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Copyright 0 1995 by the Genetics Society of America

Anecdotal, Historical And Critical Commentaries on Genetics
Edited by J a m s F. Crow and William F. Dove

Two Genes, No Enzyme: A Second Look at BARBARA MCCLINTOCK

and the 1951 Cold Spring Harbor Symposium

Nathaniel C. Comfort
Cold Spring Harbor Laboratory, Cold Spring Harbor, New York 1I724 and Department of History,
State University of New York, Stony Brook, New York 11794

“I didn’t understand a word of it, but if BARBARA

it, it must be true!” With these words, the great
Drosophila geneticist ALFRED STURTEVANT

this essay isattributed by GREEN(1992) to ALFRED STUR-
says, had
Yet the only STURTEVANT listed
attended theSymposium.
a participant
as in 1951
gave his opinion of BARBARA MCCLINTOCK’S first public is one FRANK STURTEVANT, of Northwestern University!
presentation of transposable elements, at the 1951 Cold The error is quite excusable; while his is the only p u b
Spring HarborSymposium (GREEN 1992). A legend has lished version, this story has been repeated many times
sprunguparound MCCLINTOCK’S presentation, ac- by many people, and doubtless in many versions. It is
cording to which she gave her talk with great expecta- part of the legend, amythology that continues to grow,
tion of acceptance and interest, only to be ridiculed even among demythologizers.
and ignored by her colleagues. The subtext is that A re-examination of the 1951 Symposium can help
MCCLINTOCK was discriminated against, whether be- replace the myth with a more rational explanation of
cause of her views or her sex. In her biography of the reaction to MCCLINTOCK’S paper. It also shows how
MCCLINTOCK, EVELYNFOX-KELLER writes that MCCLIN- data and scientific theory are intertwined with the cul-
TOCK’S talk “was met with stony silence. With one or ture of science. MCCLINTOCK was not apassive recipient
two exceptions, no one understood. Afterward, there of her colleagues’judgments. She defiantly and deliber-
was mumbling-even some snickering-and outright ately challenged the paradigm viewof the gene-a
complaints. It was impossible to understand. What was gutsy move, but one that brought upon herself some
this woman up to?” (KELLER 1983, p. 139). of the confusion that so distressed her.
MEL GREEN,writing in the 1992 Festschrift for It is certainly true thatMCCLINTOCK’S work was highly
MCCLINTOCK, The Dynamic Genome, seeks to redress the respected by the time she presented her data on trans-
legend: “There is a widely extant viewpoint that BARBA- posable elements. Since the 1920s she had published
RA’S research was much unappreciatedand appropriate paper after importantpaper on maizecytogenetics.
recognition was too longdelayed. . . . I believe this view- With HARRIET CREIGHTON at Cornell, MCCLINTOCK pro-
point to be a half-truth” (GREEN1992, p. 117). Some duced the firstvisual evidence of crossing-over, just
of the legend springs from MCCLINTOCK’S own lips. In barely beating out CURTSTERN’S similar work on Dro-
an interviewwith KELLER, MCCLINTOCK said, “It was sophila. Subsequent work took her ever deeper into
just a surprise that I couldn’t communicate; it was a genomic instability and chromosome structure. First
surprise that I was being ridiculed, or being told that I she showed the existence of ring chromosomes. Then
was really mad” (KELLER 1983, p. 140). Unexplained, she showed that ring chromosomes were a special case
MCCLINTOCK’S cool reception supports the idea that of broken chromosomes, one in whichthe endsbecame
her colleagues were obtuse in failing to see the truth “sticky” and fused to each other. In 1936 MCCLINTOCK
and beauty of her discovery, or worse, that they refused moved to the UniversityofMissouri, where she was
to accept her results because MCCLINTOCK was outside hired byLEWIS STADLER,whowith R. A. EMERSON,
the geneticists’ old boys’ network. MCCLINTOCK’S mentor atCornell, was one of the reign-
Some of the legend seems to stem from scientists ing lions of maize genetics. At Missouri, MCCLINTOCK
wishing to squelch it. The quote at the beginning of discovered the “breakage-fusion-bridge” cycle, a fur-
ther extension of her observations of chromosomal in-
Address fw correspondace: Cold Spring Harbor Laboratory, 1 Bung- stability. The breakage-fusion-bridge cycle led MCCLIN-
town Road, Cold Spring Harbor, NY 11724. TOCK to two important conclusions, one a prediction

