid vigor characteristic of matings between genetically iso-
chromosome losses leading to cell death, especially if a sperm or egg receives lated strains.
such a chromosome break, causing a fertilized egg to abort.
The chromosome is shown intact (1) before a chromosome break (2) fonns two
Midway in Muller's sabbaticalleave he had a physical checkup
fragments. These replicate (3), and the replicated broken tips join (4). As the which revealed two disturbing conditions, high blood press ure and
chromosome enters mitosis and coils (5), the centric and acentric pieces are evidence of a past heart attack resulting in a slightly enlarged heart
made compact (6) and are aligned (7). Note that only the centric fragment, bearing and abnormal electrocardiogram. On October 30, 1953. he wrote
the centromere, is attached by spindie fibers . When chromatids are distributed
(8), the acentric fragment cannot participate and the centric fragment is tied by
a chromosome bridge (9).
Two fates may befall the cello The bridge may prevent the nudeus from divid- 1.1S ee H. J. "Potential Hazards of Radiation, " Excerpta .'ifedica
1, 1(195i): 52 i -530.
34 8
349

i1.I',
 Genes, Radiation, and Society
to Herskowitz that he was going over the galley proofs of the two
chapters for Hollaender' s book, that he had just finished his com-
ments on McQuate' s thesis, and that he was writing a manuscript
for Columbia University's bicentennial. "Yet I'm having to go slow
as the doctor here just discovered a heart ailment (chiefly coronary
insufficiency). But please don't talk it around as I'm not letting
others know for the time being anyhow. It's not bad but it cuts
down outdoor activities-for good, I suppose. "24 The diagnosis was
a surprise to Muller because in early March, when he had his
physical in Indianapolis, his blood pressure had been at a level
considered normal for his age; his EKG was also normal (no exer-
eise test had been done), and his heart showed no symptoms other
than a lifelong systolic murmur.2.'5
Herskowitz kept MulIer's secret until the spring of 1954, when
the deluge of letters fulI of laboratory problems became too much
for Muller while he was trying to recover from minor surgery in
Honolulu. He needed rest and at least a temporary change in his
work habits. Thea objected to Herskowitz' s idea of having her
select from the incoming mail what to pass on to Muller. Hers-
kowitz agreed to cut down the flow of work by forwarding less maiI.
Assuming that handwritten letters would be shorter than typed
ones, he asked those in the laboratory to send only handwritten
letters .
Just before his heart ailment was revealed, Muller conditionally
accepted me as a graduate student and suggested to Herskowitz
that he might try to interest me in population genetics, espeeially
to study the effects of sex-linked detrimental mutations in heter-
ozygous females .26 MulIer' s methods as a graduate sponsor were
subtle. He provided a research project for a student, who would .
then be expected to develop his or her own insights into that
problem. The results would indicate to Muller whether or not the
student had developed some independenee of thought in ex-
perimental design. A student who merely followed the plans Mul-
ler had prepared would be considered limited, and M uller would
:H. J. to ITWin Herskowitz, October 30, 1953, Lilly Library.
Thea :vtuller, conversations, summer 1972 .
26
1 hated the project and dropped it, replacing it with my own thesis project on
pseudoallelism (gene structure).
35°
Human Mutations and the Radiation Danger
not reeommend hirn highl)' for a good position as a genetieist.
Students could also pursue a third route, by running their own
projeets parallel to their assigned projects; if their own work pro-
dueed significant results, they could then barter with M uller and
use their 0\\<'11 experiments as areplacement for their assigned
thesis work. In this way Muller's better students did twice the
amount of work they were initially expected to do, particularly if
they were anxious to find a niche of their own in genetics through
their Ph. D. work. 27 Funds to subsidize research were still scarce,
and most of MulIer' s students supported themselves as teaching
assistants. During the summer, however, they usually worked as
research assistants on one of Muller's many projects funded
through the RockefeIler Foundation, the Cancer Soeiety, and, la-
ter, the Atomic Energy Commisskm and the National Seience
Foundation.
.
Bv, the time Muller returned from Hawaii, he was in much
bettel' health. He still kept up his seven-day-a-week sehedule ex-
ce pt on Sundays, when he worked only half a day. He took a nap at
horne after lunch each day and walked at least ollee a day to and
from the iaboratory for exereise and invigoration. His students
followed a similal' pattern of working seven days a week, and the
lights burned late into the night as the experiments piled up. One
weekend I moved my household goods, a job carried out by hard
labor with many trips in a borrowed car to shuttle bricks, boards,
and books aeross town. came into the laboratory that
weekend looking for some information to work into a manuscript.
On Monday he stormed into my office and exclaimed, in frustrated
rage, "Don't you know that in this laboratory weekends aren't
21This was a common practice in the mid-1950's and not unique to :vtuller. In
the late 1950's and throughout the 1960'S, federal support for research became
widespread.
wasn't really an expioiting task master. He imposed the same stan-
dards on himself. He would often labor hours into the night, or on a weekend, to
compiete a manuscript or collect virgin flies for his stock designs. He found it hard
to believe that athers were less devoted to their work.
