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OCR for page 443
CURT STERN
August30, 1902-October23, 1981
BY JAMES V. NEEL
TH E R E H A V E B E E N three remarkable periods in the his-
tory of modern genetics. The first of these, in the early
years of this century, encompasses the rediscovery and
confirmation of Mendel's findings and the enunciation of the
chromosomal theory of inheritance. The seconct period is
characterizes! by a concatenation of discoveries regarding the
more precise mechanisms of chromosomal behavior, largely
based on the use of Drosophila and emanating from the "fly
room" at Columbia University. The third is the period in
which we now fins! ourselves, initiated in the 1940s by the
seminal observations of Avery and collaborators and the later
phage work demonstrating that the essential genetic material
was DNA and the demonstration, using Neurospora, that
genes have essential anc! specific roles in the synthesis of pro-
teins. This was followed by the elucidation of the structure
of DNA, leading to a cascade of discoveries concerning DNA
fine structure and how it can be manipulated. Each of these
flowerings clominatect the conceptual biological thinking of
the time.
'NOTE: This memoir is an expanded version of a manuscript published in the An-
,~ual Review of Genetics. I am deeply indebted to the many people who have contrib-
uted their remembrances and perspective to the writing of this memoir.
443
OCR for page 444
444
BIOGRAPHICAL MEMOIRS
THE EARLY YEARS
Curt Stern, the subject of this memoir, must be regarclect
as one of the principal participants (ancl last survivors) of the
fly room period. Born in Hamburg, Germany, on August
30, 1902, the first son of Barned anct Anna Stern, he early
displayed a strong interest in natural history, ranging from
microscopic studies of pondwater to the zoological collections
that enliven parental responsibilities. In these interests he
received unusual support from two extraordinary high
school teachers, who encouraged him to undertake the study
of zoology. His father, who was in the dental supply business,
and his mother, a schoolteacher, were also highly supportive
as their son's biological interests unfoIclecl. Following the
family's move to a suburb of Berlin, he entered the University
of Berlin in 1920. He must almost immediately have found
his way to the Kaiser Wilhelm Institute, where he conductect
his doctoral studies. He received a Ph.D. from the University
of Berlin in 1923, at that time the youngest person ever to
receive the degree from the university. There is no record of
an unclergraduate degree. Attendance at the university re-
quirec! one to two hours of commuting each way. To achieve
his degree so early under these circumstances was an early,
clear signal of the remarkable combination of high intellec-
tual ability, photographic memory, and stamina that was to
characterize his career.
Stern's Ph.D. thesis was a descriptive cytological study of
mitosis in a protozoan of the order Heliozoa, uncler the di-
rection of one of the truly eminent protozoologists of that
era, Max Hartmann. One perceives here the momentum of
a high school fascination with microscopic pond life. By the
time he finished his thesis, he knew he would not remain a
' The only remaining survivor of the "fly room" known to the author is Helen
Redfield (firs. Jack) Schultz.
OCR for page 445
CURT STERN
445
protozoologist. Reading widely, he became intrigued by ge-
netics. A paper by Richard Goldschmidt, then director of the
Institute, on the basis for crossing-over caught his attention.
Stern felt Goldschmidt's interpretation was incorrect and
wrote a critique, which, after great hesitation, he submitted
to Goldschmidt. (He was at that time probably the most jun-
ior fellow at the Kaiser Wilhelm Institute.) Some six months
later, Goldschmidt returned the paper without comment, but
shortly thereafter called Stern into his office. Those of us
privileged to see Goldschmidt in action after his immigration
to the United States will recall the personification at least
superficially of the Geheimrat Professor. One can imagine
the trepidation with which Stern approached the meeting.
With few preliminaries, Goldschmidt offered Stern a fellow-
ship, recently funded by the International Education Board
of the Rockefeller Foundation, to study genetics with the
Morgan group. There apparently was never any discussion
of the critique.