Genetics 1 4 0 1161-1166 (August, 1995)

more [before transposons], MCCI.INTO(:K would have
become a major figure in the history of genetics” (FED-
and the other hermost profound discovery. The obser- OROFF and BOTSTEIN1992).
vation of the “stickiness” of the broken ends of chro- The 1951 Symposium took place in the years between
mosomes led MCCLINTOCK to the prediction of special OSWALD AVERY’S demonstration that the transforming
structures ontheends of chromosomesneeded to principle in Pneumococcus wasDNA, and the double
maintainthe stability lost when thechromosomes revelations of ALFRED HERSHEY and MARTHA CHASE’S
broke. She called them telomeres, the study of which “blender experiment” andWATSONAND CRICK’S model
today is undergoing rapid growth. When MCCLINTOCK of the DNA double helix. The nature of the gene was
left Missouri for Cold Spring Harbor in 1941, she took the central questionin many biologists’ minds. Bacterio-
with her strains of maize that underwent the breakage- phage and bacteria were the model systems of choice,
fusion-bridge cycle. From them MCCLINTOCK deduced with Neurospora and Drosophila close behind. Scien-
the existence of transposable elements. The first set of tists working with these organisms were heavily invested
experiment$ in which she observed transposition were in the classical T. H. MORGANmodel of the gene. MOR-
done in 1944 (MCCI~INTOCK 1984). She spent the next GAN’S group had shown the enormous intellectual pro-
six years pursuing, confirmingand verifying her results. ductivity of viewing genes as unitary, particulate struc-
By 1951 MCCLINTOCK was among the best-respected tures, arranged linearly and statically along the
cytogeneticists in the country. Even ten years earlier, chromosomes. BEXDLEAND TATUM’S one-gene oneen-
when the director of the Carnegie Department, MILI- zyme hypothesis, first developed in the early 1940s, lay
SLAV DEMEREC, told staff scientist E. CARLETON MAG atop this foundation. It confirmed that geneswere uni-
DOU‘EILthat he had succeeded in hiring MCCI.INTOCK, tary by showing they had unitary effects on metabolic
MACDOWELI. “jumpedup in theairand said, ‘We pathways. MILISLAVDEMEREC,director of the Cold
should mark today’s date with red letters in the Depart- Spring Harbor Biological Laboratory and organizer o f
ment calendar!’ That expresses the feeling that is gen- the 1951 Symposium, acknowledged that the gene con-
eral among our members” (DEMEREC1942). In 1939 cept had been unraveling in recent years. Genes, DIG-
she was elected vice-president, and in 1945 president, MEREC said, “are regarded as much more loosely de-
of the Genetics Society of America. By 1951 she had fined parts of an aggregate, the chromosome, which in
received the Achievement Award of the Association of itself is a unit andreacts readily to certain changesin the
University Women,beenawarded two honoraryde- environment”(DEMEREC 1951). Whatever the gene’s
grees, and been a member of theNational Academy of precise molecularnature, the working model of the
Sciences for seven years. “The influence of her early gene was of a unitary entity that acted independently to
work is greater than that of any of her peers, with the produce a physiologically active molecule. While many
possible exception ofAL.FRED STURTEVANT,” wrote NINA at the time recognized that it was an oversimplification,
FEDOROFF and DAVIDBOTSTEIN.“Had she done no the view of the genomeas a static, linear series of partic-
Perspectives 1163