35 1

[25]
The Fallout Controversy
On November 1, 1952, in the remote Pacific near Eniwetok
atoll, an explosion occurred, producing a fireball three mHes in
diameter. After years of controversy among atomic physicists, the
view of Edward Teller had won out over that of J. Robert
Oppenheimer, and the United States had succeeded in construct-
ing a device which could be used to make hydrogen bombs. No
longer were atomic bombs to be limited to the equivalent of sever-
al tens of thousands of tons of TNT in destructive power, as the
twenty-kiloton Hiroshima bomb had been; the new hydrogen
bombs could be made in almost any destructive capacity desired.
The megaton replaced the kilo ton in military jargon, and the world
slowlv realized there was no longer any assurance of survival.
more ominous, about a year later, was another H-bomb
explosion which released an unexpectedly long trail of radioactive
debris that settled on the Marshali Islands, causing some 3 00 of the
inhabitants to suffer mild symptoms of radiation sickness and forc-
ing them to. abandon their hornes far more than a decade. The
H-bomb did more than annihilate alilife in the vicinity of its fire-
ball; it released ablanket of fallout which could kill all unprotected
survivorsin its path for a distance of 100 or more miles. Another
incident took place some ninety miles from Eniwetok in March
1954, when the skipper and crew of the Fukuryu Maru (the Lucky
Dragon) were concluding an active day' s fishing. Their ship was
caught in a sudden rainstorm, whose clouds contained the highly
radioactive debris which had been fonned from the minerals in the
35 2
The Fallout C ontroversy
ocean and the vaporized coral island on which the bomb had been
detonated. 1
The accidents were at first kept quiet, with the AEC physicians
vigorously denying that the death of the radio operator on the
Japanese fishing boat was connected with the fallout. All the crew
had suffered radiation sickness, and they had been hospitalized
after their craft reached horne. The accusations and the denials
became known throughout the world and in the U.S. press, but
Muller feit that the actual infonnation on which to base judg-
ments of the severity of the fallout and its distribution remained
classified. Z
Thus the public did not know that during "Operation Cross-
roads," in 1946 at Bikini atoll, an underwater atomic bomb re-
leased so much radioactive material that the fish, clams, lobsters,
sea urchins, algae, and other marine life became highly radioac-
tive. The accumulation of radioactive trace elements in living tis-
sue results in a biological magllification of radiation. This effect
soon became apparent to Navy scientists studying the tood chain in
the surviving organisms at Bikini. Even the barnacles below the
waterlines of the Navy ships stationed at the atoll became so
radioactive that they could be detected through the steel hulls.
Despite these massive dos es of radiation, the natural marine ecol-
ogy soon returned to its fonner abundance and the Navv scientists
feIt relieved that, contrary to their expectation, there was no evi-
dence of mutations in any of the thousands of plants and animals
they examined. Their expectation was false; it was based on the
naive view that such mutations show up in the Flor F 2 as deformi-
ties or monstrosities or visible mutations.
The United States was not alone in its efforts to develop mili-
tary sophistication in atomic weapons. In 1949, President Truman
announced the first Soviet atomic test, and in 1952 Great Britain
lAccounts ofthe early tests ofhydrogen bombs and the discovery offallout are
from Encyclopaedia Britannica, 1959 ed., 3:559.
'''It would be very strange if the Russians did not alreadv have this information
as a result of their own experiments and measurements. It also seems strange,
therefore, that it should be 'reasons of securitv' which makes such information
unavailable to the public in the democratic (H . J. to Yl. Hers-
nat, December 11, 1954, Lilly Library). y(uller cites a .Yew York Tirnes article of
December 3, 1954, as the basis for his judgment.
353
 Genes , Radiation , and Society
detonated its first atomic bomb in the Pacmc. In 1953 the Soviet
Union exploded its own hydrogen bomb, and public terror of
mutual annihilation continued to growas the tests continued.