DR OS OPHILA
Stern arrived at Columbia in 1924. The fly room was near
or at its zenith. Morgan in ~ 9 ~ 0 had demonstrated sex-linked
inheritance in Drosophila and then the recombination of two
sex-linked alleles, one responsible for white eye and the other
for rudimentary wing. These exciting discoveries, along with
the obvious potentiality of Drosophila as an experimental or-
ganism, had attracted to Morgan's laboratory a now historic
triumvirate Sturtevant, Bridges, and Muller. The publica-
tion by Morgan, ~ ' ~3
biturtevant, Muller, and Bridges of "The
Mechanisms of Mendelian Heredity" in 1915, which brought
together the early data on autosomal as well as X-chromo-
some inheritance and linkage, with linkage maps and the evi-
dence for nondisjunction, had made clear how major the de-
velopments at Columbia were. One result was a rising
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446
BIOGRAPHICAL MEMOIRS
number of students and visiting investigators, who, together
with Sturtevant, Muller, and Bridges, were all crowcled into
a room 16 by 23 feet in size, which contained eight working
desks! (For a forthright statement of the physical "together-
ness" of this setting, and especially the space allotted to Stern,
see Provine ~ 98 ~ .)
There quickly ensucc! several "beginner papers," but the
first major result of this fellowship, published in 1926-1927,
was the demonstration by a combination of genetic and cy-
tological techniques that the anomalous genetic behavior of
the "bobbed bristles" trait couIcT be explained by the Y-linkecl
inheritance of the responsible allele (plus a homologous locus
on the X). Until then, the Y-chromosome of Drosophila, al-
though associated with male fertility, had been consiclered as
otherwise genetically empty. Although Y-linkocI inheritance
had previously been demonstratecI by Schmidt in the fish,
Lebistes reticulates, this demonstration that abnormalities of Y-
chromosome behavior (that is, the occurrence of XXY fe
males) accounted for abnormalities in the inheritance of the
bobbed trait was unusually elegant for the times, no doubt
benefiting greatly from the cytological clemancis of Stern's
Ph.D. thesis.
His second major contribution appeared in 1931: the
demonstration,
a ~ 1
using cytologically abnormal X-chromo-
somes, one with an X-Y transIocation, one with an X-IV
transIocation, that the genetic phenomenon of crossing-over
was accompanies! by a physical exchange between the chro-
mosomes. (Simultaneously, Creighton and McClintock quite
independently clemonstrated the same phenomenon in
corn.) This was shortly followed by an ingenious clemonstra-
tion (back to the Y-chromosome) that as he acIdect supernu-
merary Y-chromosomes bearing the bobbed allele to Droso-
phila, the trait gradually ctisappearect, understandable now
that we know the bobber] alleles are characterized by varying
OCR for page 447
C U RT S T E RN
447
degrees of unclerproduction of ribosomal RNA. At that time
the demonstration that adding enough defective genes wouIcI
produce normality was novel.
Stern's last major contribution to our understanding of
the chromosomal basis of inheritance was published in 1936.
He had returned to Germany in 1926, but had come back to
the United States in 1932, on a second fellowship from the
Rockefeller Foundation, spending the year at the California
Institute of Technology in the company of a remarkable
collection of geneticists: Morgan, Sturtevant, Dobzhansky,
Bridges, Schultz, Emerson, Darlington, Kaufmann, and Lin-
degren. He had married an American citizen, Evelyn Som-
merfield, in 1931. In 1933, when he was clue to return to
Germany, Hitler came to power, a clevelopment whose tragic
implications for German Jews has been only too well docu-
mentecI. While Stern remained in the United States, Evelyn
returned to Germany to seek some cautious acivice from his
colleagues. What she Earned convinced them it wouIcI be
wise not to return. (He became a U.S. citizen in 1939.) Stern
accepted a temporary position at Western Reserve University,
but quickly moved to the University of Rochester, where he
was to remain until 1947, serving from 1941 to 1947 both as
chairman of the Department of Zoology and chairman of the
Division of Biological Sciences.
Shortly after he arrived at Rochester, he began to inves-
tigate what was then a puzzling phenomenon: the occurrence
in female flies heterozygous for one or several sex-linkoct re-
cessive alleles of epidermal spots manifesting the effects of
one or all of these alleles or even, if the alleles were on ctif-
ferent but homologous chromosomes, "twin spots," exhibit-
ing the phenotypes associated with both alleles. A classical
analysis revealed that the only consistent explanation re-
quired a previously unrecognized phenomenon, mitotic
crossing-over. ~ wouIcI like to suggest that this paper was the
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448
BIOGRAPHICAL MEMOIRS
ens! of the "chromosomal era" in the history of Drosophila as
an experimental organism. It was to continue to provide in-
sights on other problems, but the story of how its chromo-
somes behaved (in the classical sense) was now essentially
complete.