ulate, independently acting genes proved so successful of the original gene” (GOLDSCHMIDT 1951, p. 1).
in explaining genetic observations and generating new Though he cited examples drawn from work on muta-
experiments that few had cause to doubt it. ble genes in Drosophila and other organisms, promi-
One who did doubt it was R ~ C H A R DGOLDSCHMIDT. nent among GOLDSCHMIDT’S remarks are several glow-
For decades, GOLDSCHMIDT had been a gadfly to the ing mentions of MCCLINTOCK’S description of the Ac/
genetics community. Brilliant and cantankerous, GOLD- Ds system in maize. For GOLDSCHMIDT, transposable ele-
SCHMIDT delighted in challenging assumptions and ments provided a shining example of position effects
pointing out logical inconsistencies in the evolving ge- and a dynamic genome. He referred to transposable
netic theory. Since 1938, GOLDSCHMIDT had been ar- elements as “invisible” (i.e., submicroscopic) rear-
guing against the theory of the gene. His observations rangements and position effects. To his fullsatisfaction,
of Bar eye in Drosophila led him to conclude that the MCCLINTOCK had proved that in Zea, “mutable loci are
chromosome, not the gene,was the unitary element of actually position effects produced by genetically con-
heredity (GOLDSCHMIDT 1938; DUNN1965). GOLD- trolled and repeating transpositions and transloca-
SCHMIDT argued from translocation data that the posi- tions” (GOLDSCHMIDT 1951, p. 4). GOLDSCHMIDT waxed
tion of a locus on the chromosome determined its func- poetic in his analogy for how genetic function derived
tion. When a locus moved to a different site, its function from position: “If the A-string on a violin is stopped an
changed. GOLDSCHMIDT argued in 1938 that “the whole inch from the end the tone C is produced. Something
conception of the gene”was “obsolete.” GOLDSCHMIDT has been done to a locus in the string, it has been
and BEADLE took up opposing sides in a debate that changed in regard to its function. But nobody would
lasted over a decade. GOLDSCHMIDT’S was a dynamic conclude that there is a Gbody at that point” (GOLD-
genome, not a static one. By playing the devil’s advo- SCHMIDT 1951, p. 7 ) . MCCLINTOCK’S data must have
cate, GOLDSCHMIDT forced geneticists to reconsider seemed a godsend to GOLDSCHMIDT, an example tailor-
their assumptions. Though his style was combative and made to support his attack on the gene concept.
his views extreme, position effects havebeen supported In her own paper, directly following GOLDSCHMIDT’S
by molecular analyses. To besure, GOLDSCHMIDT in the Symposium volume, MCCLINTOCK returned the
seemed to set up a straw man; his notion of the prevail- favor.Bothimplicitly and explicitly, she repeatedly
ing gene concept seemed to be a globular molecule aligned herself with GOLDSCHMIDT. Twenty-three pages
situated on the chromosome, an almost literal “bead into her paper, she said, “It will be noted that use
on a string.” This caricature doubtless made it easier of the term gene has been avoided in the foregoing
for him to ridicule the gene theory, but it also made discussion of instability”(MCCLINTOCK1951, p. 36).
his alternative impossible for most geneticists to accept. She went on to say that this “does not imply a denialof
For all his crotchetiness, GOLDSCHMIDT was and is an the existence within chromosomes of units or elements
important figure in genetics. DEMEREC gave GOLD- having specific functions. The evidence for such units
SCHMIDT the honor of presenting the opening talk of seems clear.” But the gene concept, she said, “stems
the 1951 Symposiumin a special session entitled “The- from studies of mutation.” She then made the link
ory of the Gene.” The session was filled out by MCCLIN- with GOLDSCHMIDT explicit: “The author agrees with
TOCK herself, LEWISJ.STADLER, who also critiqued the Goldschmidt that itis not possible to arrive at any clear
gene model, but in a way less infuriating to geneticists, understanding of the nature of a gene, or the nature
and N. H. HOROWITZ and URS LEUPOLD, speaking on of a changein a gene,from mutational evidence alone”
new evidence for and implications of the one-gene one- (MCCLINTOCK 1951, p. 36). MCCLINTOCK invoked
enzyme theory. The “Theory of the Gene” session thus GOLDSCHMIDT again in her conclusion: “Evidence, de-
had three scientists critical of the standard model and rived from Drosophila experimentation, of the influ-
one supportive of it. Much of the rest of the meeting ences of various knownmodifiers on expression of phe-
was given over to the microbial geneticists. Three ses- notypic characters has led Goldschmidt (1949, 1951)
sions weredevoted entirely to bacteriophage and bacte- to conclusions that are essentially similar to those given
ria, with manymore papers on microbial genetics sprin- here” (MCCLINTOCK 1951, p. 46). In thebattle between
kled throughout the remaining sessions. Most of the GOLDSCHMIDT and BEADLE,it could not have been more
remainder concerned Drosophila or Neurospora. On plain with whom MCCLINTOCK allied herself.
both sides of the podium, discussion was dominated by It is clear what appealed to MCCLINTOCK in GOLD
classical genetics, microorganisms, and the one-gene SCHMIDT’S work. Like MCCLINTOCK, GOLDSCHMIDT was
one-enzyme vision of genetic function. arguing for a dynamic genome, a self-regulating system
In theopeningpaper of the Symposium, GOLD- defined by the interactions among its parts.To both,the
SCHMIDT revisited the arguments many had heard be- static concept of linear beads on a string, though it had
fore. He had new evidence to support it, however. He enormous predictive value,was a vast oversimplification.
took on BEADLE’Smodel early on, accusing BEADLEof Yet MCCLINTOCK undersold herself somewhat by align-
“extrapolation from the mutantaction to the existence ing herself so closely withGOLDSCHMIDT. Her model was
1164 N. C. Comfort