The misfortune of the few victims in the far Pacific was not the
sole worry of apopulace fearing for its own survival; throughout the
world scientists could detect the tests carried out in the Pacific, in
the southwestern United States, and in Siberia, bv the telltale world-
wide fallout of a far less terrifying dose which' slowly permeated
the atmosphere. No longer were the unlucky few in faraway places
the victims of radiation exposure. All of humanity now inhaled a
common thinly polluted air, whose faint levels of radioactive iso-
topes would rise and fall in response to the tests carried out by the
United States and the USSR. 3
The reality of these new developments still remained quiescent
when Muller and his family sailed to Honolulu in June 1953. In less
than a year, nearly fifty explosions would be set off bv the United
States, although only a few of these were publicly at the
time they occurred. M uller endorsed the necessity of weapons
testing, asserting that the "destruction of civilization would be
much more likely to occur if such weapons as the atom bomb were
not developed by scientists in the United States, since thev surelv
would be developed by those in the Soviet Union anyway and
that case would be used, either directly or through the pressure
created by the threat of their use, to accomplish the enslavement of
all mankind. "4
MuHer worried about radiation effects, of course, but he feIt the
protection given by new and formidable weapons that could deter
any Soviet attempt to overrun Europe or engage in warfare with
the United States was weIl worth the brief exposure from a few
tests. In the meantime he was directing his radiation wamings to
the medical profession:
Sometimes the warnings have been sensationalized and exagger-
ated so that people got the idea that their children were likely to
J"Local" fallout from hydrogen bombs involves hundreds of roentgens to the
exposed individual; "worldwide" fallout is measured as a verv small fraction of a
roentgen. Thus the individual risk is low for worldwide fallout, but it involves the
entire world population, rather than a small segment of it.
'H. J. Muller to Henri Corbiere , February 1.8, 1953, Lilly Library.
354
The Fallout Controversy
be monsters, but more usually the warnings were quietly side-
tracked by the people dealing with radiation, both medical men
and others, because they did not want to have the public think that
they might be doing some damage. When referring to my warn-
ings, they usually made it appear that I had indulged in the same
kind of sensational presentation of them as that of the alarmists,
because, of course, that is the kind of claim that can more readily
be disproved and dismissed. In other words thev raised a straw-
man and then knocked hirn down and distracted attention
from the real issue. 5 •
The "real issue" came to .MuHer via leaks from journalists who
reported speeches given by AEC spokesmen. J. C. Bugher, for
example, director of the Division of Biology and :VIedicine for the
AEC, presented a paper on the medical effects of atomic blasts in
September 1954. He outlined the physical details of the H-bomb
explosion. About 6 per cent 01' the bomb' s energy was released as
immediate nuclear radiation in the firebaH and an additionalu per
cent was slowly released through the fission products of the bomb
and the induced radioactivity from the air, water, or minerals in
the vicinity of the fireball. The Pacific tests released a worldwide
fallout of 100 millicuries per square mile, resulting in an external
whole-body dose of 0.01 milliroentgen per day, which Bugher esti-
mated was less than 5 per cent of what the bodv receives from
normal background radiation. He admitted that in' some instances
there might be "biological amplification" as grazing animals ate
contaminated grU?s and accumulated radioactive iodine-131 in the
thyroid glands or strontium-90 in the bones and milk. These, in
turn, would be concentrated in the thyroid glands and bones of
children who drank the milk. 6
uHer feIt Bugher' s paper was insufficient in its coverage,
since it neglected all but the external gamma radiation of fallout,
but he was, "inclined to think that the danger from the tests alone
has been much overrated .... Nevertheless, the damage from the
use of nuclear energy in war itself remains quite another thing
and so, for that matter, does that created by current medical
'H.J. Yluller to :Vlrs. C. C. Little, Ylarch 31, 1953, Lilly Library.
0J.C. Bugher, ''The Yledical Effects of Atomic Blasts," .\EC press release,
September 1.3.. 1954. Lilly Library.
355
 Genes , Radiation, and Society
practices.'" M uller maintained this cautious, conservative view
throughout 1953 and 1954· When Sturtevant presented an address
to a health conference in Houston, Texas, on the radiation danger, 8
MuHer was at first pleased that the conscience of geneticists was
leading them to speak out on public issues. A news account of it
that attributed to Sturtevant a "fear of monsters" brought a sharp
rebuke to the newspaper from MuHer. The reporter, Stewart
Alsop, wrote to Sturtevant, sending a copy of MulIer's letter, and
received approval of what he had written. M uller was appaIled: "1
had no idea, when 1 read Sturtevant's article, that he really did
subscribe to the popular view that it is the production of monsters
which constitutes the chief danger of radiation to posterity."9
It was discouraging to try to steer a balanced course in apprais-
ing the radiation danger when errors like Sturtevant' s were made.
Muller did not know why Sturtevant had fallen into such ablunder:
"it may be that the misunderstanding has an emotional background
since I have reason to believe that he has long had a personal
resentment against me since the early days of the Drosophila work.
I cannot believe that Sturtevant is simply so stupid as to misunder-
stand for I have the highest respect for hirn professionally. "10
A change in attitude seemed to be developing when M uller
received a telephone call in March 1955 from Earl Green of the
AEC. He asked MuHer to prepare a paper on "the mechanism of
production of mutations by radiation" for the United Nations Inter-
national Conference on the Peaceful Uses of Atomic Enerl!V which
was to be held in Geneva in August. This was to be the fi;st such
conference bringing together experts from the USSR and the
Western world. M uller agreed, but he asked if the title could be
changed to "How Radiation Changes the Genetic Constitution."