In several autobiographical sketches, Stern has empha-
sizect that he never attempted any grand research design, but
simply "followocl his nose." The beauty of Drosophila was how
quickly one could move from one major issue to the next,
providing you knew how to manipulate the fly stocks at
which Stern was probably seconc! only to Muller. That nose
led him unerringly to basic issues. Space permits mention of
only three of the outstanding contributions subsequent to
1936 that depended on experiments with Drosophila.
T. Isoalleles. The recessive allele cubitus interruptus (ci), lo-
cated in the fourth chromosome, causes a gap of variable
length in the fourth wing vein. Working with strains thought
to be isogenic except for fourth chromosomes of different
origins, as well as with strains in which a deficiency of the
region encompassing the ci locus was present, and manipu-
lating temperature, Stern in 1943 demonstratecl that normal
alleles of ci differed greatly in their potency, as measured by
their ability to mollify the expression of the ci trait in hetero-
zygotes or hemizygotes for this locus. He termed these dif-
ferent normal alleles "isoalleles." This demonstration of a
range of genetic variation beyond that easily envisioned pre-
saged (and may now find an explanation in) the demonstra-
tion years later of extensive inapparent biochemical variation.
At that time there was still concern amounting to clisbelief
among some biologists and paleobiologists that the kinds of
traits arising through mutation in Drosophila couIct possibly
serve as the stuff of evolution. Since these alleles of small
effect presumably arose from the mutational process, this cli-
rection of attention to traits of lesser effects player! a signifi
OCR for page 449
CURT STERN
449
cant role in what Mayr has termec! "the evolutionary synthe-
sis" (Mayr and Provine ~9801.
2. Genetic effects of low-level racliation. During and after
World War Il. Stern, in collaboration with Spencer, Caspari,
and Uphoff, was drawn into studies of low-level radiation
effects, studies sponsored by the Rochester branch of the U.S.
Army's Manhattan Engineering District. The question, in the
context of the advent of the atomic bomb, was obvious: Was
there a threshold in the genetic effects of radiation? The fincI-
ing now a cornerstone of racliation genetics was clear:
"Viewing all experiments together, it appears that radiation
at low doses, administered at low intensity, induces mutation
in Drosophila sperm. There is no threshold below which ra-
cliation fails to induce mutations" (Uphoff and Stern 19491.
It was un(loubtedly this background that led to a term ~ ~ 950-
1953) on the Advisory Committee to the Division of Biology
anc! Medicine of the Atomic Energy Commission, a critical
period in the (development of the AEC's policy of broadly
basect research into radiation effects.
The work on the effects of low-level radiation hac! used
sex-linkocI lethals as indicator traits. Since it was felt that ra-
cliation produces disproportionate numbers of lethals (as the
normal spectrum of mutation is unclerstood), an important
question was the effect of these "recessives" when heterozy-
gous. Given the ratio of heterozygotes to homozygotes pre-
clictect by the Harcly-Weinberg formulation, for a rare allele
even a small heterozygote effect for an autosomally inherited
recessive lethal coup outweigh the impact of the occasional
homozygote. Stern-with Carson, Kinst, Novitski, and Up-
hoff in 1952 established that under their conditions, the
average viability of heterozygotes for sex-linkecl lethals was
96.5 percent normal, a figure still standard. Their ciata clict
not permit any distinction between the effects on viability of
spontaneous anc! inclucecI mutation.
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450
BIOGRAPHICAL MEMOIRS
3. The extent of cell autonomy in gene expression and the genetic
control of patterns. In 1947, Stern left Rochester to become
professor of zoology an(l, in 1958, professor of genetics as
well, at the University of California, Berkeley; he retirect
from these positions in 1970. At Berkeley he returned to an
oIc] problem; the basis of the attack had been laid with the
1936 paper on somatic crossing-over and segregation. These
.
studies had demonstrated a high degree of cell autonomy In
the expression of genetic constitution in Drosophila. On the
other hanct, transplantation experiments (with Hadorn) hac!
demonstrated that the color of the vase efferentia was depen-
clent on that of the testis attacher! to them and not on the
genetic constitution of the ducts themselves, and that the
shape of the testis (spiral or oval) was dependent on the ge-
netic constitution of the sperm ducts to which it was attached
~ ~ 939-l 94 ~ ). In the ~ 950s ant! much of the ~ 960s, most of
Stern's research efforts, often in collaboration with his stu-
clents, were directed toward the difficult problem of the ge-
netic control of differentiation, and especially of patterns, still
using Drosophila as an experimental organism.