considerably more sophisticated than GOLDSCHMIDT’S. containapentosenucleoproteinthat continually re-

Unlike GOLDSCHMIDT, MCCLINTOCK believed in genes; leases polynucleotide into the cytoplasm.Thispolynu-
she just didn’t believe they were the entire story. Her cleotide, presumably RNA, was presented in the context
“controlling elements” were presented as a new sort of of the one-gene one-enzyme theory as direct evidence
genetic material, not genes but regulators of genes. that unitary genes have discrete, independent effects in
While GOLDSCHMIDT deniedthe BEADLE and TATUM the cytoplasm. MARSHAK suggested that MCCLINTOCK’S
model, MCCLINTOCK was working outside of it. Activatorwas involved with the formation of nucleopro-
Equally clear, however, iswhy MCCLINTOCK’S work teins from raw histones and polynucleotides, much like
would have receiveda cool reaction from the assembled a conventional gene. MCCLINTOCK’S data, he says, are
audience. The differences between MCCLINTOCK’S and readily explained on the basis of the hypothesis ifwe
GOLDSCHMIDT’S views wouldhave been obscured by suppose that Ds and the other similar factors represent
their obvious mutual admiration. Hadn’t MCCLINTOCK chromosome segments containing greater amounts of
avoidedeven the use of the term gene for the first DNA than those which were previously associated with
two thirds of her talk? Hadn’t she concludedby saying the locus in question. . . . If the “activator” is involved
in the production of the pentosenucleoprotein[,] as for
GOLDSCHMIDT’S conclusions were essectially the same example from raw materials such as histones and poly-
as her own? To an audiencefull of Neurosporists, Dro- nucleotidesstoredin the nucleolus[,] an alterationin
sophilists and phageologists, it must have seemed that the amount of the substance we have assumed to govern
they had just had to endure their second GOLDSCHMIDT the rate of activity of the locus will result and hence a
talk! mutated condition: or if the intensity of the activator
action varies, in a mutable condition. In this connection
Although many geneticists were relaxing their con- it appears significant that the simultaneous presence in
cept of the gene,all wereemploying it. Whatever a gene the genome of a Ds or Ds-like region together with an
was, it was unitary and it acted independently. Eleven “activator” results in a “fusion-breakage event” which
papers in the Symposium concerned the mutagenic ef- further suggests involvement of transport andutilization
fects of radiation and chemicals. Further, “mutation” of nucleolar materials (MARSHAK1951, p. 157).
and “gene” were defined in ways that were internally In this rather convoluted passage, MARSHAK attempts
consistent but entirely self-contained. A mutation dis- to explain MCCLINTOCK’S observations from a BEADLE-
ruptedagene. In his paper on pseudoallelism and and-TATUM point of view. Although MCCLINTOCK em-
crossing over, E. B. LEWISdefined the geneas a physical phasized that Ac and Ds were not conventional genes,
unit within which there is no crossing over. A static, MARSHAK’S explanation hinges on Ac being involved in
discrete unit was inherent in his definition: “The defi- the production of nucleic acids-in short, acting like
nition of any particulate unit must be in terms of its a gene. Significantly, MARSHAK says that “while other
indestructibility by some breakage or splitting process” interpretations may seem equally plausible, this one . . .
(LEWIS1951, p. 174). LEWIS’Sconcept of a gene com- is amenable to being tested experimentally,” suggesting
plex became a paradigm in Drosophila developmental that MCCLINTOCK’S interpretation is not.
genetics, but the regulation of the bithorax complex In their paper on chemically induced mutations in
by chromosomal components has proved to be a key Neurospora, K. A. JENSEN,INGERKIRK, G. KOLMARK,
element in understanding its operation. While GOLD- and M. WESTERGAARD offer a different interpretation