This broader topic would permit hirn to discuss chromosome
breakage as weIl as gene mutations; furthermore, it would not limit
hirn merely to the production of mutations but would include the
'H. J. Muller to Eugene Rabinowich, December 23, 1954, Lilly Library.
8A. H. Sturtevant, 'The Genetic Effects ofHigh-Energy Irradiation ofHuman
Populations," Engineering and Science 18(1955hrlZ. .
J. Muller to Stewart Alsop, October 16, 1954, Lilly Library.
lOH. J. Muller to Benjamin Sonnenblick, November 5, 1954, Lilly Library.
Note, incidentally, the attribution to Sturtevant, not himself, of responsibility for
the alienation in the Drosophila group.
The Fallout Controversy
consequences of such mutations in the population. Green told
M uller that his paper would be one of several submitted and that a
final selection of U.S. delegates would be made in June. He
assured MuHer, however, that this would be only a formality in his
case; in fact, he wanted MuHer's assurance that he would attend
the conference. He also apologized for the inconvenience uHer
would face in preparing a lengthy paper on such short notice.
A form letter soon followed, indicating that the UN woulc.l notify
the AEC of the selected papers on June 15, 1955, and the AEC
would transmit this information to the chosen delegates shortly
afterward. A preliminary agenda, sent in showed an AEC-
recommended panel of Muller, W. L. RusseH, John Gowen, Nor-
man Giles, Bruce \VaHace, and E. L. Powers. All except Powers
were radiation geneticists. Shortly afterreceiving this, sent
offhis manuscript, making sure it would arrive in time for the AEC
to transmit it to the UN selection committee. ll
Muller decided the trip to Europe would be a good occasion for
Thea and Helen to see their relatives in London and Germany.
Thea had not seen her brother in twenty years, and neither her
father nor her brother had ever seen Helen. MuHer planned to
combine this trip with a separate talk he intended to give at the
annual conference of Nobel Prize winners at Lindau on July 10.
The Geneva conference was scheduled for August 15, and this
would give the Mullers a month's vacation in Switzerland. On June
18 they flew to Europe and hoped the official word from the AEC
would be sent shortly after so the arrangements could be com-
pleted.
A month passed quickly, and by July 20 there was still no ward;
now MuIler was beginning to worry about the delay. He wrote a
letter of inquiry that day to the AEC. lz In the meantime a letter
dated July 18 was se nt by the AEC to MuHer at Indiana University, .
infonning hirn that his paper had not been accepted for oral deliv-
ery and because of a limitation on the number of allotted delegates
llThe account ofhow Yluller received news of the conference and prepared for
it is from his "Chronology re!ating to the paper submitted by me for the Interna-
tional Conference on the Peaceful Uses of Atomic Energy ," dated October .f,
1955, Lilly Library.
12Thea Ytuller, personal communication, summer 1973.
357
 --- - -- - - -
Genes, Radiation, and Society
Herskowitz, who attended to Muller's mail, immediately called
Switzerland to convey the shocking news. In his response to Mul-
ler's July 20 letter ofinquiry, G. C. Weil, technical director ofthe
AEC, regretted the inconvenience to hirn, but he pointed out that
the AEC did not receive the UN decision withdrawing Muller's
paper until just before he wrote to Muller on July 18. This was
misleading. The UN had been told by the U.S. Atomic Energy
Commission to withdraw the paper. Thus, the UN, not the AEC,
was made the apparent instigator of the rejection of Muller' s manu-
script. Weil had clearly misled Muller on July 18, when he wrote
that "we have recentlv learned on the basis of a tentative agenda
prepared by the Nations for the Conference that your pa-
per has not been requested for oral presentation. "13
Delegates to the conference began to arrive for vacations in
Switzerland in early August, and Muller learned from them that
the AEC panel had now been pared to Russell and Wallace as
American speakers; T. C. Carter from England was selected by the
UN program committee to complete the radiation genetics section.
Originally five speakers had been planned, each with thirty min-
utes' time for presentation. Now only three were to present pa-
pers, with twenty minutes allotted for each. The session would be
chaired by Tage Kemp from Denmark; and Ake Gustafsson, from
Sweden, would be vice-chairman. One of the Scandinavian dele-
gates gave Muller an observer' s card so that he could attend as a
member of the audience. Numerous rumors began to circulate
about Muller's absence from the invited speakers. According to
one source, the AEC had given his paper a high recommendation
but word of Muller's heart condition had led the selection commit-
tee astray and he had not been expected to attend. 14 Kemp later
l3The AEC's request to the UN to withdraw Muller's paper was relayed to
YluHer. See H. J. MuHer to M. Westergaard. August 31, 1955: and G. W. Beadle
to H. J. Muller. September 17. 1955. The implication that the UN withdrew the
paper is conveyed by G. C. Weil to H . J. Muller, July 18, 1955. A letter from Paul
W . McDaniel, technical papers officer for the AEC, to H. J. Muller, July 22,
1955, repeated this erroneous impression: "we were not informed of the final
decision taken by the United Nations until just before we wrote you." Allletters
are in the Lillv Librarv.