The various types of regularly arranged chaetae, so ob-
v~ous when one inspects a fly, proved most useful in these
interests. Studies with Hannah-Alava, employing genetic mo-
saics of various derivations, demonstrated a new level of com-
plexity in embryological determination: Differentiation of
the male sex-comb (a specialized row of chaetae) clependec!
on a field effect within which the development of the sex
.
comb teeth was cletermined down to very small patches of
cells by the sex (maleness) of the cells. These and other
studies led to what Stern in 1954 termed the "prepattern
hypothesis." Prepattern was a descriptive term for any kind
of spatial differentiation in development, development being
regarded as a succession of prepatterns. Within the prepat
OCR for page 451
CURT STERN
451
tern there are singularities, to which developing cells respond
according to their genetic competences. Much of his later
work on this subject, well summarized in 1978 by his col-
league and collaborator Dr. C. Tokunaga, was directed to-
war(1 defining, largely through the use of genetic mosaics,
the nature of prepatterns, singularities, and competences.
The familiarity with Drosophila mosaics these inquiries cle-
mancled inspired an omnivorous interest in mosaics of all
types, reflected in the Prather Lectures at Harvard in 1965
in which he summarized his activities in this fielcI.
One of Stern's last papers on Drosophila, in 1969, once
again illustrated his ability to take full advantage of Drosophila
as an experimental organism. The earlier studies on somatic
cell crossing-over hacI not provicled a clear approach to the
relative frequency of somatic cell versus meiotic crossing-
over. Since then, in the 1950s, compound or complex loci
hacI been recognized in Drosophila melanogaster, such as the
"white eye" (w) locus, within which meiotic crossing-over
could occur. As a manifestation of the remarkable degree of
cell autonomy in Drosophila, each incliviclual facet of its com-
pound eye expresses its genotype as regards pigmentation
independently of the other. Stern scored female flies heter-
ozygous for two different w alleles between whose mutational
sites the frequency of meiotic crossing-over had been cleter-
minect, for red-colored spots in their white eyes. Four such
spots were found in a total of 6,137 flies. No such spots were
observed in the eyes of 27,557 controls. On the assumption
that the rect spots resulted from somatic cell recombination
in a cell of the developing eye disc, and that the eyes of the
flies scored collectively provided a minimum of 9 x lob mi-
toses in which the results of somatic recombination couIct be
observecl, these 4 spots suggested a frequency of recombi-
nation between these mutational sites of less than ~ or 2 in
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452
BIOGRAPHICAL MEMOIRS
2 x lO`; mitoses (! if the exchange was nonreciprocal, 2 if
reciprocal). This was 400 to 800 times less common than
. . · · .
melotlc crossing-over in t ne same region.
HUMAN GENETICS
Technically, Stern's advent into the other fielc! of genetics
with which his name is so prominently associated, human
genetics, dates from a paper entitlecl "Welche Moglichkeiten
die Ergebnisse der experimentellen Vererbungslehre ciafur,
class clurch verschieclens Symptome charakterisierte Ner-
venkrankheiten auf gleicher erblicher Grundlage beruhen?"
published in 1928 in Nervenarzt. It is a very clear statement,
directed to physicians, of a principle now commonly ac-
ceptecI: Indistinguishable phenotypes may have very (liffer-
ent genetic bases. His serious entry into the fielct, however,
can be ciated more precisely to 1939. That year he supervised
his first graduate student seminar in the field of human ge-
netics. As one of the half-clozen students who met weekly for
a semester, T still have the list of papers he chose for review.
Two impressions stand out. First, he hac! manager! to select
what little solid ciata existed; the contrast between the then
anct the now of human genetics, developments within a single
generation, is simply staggering. Second, given the excesses
of American eugenicists, and especially the Nazi racism from
which he personally had suffered, one might have expected
some bitterness and an occasional diatribe on these mon-
strous perversions; Stern kept the discussion all science.
This occasion must mark the beginning of the most suc-
cessful textbook on human genetics ever written. In the
course of three English editions ~ 1949, ~ 960,1973),Principles
of Human Genetics sold 62,337 copies; there is no way to esti-
mate accurately the number of copies soIct of the German,
Japanese, Spanish, Portuguese, Hincti, Polish, and Russian
OCR for page 463
CURT STERN
463
The determination of color in the vase efferentia of Drosophila me-
lanogaster. Science, 86:408.