SCHMIDT and MCCLINTOCK’S 1951 visionof genetic reg- of MCCLINTOCK’S results. They cite MCCLINTOCK’S con-
ulation differs from the modern conception, the paral- trolling elements as evidence for their theory that muta-
lels are in many ways more striking than thedifferences. ble genes might result from unstable precursors accu-
MCCLINTOCK’S controlling elements were non-genes, mulating - in the nucleus, the result of blocked steps in
genetic particles outside the framework of one-gene a BEADLEand TATUM-Style biochemical pathway:
one-enzyme biochemical genetics.
We know from the work on biochemical mutants of
MCCLINTOCKdid not go unnoticed in the Sympo- Neurospora that when a specific intermediate step in the
sium. Several speakers in later sessions referred to her synthesis of a vitamin or an amino acid is blocked by
work, either in their papers or in the reprinted discus- mutation, a precursor may accumulate in the cells. We
sion that followedmanyof the papers. STADLER,of also know that some of these precursors are highly stable
course, cited her graciously,though hedid so nonanalyt- and may be extractedfrom themycelium, and that others
are very unstable and areconverted into by-products, like
ically.JACKSCHULTZcited her 1933 work on association the purple pigment in our adenineless strain, whereas
of nonhomologous chromosomes during prophase of others disappear completely. We want to suggest that
meiosis and NORMANGILEScited her seminal 1945 pa- some of the unstable precursors may act as mutagens
per on thecytology of Neurospora. Two others referred during the decomposition. This would be a very specific
to MCCLINTOCK’S work on transposons. ALFRED MAR- type of mutagens [sic], since, as primary gene products,
they may be produced within the nucleus. It does not
SHAK, commenting on DAVIDBONNER’S paper on gene- seem improbable that some of these precursors would
enzyme relationships in Neurospora, cited MCCLIN- be so unstable that they may only react over a very short
TOCK’S work in connection with observations that nuclei distance, thus inducing mutations in loci only within a
Perspectives 11
short distance from the “blocked” gene. The so called systems.” She wrote that “it was the patternof behavior,
unstableormutablelocinow under investigation by rather than the change in expression of the particular
precursor” mechanism UENSENet al. 1951, p. 258). phenotypic character,that was obviouslyof impor-
tance” (MCCLINTOCK 1951, p. 43). She felt it was neces-
Neither ~ ~ R S H AnorK JENSEN et al. ignored MCCLIN- sary “to consider these various widely different levels
TOCK’S work; neither, however, accepted her interpreta- of unitary control and how they may operate in the
tion of her data. MARSHAKand JENSEN were working working nucleus, and also to consider the natureof the
within the dominant paradigm of the BEADLEand TA- changes that can affect their operation” (MCCLINTOCK
TUM model, and attemptedto jam MCCLINTOCK’S 1951, p. 37). This was pretty cosmic stuffto a biochemi-
anomalouscontrollingelementsintothat prevailing cal geneticist or phage group member.
and familiar conceptual framework. These examples The controversy between the MCCLINTOCK/GOLD-
neither illustrate cloddishness on the part of MARSHAK SCHMIDT view of the gene and the BEADLE/TATUM view
or JENSEN, nor demonstrate that MARSHAK and JENSEN was resolved withthe advent of modern molecular tech-
“misunderstood” MCCLINTOCK’Swork.Rather, they niques. As is often the case, both sides were partlyright.
suggest scientists who listened to and believed MCCLIN- Transposition became widely accepted in the 1970s as
TOCK’S data but thoughther interpretation overly spec- SAEDLER, STARLINGER, SHAPIRO, BOTSTEIN,FINK,FEDOR-
ulative and even untestable. E. B. LEWIShad a similar OFF and others identified transposable elements in mi-
reaction. He said of MCCLINTOCK, “Everybody recog- crobial and animal systems and elucidated the mecha-