'<Earl to H'- J. Muller, October 20, 1955, Lilly Library .
The Fallout Controversy
asked Muller ifhe would like to speak for about five minutes to the
audience during the thirty-minute question-and-answer session
following the presentation of papers. He said he had checked with
Walter Whitman, the general secretary of the conference, and
received approval. MuHer agreed to this proposal and spent many
hours condensing his paper to a five-minute presentation. But at
the meeting, on August 15, Muller was told by Kemp and later by
Whitman himself that his offer had to be withdrawn because he
was not an official U. S. delegate. 13
Russei!' s paper was on the genetic effects of radiation on a small
number of genes in mice. By mating normal X-rayed mice to mice
carrying recessive gene mutations, Russell could pick up any newly
induced mutations of these particular genes. This was an applica-
tion of what Muller called the "specific visible test" in Drosophila,
a method he and his colleagues had used on several occasions to
obtain frequencies of mutations for a selected number of individual
genes. From Russell' s data there was no question that X rays pro-
duce gene mutations in mice, and thus the paper was, in Muller's
view, an important contribution wh ich was representative and
worthy of selection for a panel on radiation effects. Muller had
reservations, however, about the choice onVallace's paper. 16
Wallace had studied different populations of flies under com-
petition. One strain had received chronic radiation exposure and
the other had not. By counting survivors of the two marked strains,
\Vallace believed he could tell how effective selection was in elimi-
nating the X-rayed strain. MuHer disagreed with the design of the
experiment. Most flies, treated or untreated, fail to survive ex-
treme crowding. Thus if 90 per cent or more of the eggs laid did
not become adult flies, the few surviving progeny of irradiated flies
might be those without induced mutations, or they might be flies
containing rare compensating beneficial mutations. If the latter
were true, the absence of a difference in the ratio of X-rayed to
control flies would not be remarkable. But if all the eggs laid were
allowed to survive, then there would have been a pronounced
" H. J. Muller to A. H. Rosenfeld, August 31, 1955, Lilly Library.
1·"1 am sure that Bruce Wallace is sincere, but also that he is very much
mistaken, and 1 think that the whole Lerner-Dobzhansky group is 'barking up the
wrong tree'" (H . J. Muller to G. Bonnier, July 18, 1958, Lilly Libraryl.
359
 Genes, Radiation, and Society
difference in the viability of the adult flies emerging, as weU as of
later generations among those carrying induced mutations. In hu-
man populations, particularly in those countries with good public-
health pro grams and medical care, the survival rate of children in a
family is high. There is also a tendency to identify biological find-
ings using other organisms with the human condition. Thus Wal-
lace' s experiments would lead an audience (most of whom not
geneticists) to conclude that radiation would not affect the descen-
dants of even a heavily irradiated human .population.
At the conference in the role of observer, MuHer found an
unusual response to the AEC snub. Almost every speaker referred
to MuHer' s contributions to radiation genetics, and each empha-
sized his role in the development of this field, using phrases such
as "the father of radiation genetics." Several mentioned, in their
oral presentations, that M uHer was sitting in the audience, arid
they deplored the fact that he was not allowed to speak. Gustafsson
criticized the official policy which kept Muller muzzled, and at the
end of the second session, he suggested that the audience show its
appreciation to MuHer by standing in tribute. l7 The speakers,
chairmen, delegates, and observers then rose and gave MuHer a
prolonged ovation, to his surprise and inner pleasure.
Kemp told MuHer he had checked out the AEC response to
Muller's inquiry of July 20 and could definitely deny that the UN
had rejected his speech. Quite the contrary was true. It was rated
high by the UN selection committee and never turned down by
them.
When asked why he thought he was banned, M uller said the
main reason was fear by the AEC that public knowledge of the
genetic effects of radiation would create an outcry against military
and industrial radiation. Rather than try to educate the public
about the magnitude of radiation hazards, the AEC, Muller be-
lieved, adopted a policy of denial and censorship to delude the
public into believing there was no genetic effect of radiation. 18
M uller had sufficient cause to believe this from the instances over
" Ake Gustafsson, personal communication, August 1973, Berkeley, California
(at Thirteenth International Conference of Genetics). Also H. J. MuHer to M.
Westergaard, August 31, 1955, Lilly Library.
'"H. J. MuHer to J. R. Kirk, September 10, 1957, LiHy Library.
3 60
The Fallout C dntroversy
the previous ten years when the AEC showed poor judgment by
presenting their viewpoint with "experts" who were virtually un-
known to the body of radiation geneticists.