Interaction between cell nucleus and cytoplasm. Nature, 140:770-
71.
Methylene blue-staining of peripheral nervous system in Droso-
phila melanogaster. DIS, 8:90.
1938
With E. Hadorn. The determination of sterility in Drosophila males
without a complete Y-chromosome. Am. Nat., 72:42-52.
The innervation of setae in Drosophila. Genetics, 23: 172-73.
During which stage in the nuclear cycle do the genes produce their
effects in the cytoplasm? Am. Nat., 72:350-57.
Control of a species-difference by means of a difference in an in-
ductor. Nature, 142: 158.
Biology. In: An Orientation in Science, ed. C. W. Watkeys et al., pp.
256-332. New York: McGraw-Hill.
1939
Somatic crossing-over and somatic translocations. Am. Nat.,
73:95 -96.
With E. Hadorn. The relation between the color of testes and vase
efferentia in Drosophila. Genetics, 24: 162-79.
1940
Recent work on the relation between genes and developmental
processes. Growth, Suppl. 1:19-36.
The prospective significance of imaginal discs in Drosophila. J. Mor-
phol., 67: 107-22.
On dependent growth and form of the testes in various species of
Drosophila. Collect. Net., 15: 1-4.
Growth in vitro of the testis of Drosophila. Growth, 4:337-82.
1941
The growth of testes in Drosophila. I. The relation between vas
deferens and testis within various species. i. Exp. Zool., 87: 113-
58.
The growth of testes in Drosophila. II. The nature of interspecific
differences. i. Exp. Zool., 87:159-80.
OCR for page 464
464
BIOGRAPHICAL MEMOIRS
With A. Brasted. An analysis of the expression of the mutant "en-
~railed" in Drosophila melanogaster. (genetics. 26: 347-73.
~ .
1942
With L. Birmingham. Boundaries of differentiation of cephalic
imaginal discs in Drosophila. ]. Exp. Zool., 91:345-63.
1943
The Hardy-Weinberg law. Science, 97: 137-38.
Genic action as studied by means of the effects of different doses
and combinations of alleles. Genetics, 28:441-75.
Some new types of class experiments with Drosophila at the Univer-
sity of Rochester. Ward's Nat. Sci. Bull., 16:67.
Effect of environmental differences, and a "Lamarckian" experi-
ment. Ward's Nat. Sci. Bull., 16:93.
Cumulative and competitive action of alleles and their bearing on
the position eject. Genetics, 28:92-93.
With E. W. Schaeffer. On primary attributes of alleles in Drosophila
melanogastex Proc. Natl. Acad. Sci. USA, 29:351-61.
With E. SchaeEer. On wild-type iso-alleles in Drosophila melanogaster.
Proc. Natl. Acad. Sci. USA, 29:361-67.
1944
The journey not the goal. Sci. Mon., 58 :96 -100.
With E. W. Schaeffer and W. P. Spencer. The genetic basis of dif-
ferences between two species of Drosophila. Am. Nat., 78: 183-
88.
Peace time research in war time. Science, 99:278-80.
A study of race. l. Hered., 35:314-16.
With G. Heidenthal. Materials for the study of the position effect
of normal and mutant genes. Proc. Natl. Acad. Sci. USA,
30: 197-205.
1945
With D. R. Charles. The Rhesus gene and the effect of consan-
guinity. Science, 101 :305 - 7.
A letter a college president might write. Bull. Am. Assoc. Univ.
Prof., 31:1.
Review of Genetics, by E. Altenburg. Science, 102:514-15.
OCR for page 465
CURT STERN
465
1946
With E. W. Schaeffer and G. Heidenthal. A comparison between
the position effects of normal and mutant alleles. Proc. Natl.
Acad. Sci. USA, 32:26-33.
Review of Heredity and its Variability, by T. D. Lysenko, Am. Nat.,
80:241-43.
With G. LeClerc. Occurrence of mitotic crossing-over without
meiotic crossing-over. Science, 103:553 -54.
With R. MacKnight and M. Kodani. The phenotypes of hemizy-
gotes of position alleles and of heterozygotes between alleles in
normal and translocated position. Genetics, 31:598-619.
With M. Kodani. An invisible chromosome. Science, 104:620-21.
1947
The skin color of children from white by near-white marriages. I.
Hered., 38:233-34.
1948
With W. Spencer. Experiments to test the validity of the linear R-
dose/mutation frequency relation in Drosophila at low dosage.