nized her as a great scientist. You could trust what she nism of transposition. It became clear that transposable
found out experimentally, but it was the speculation elements can encode proteins that regulate other
that gotout of hand. This reinforced her mystical ideas. genes. Ac was shown to encode transposase, which acti-
Youjust couldn’t fit it into the standard genetic theory” vates Ds.Genome instability, position effects, and geno-
(E. B. LEWIS,personal communication). LEWISeven mic regulation have become hot topics as molecular
suggests that MCCLINTOCK and GOLDSCHMIDT may have techniques allow confirmation and modification of
been seen as supporting the anti-genetic stance of T. D. MCCLINTOCK’S ideas. MCCLINTOCK’S vision of biology
LYSENKO-a view MCCLINTOCK and GOLDSCHMIDT needed to be filtered through the accepted methodol-
would have found absurd, but one that illustrates how ogy and language of genetics and molecular biology in
far out they seemed to mainstream geneticists. MCCLIN- order forit to significantly influence our understanding
TOCK’S peers were struggling to re-interpret anomalous of the gene. Further, thenew techniques made it possi-
data into terms with which they were comfortable. ble to confirm what had seemed speculations, and to
Other factors doubtless also contributed to theconfu- do it in multiple species.
sion about and lack of acceptance of MCCLINTOCK’S MCCLINTOCK was universally admired as an experi-
conclusions. Her model systemmayhave had some mentalist, but to some she seemed to be extrapolating
alienating effect, but it would seem to have been small. too much from her data. Those unfamiliar with her
True, STADLER was the only other maize geneticist on work probably associated her paper with the one pre-
the program, yet there were several other plant geneti- ceding it and considered MCCLINTOCK just another
cists and severalcytologists.And although maize was Goldschmidtian. MCCLINTOCK herself encouraged this
becoming an unfashionable organism in genetics, it perception by explicitly aligning herself withthe notori-
wasn’t so far out ofstyle as to be incomprehensible. ously controversial GOLDSCHMIDT. And GOLDSCHMIDT
MCCLINTOCK’S presentation may have been something was, in fact, the closest to MCCLINTOCK in his concept
of a barrier,however. Non-trivial, surely, was her paper’s of the gene. But unlike GOLDSCHMIDT, MCCLINTOCK
considerable length. Itfilled 35 pages in the Symposium didn’trefutethe paradigm-she stepped outside it.
volume and was by far the longest of anyat themeeting. Among those who knew and respected her, some tried
MCCLINTOCK presented a welter of data, with long sec- to squeeze her findings into their conception of the
tions of background on each important mutation and gene, while some, like STURTEVANT-Or whoever it
detailed descriptions of her crosses. Her talk was essen- was-didn’t understand a word of what she said.
tially historical in structure, rather than the more con-
Thanks to ROB MARTIENSSEN for critical reading and discussion of
ventional format of Introduction, Results and Conclu- the manuscript, to CAROLGREIDER for critical reading and more,
sion, so it tended to ramble. Also, she used highly and to CLARE BUNCEfor archival research. The portraits of BARBARA
unconventional language. Beyond dispensing with the MCCLINTOCK and RICHARD GOLDSCHMIDT were provided by the Cold
word “gene,” she invented new terms to describe her Spring Harbor Laboratory Research Library Archives.
new phenomena. In addition to “activators” and “dis-
sociators”-terms that even those scientists who tack- LITERATURE CITED
led her data handled withverbalkidgloves, such as
DEMEREC,M., 1942 Letter to W. M. GILBERT,Executive Director of
quotation marks and “so-cal1eds””she spoke of “al- the Carnegie Institution of Washington, 6 April 1942. MCCLIN-
tered states,” “out-of-phase timing,” and “controlling TOCK file, Cold Spring Harbor Laboratory Archives.
1166 Comfort N. C.