After the conference, Muller retumed to join his family in Gun-
ten, at the Lake of Thun, and shortly thereafter they retumed to
Bloomington so Muller could prepare for the fall semester. In the
meantime, the press had begun its own investigations of the AEC
gag on Muller, and Warren Unna ofthe Washington Post broke the
story on September 17 as a front-page item. Kemp's findings. had
been confirmed, and the UN was irritated with the AEC's attempt
to pass the blame to them. Brian Urquhart, deputy to Whitman,
claimed the UN had favored MuHer' s paper but the AEC withdrew
it. He cited a letter ofJune 30 sent to hirn by G. C. Weil request-
ing the withdrawal and stating that Muller would not be a U.S.
delegate. 19 The AEC did not deny this accusation but it claimed
that Muller'!) paper included reference to the atomic bombings in
Japan and "such a paper, presented orally, would have opened the
Conference session tor discussion. A discussion on the bombing of
Hiroshima would have been out ofbounds for the
This was not, in Muller' s mind, a valid basis. If they had re-
quested it ofhim, he would have deleted the reference, and, at any
rate, he did not think that "this case should be judged on the basis
oflegalistic roIes." :\Iuller's reference to Hiroshima was not polem-
ical; "Many persons unfamiliar with genetics have regarded the
seeming normality of children born to survivors of the Hiroshima
and Nagasaki bombings as evidence against the conclusion that the
amount of radiation there received produced a significant amount
of genetic damage. "21
Legally, of course, the AEC was within its rights to recommend
delegates of its own choosing. Muller did not contest that right; but
he did object to the agency's use of public lies to cover up its
'reasons and to the devious practices it used at the Geneva
conference.:!2 Muller's speech, for example, which was to have
'"H. J. MuHer, "Chronology relating to the paper ... "
"'Statement by Lewis L. Strauss at AEC press conference, October 3, 1955,
mimeographed press release, LiHy Library .
21 H . J. MuHer, "How Radiation Changes the Genetic Constitution," Bulletin
of the Atomic Scientists 11(1955):329-339; p. 336.
'"H . J. to J. S. Huxley, November 11 , 1955, Lilly Library.
3 61
 Genes, Radiation, and Society
been available in mimeographed form at the conference, was not
distributed to anyone but the official U. S. delegates. But this
offense was minor when compared to the apparent censorship of
ideas: "material which was obviously considered by the scientific
personnel in the subject concerned to be of importance for the
conference was prevented from being presented through the influ-
ence of the administrative personnel or personnel at least re mo te
from the subject concerned. In other words, there was an attempt
by an administrative arm of the government to influence thought
and theory in scientific matters and the dissemination of the con-
clusions reached by representative scientists."2.3
Russell was furious over the way things turned out. Just before
he left for Geneva, Russell had heard "an odd rumor" about Mul-
ler' s Geneva paper and tried to find out wh at had happencd. At
Geneva he sent MuHer the information on the genetics program
with its three speakers and he denounced the "utterly despicable
behavior" of the persons who were responsible for the decision to
bar Muller. He considered withdrawing in protest, but this would
have been self-defeating, leaving Wallace alone to represent the
United States with the implication that small doses of radiation may
even be beneficial (by causing hybrid vigor). Carter would offer his
overly conservative appraisal that nothing was really known about
the genetic effects of radiation. Russell suspected, correctly, that
the AEC and not the UN was behind this, but ali his attempts to
find out were met with buck passing. 24
Muller returned to Bloomington with his fall dass awaiting his
attention. The story of the Geneva snub had not yet become public
knowledge. His manuscript, which he had sent before he left
Bloomington to the Bulletin of the Atomic Scientists, was to be
published in November. L. C. Dunn, who was editing this special
issue of the Bulletin, thought the ban was imposed by the AEC
because of the length and comprehensiveness of Muller's paper. Z5
The publicity following the Washington Post account of the
AEC ban on Muller's participation was embarrassing to the
Eisenhower administration. It was a clear case of the AEC' s having
"'H. J. MuHer, "Chronology relating to the paper .. . "
.. w. L. Russell to H. J. MuHer, July 19, 1955, Lilly Library.
"'L. C. Dunn to H. J. MuHer, July z8, 1955, Lilly Library.
3 620
The FaUout Controversy
been caught with a documented refutation of its charges by the
United Nations; and thus the AEC was forced to acknowledge its
deception or to somehow rationalize it away. At first the AEC did
not reply, and the furor gathered momentum. George Beadle,
then president of the AAAS, received a call on September 17 from
an AEC staff memher who mentioned the Washington Post story
and suggested to Beadle that the ban on MuHer was decided by
persons higher up than the Division of Biology and :\--ledicine.
These officials had initiated the withdrawal of the paper because of
their disapproval of Muller' s views of genetic damage from fallout.
Beadle feIt this was too highhanded an approach for a scientinc
issue and he proposed to Muller that he would take up the Geneva
incident at the next board meeting of the AAAS. 26 The National
Academy of Sciences was also considering a review of the radiation
controversy, but its role as an advisory hody to the federal govern-
me nt made it seem suspect, t::spedally since a high-level security
clearance was rumored to be aprerequisite for the participants
selected for the study. Beadle preferred a civilian, nonclassified,
approach to the assessment, and he wanted the AAAS to carry out
its own investigation of the genetic effeet of radiation.
Newsweek in its Oetober 3 issue presented a sharp criticism of
the AEC policy on Muller's paper. After consulting with i\ luller hy
phone, they cited his contempt for the AEC's "ostrich" poliey,
which ignored the realitY of genetic damage. They had learned of
Beadle's plans for the AAAS, and they also reported that India was
going to request a UN study of the hazards of nuclear radiation.
The AEC rejected Muller's speech, they reported, because it con-
tained references to Hiroshima and Nagasaki. 'i.7 This view was re-
peated on Octoher 3 hy Lewis Strauss, chairman of the AEC. He
may have hastened to call in the press after seeing the Newsweek
article . He daimed the work involved for the conference was so
overwhelming that he had not read NI uHer' s paper carefully and
had set it aside for rejection when he came across the references
to Hiroshima. He apologized for the "snafu" and hoped this would
end the incident. Strauss denied that "Dr. had been ex-
cluded, expelled, or worse from the Geneva Conference hy the
"'G. W. Beadle to H. J. :'v1uHer, September 17, 1955, LiHy Library.
rTNewsweek, October 3, 1955.
 Genes, Radiation, and Society
Commission, or specifically by me." After citing the reference to
Hiroshima as the reason for excluding the paper, he apologized for
not having "communicated with Dr. MuHer personaHy and asked
hirn to alter his paper." He claimed he was flooded with work for
the conference and, "as I say, I am sorry for it, and I take the full
personal responsibility for it. "28
No doubt Strauss hoped this would head off the AAAS attempt
to launch a study of radiation and help quiet the unexpectedly loud
clamor for information on radiation effects which had arisen in
response to the poorly handled censorship reported by the press.
M uHer never knew for certain how the decision to withdraw his
paper came about. He assumed thcrt the desire to play down radia-
tion damage was the principal force behind AEC poliey. To so me
extent this was true, but the major issue involved MuHer himself,
as I learned when I interviewed Willard F. Libby in 1971. 29
Libby was an AEC commissioner in 1955. He had established
his farne as a chemist, achieving national prominence for dating
archaeological materials by their carbon-14 content. This radioac-
tive element, which loses half its radioactivity every five thousand
years, serves as an indicator of age in organic relics. On ce an
organism dies it no longer recYcles, through metabolism, the car-
bon compounds in its cells. A wooden log of unknown age can be
dated by comparing its ratio of carbon-14 to nonradioactive carbon
(carbon-12). The Iower the ratio of carbon-14 to carbon-12, the
oider the log. Eventually this discovery led to a Nobel Prize for
Libbv. This and his interest in designing equipment to detect low
of radiation made hirn a valuable member of the AEC during
its weapons-testing program.
Libby was conservative in his political views. He despised J. R.
Oppenheimer from his graduate-school days at Berkeley when
"Statement bv Lewis L. Strauss at AEC Press Conference, October 3, 1955.
2OE. A. Carlsa'n, "Interview with Willard F. Libbyon H. J. MuHer and Radia-
tion Hazards," April 8, 1971, UCLA; manuscript copy filed at LiHy Library.
Muller wrote, "It happens that Libby was the man who intervened to prevent my
participation in the 1955 Geneva Conference, although I am not supposed to
know that" (H. J. Muller to C. W. Collier, April 11, 1958, Lilly Library). In
apparent contradiction is a letter to Muller attributing the ban to a minor screen-
ing official on Commissioner Strauss's staff (K. A. Cox to H. J. MuHer, April 23,
1959, Lilly Library).
The Fallout Controversy
Oppenheimer, he claimed, served as a recruiter for Communist-
front organizations. 30 During the war, Libby was active in the inter-
national development of the atomic bomb. He reguIarly flew to
England to discuss the progress made by his coHeagues at Harwell.
There was onIy one man he encountered who was more security-
conscious than he and whose reliability he thus thought was unim-
peachable: Klaus Fuchs! It was Fuchs, as wen as Bruno Pontecorvo
(apparently working independently), who sent some of the British
atomie-bomb secrets to the USSR. Fuchs, who studied physies
earIy during the war years at Edinburgh with :\'la.x Born, was even-
tually convicted of being a spy and served a long sentenee, but
Pontecorvo, who had studied with Fermi in Italy and JoHot-Curie
in Paris, fled to the USSR, where he became a professor of physics
at Moseow Universitv.