DEMEREC, M., 1951 Foreword. Cold Spring Harbor Symp. Quant. o f h e t i c s , edited by N. FEDOROFFand D. BOTSTEIN. Cold Spring
Biol. 16: v. Harbor Laboratory Press, Cold Spring Harbor, NY.
DUNN,L. C., 1965 A ShortHistmy ofGenetics McGraw-Hill, NewYork. JENSEN,K.A.,I.KIRK,G.KOLMARK~~~M.WESTERGAARD,~~~~ Chemi-
FEDOROFF, N., and D. BOTSTEIN, 1992 Introduction, pp. 1-4 in The cally induced mutations in Neurospora. Cold Spring Harbor
Qnamic Genome: Barbara McClintock ‘s Ideas in the Centu? of Genet- Symp. Quant. Biol. 16: 245-261.
ics, edited by N. FEDOROFFand D. BOTSTEIN. Cold Spring Harbor KEI.I.ER, E. F., 1983 A Feeling for the Organism. W. H. Freeman, New
Laboratory Press, Cold Spring Harbor, N Y . York.
LEWIS,E. B., 1951 Pseudoallelism and gene evolution. Cold Spring
GOLDSCHMIDT, R., 1938 Physiologzcul
Genetics. McGraw-Hill,New
Harbor Symp. Quant. Biol. 16: 159-174.
York. MARSHAK, A,,1951 Discussion. Cold Spring Harbor Symp. Quant.
GOLDSCHMIDT, R.,1951 Chromosomes and genes. Cold Spring Har- Biol. 16: 156-157.
bor Symp. Quant. Biol. 16: 1-11. MCCLINTOCK, B., 1951 Chromosome organization and genic ex-
GREEN,M., 1992 Annals of mobile DNA elements in Drosophila. pression. Cold Spring Harbor Symp. Quant. Biol. 16: 13-47.
The impact and influence of Barbara McClintock, pp. 117-122 MCCLINTOCK, B., 1984 The significance of responses of the genome
in The Dynamic Genome: Barbara McClintock’s Ideas in the Centuly to challenge. Science 226: 792-801.