A eombination of eoincidenees may have built up in Libby' s
mind when he reviewed ;VI uller' s FBI file: M uller was reported to
be a Communist bv, the FBI or at least to have attended their
meetings in Austin just before his departure for Berlin; he was
known to be an editor and distributor of the Spark; he lived in the
USSR for almost four years, oecasionally writing seathing attaeks on
capitalism in his eugenic ::md popular essays; he volunteered to
serve with the Loyalists in Spain and worked in the Canadian blood
unit under Norman Bethune, a hero of :\-Iao Tse-tung's Eighth
Anny; also in the unit was one Allen Canadian with the
same name as Allen Nunn May, the English physicist who was
never in Spain but who was the first of the atomie spies to be
arrested. When M uner was settled in Edinburgh, who was his best
student? Guido Pontecorvo, brother of the atomie spy! Whether
the FBI or Willard Libby distinguished or confused these eoinci-
denees is still shrouded in secreey. JI
Other items may have been in the FBI file. vVhen Muller's
papers from Edinburgh were sent to Bloomington after the war
ended, they included all his pamphlets, correspondenee, journals,
and batches of propaganda leaflets whieh he had received as a
"'Carlson, "Interview with Willard F. Libby," p. 3.
31 For a detruled account of the backgrounds of Klaus Fuchs, AHen Nunn ylay,
and Bruno Pontecorvo, see Alan \-1oorehead's The Traitors (London: Hamish
Hamilton, 195z).
 Genes, Radiation, and Society
visiting scientist in the USSR. Since the Mullers had left Edin-
burgh on short notice, they never had a chance to sort out the
rubbish among their belongings. Some of the crates were never
opened; some they opened the lids of, but only to look for Muller's
reprints. When this shipment arrived in Inwanapolis in 1946 the
customs inspector remarked that he had never seen so much Com-
munist propaganda. 32 .
In Libbv's view, Muller could not be trusted. 33 Muller's anticom-
munism au'd his attacks on Lysenko may have been nothing more
than a cover, just as Fuchs had succeeded in camouflaging his
left-wing zeal with a passionate concern for security precuations.
But if this were true, what would be Muller' s role in addressing the
Geneva conference? Libby feared that concern over radiation dam-
age would be exploited to prevent the proper development of the
hydrogen bomb or to sabotage its further development. He also
feIt a personal hatred for communists and fellow travelers, whom
he equated with traitors, and he could not see anyone with Mul-
ler's past as an official U.S. delegate appointed by his own Atomic
Energy Commission.:J.I
This philosophy of "once burned, twice shy" prevailed, and the
recommendation to accept Muller's paper by the Division ofBiolo-
gy and Medicine was overruled. The AEC staff was not sufficiently
familiar with the radiation work which had been developed pains-
takingly in Drosophila since 1927; their deep suspicions of espion-
age and security risk were echoing from the cases of Fuchs, the
Rosenbergs, and Oppenheimer. This combination of circumstances
made it possible for the commissioners to serve as prosecutor,
judge, and jury in evaluating Muller's Geneva speech.
The political decision was, I believe, a bad error. Muller was un-
justly assigned a communist sympathy he no Ion ger had. His in-
tegrity as a geneticist was too high to permit its perversion for some
cryptic allegiance to the government that killed Vavilov, Levit, and
Agol and that dismantled the energetic research in basic genetics
which he worked so hard to build up. The decision was costly, tao,
because the AEC now found itself with far more public concern
J2Thea Muller, conversations. Bloomington, Indiana, summer 1973.
"'Carlson, "Interview with Willard .F. Libby," p. 2.
J<lbid.
The Fallout Controversy
over radiation damage than if Muller had been allowed to present
his moderate views. Muller accepted the accuracy of the data re-
leased by Strauss on the amount of fallout received worldwide, but
he denied that the small risk to the individual was of no conse-
quence to the population.
The weak public "apology" (which Strauss never sent as a person-
al message to Muller) also had a wholesome effect because it sug-
gested that a more liberal attitude would now prevail in the assess-
ment ofradiation damage. was pleased when he was asked
by the National Academy of Sciences to serve on its planned study
of the biological effects of radiation. This change in attitude among
scientists was brought horne to Muller by the AEC's Earl Green,
who assured hirn that the Division of Biology and .\-ledicine recog-
nized the risks of genetic damage.
The gag, however, made Green wonder whether the administra-
tive wing of the AEC had the same.awareness as the Division of
Biology and Medicine, especially since he had not known of the
reasons for the withdrawal, and he had assumed that the heart con-
dition wh ich MuHer mentioned in his first telephone conversation
was the reason. Green asserted that Muller's statements on fallout
had been welcomed as balanced and free ofbias by the members of
his division, and that the political, not the biological, authority was
in error in making the final judgment. 33
The Geneva affair was one of several episodes that eventually
led to Strauss's rejection by the D.S. Senate for a cabinet position
in the Eisenhower administration. The AEC no Ion ger pursued its
policy of absolutely denying the danger of low-dose radiation al-
though it properly minimized the individual risk from such ex-
posures. The public participated in a considerable amount of open
debate. The protection of the population became an important
issue and the demand increased for political, rather than military>
approaches to diminish East-West tensions.
JSEarl Green to H. J. Yluller. October 20, 1955, Lilly Library.