Genetics, 33:43-74.
With E. Caspari. The influence of chronic irradiation with gamma
rays at low dosages on the mutation rate in Drosophila melano-
gaster. Genetics, 33:75-95.
Negative heterosis and decreased effectiveness of alleles in heter-
ozygotes. Genetics, 33:215 -19.
With Trudy Enders. The frequencies of twins, relative to age of
mothers, in American populations. Genetics, 33:263-72.
With E. Novitski. The viability of individuals heterozygous for re-
cessive lethals. Science, 108: 538-39.
The effects of changes in quantity, combination and position of
genes. Science, 108:615 -21.
1949
Gene and character. In: Genetics, Paleontology and Evolution, ed.
G. L. tepsen, pp. 13-22. Princeton: Princeton University Press.
With D. E. Uphoff. The genetic effects of low intensity irradiation.
Science, 109:609-10.
Selection and eugenics. Science, 110: 1-8.
OCR for page 466
466
BIOGRAPHICAL MEMOIRS
Principles of Human Genetics. San Francisco: W. H. Freeman and Co.
617 pp.
1950
Anomalies of genetic origin. Pediatrics, 5:324-28.
Reply to Bernhard Stern. Science, 111:698.
Genetic aspects of sterility. Fertil. Steril., 1 :407-14.
Boveri and the early days of genetics. Nature, 166:466.
With E. R. Sherwood. The migration of testis sheath cells in Dro-
sophila virilis. In: Moderne Biology. Festschrift fur Hans Nachtsheim,
ed. H. Gruneberg, pp. 236-40. Berlin: F. W. Peters.
1951
Concluding remarks of the chairman. Cold Spring Harbor Symp.
Quant. Biol., 15 :409-12.
With A. Hannah. The sex combs in gynanders of Drosophila melan-
ogaster. Port. Acta Biol., Ser. A:798-812.
With S. T. Fung. The seriation of fourth chromosome loci in Dro-
sophila melanogaster. Proc. Natl. Acad. Sci. USA, 37:403-4.
Probleme der menschliche Erbforschung. Mitt. Naturforsch. Ges.
Bern, 9.
Problems of radiobiology with emphasis on radiation genetics. In:
Proc. Annul Biol. Colloq., pp. 7-16. Corvallis: Oregon State Uni-
versity Press.
The genetic future of man. In: Proc. Annul Biol. Colloq., pp.43-51.
Corvallis: Oregon State University Press.
1952
Man's genetic future. Sci. Am., 186:68-74.
Genetics and the world today. In: The Scientists Look at Our World,
ed. }. M. Fogg, pp. 61-82. Philadelphia: University of Pennsyl-
vania Press. (Also in: Smithson. Year Annul Rep., Smithson.
Inst. for 1953, pp. 263-276.)
1953
With G. Carson, M. Kinst, E. Novitski, and D. Uphoff. The viability
of heterozygotes for lethals. Genetics, 37:413-49.
With G. Belar. Race crossing in paradise? }. Hered., 49: 154-55.
OCR for page 467
C U RT S T E RN
467
The geneticist's analysis of the material and the means of evolution.
Sci. Mon., 77: 190-97.
Model estimates of the frequency of white and near-white segre-
gants in the American Negro. Acta Genet. Stat. Med., 4:281-
98.
1954
Listings of unpublished articles. Science, 119:221.
George W. Beadle. Science, 119:229 -30.
Two or three bristles. Am. Sci., 42:213-47
Needed research. Eugen. Q., 1:161-65.
The biology of the Negro. Sci. Am., 191:81-85.
The facts of life. Popul. Stud., London, 8: 188-91.
Guest editorial: One scientist speaks up. Science, 120:5A.
Genes and developmental patterns. In: Caryologia, Proc. 9th Int.
Congr. Gen., Part I, pp. 355-69. Firenze: Industria Tipografic
. . .
. ~ 1orentlua.
.
1955
A professor's days. Calif. Mon., 65: 15 -17.
Gene action. In: Analysis of Development, ed. B. Willier, P. Weiss, and
V. Hamburger, pp. 151-69. Philadelphia: W. B. Saunders Co.
With M. Kodani. Studies on the position effect at the cubitus inter-
ruptus locus of Drosophila melanogastex Genetics, 40:333-73.
Qualitative aspects of the population problem. Science, 121:683-
86.
Grundlagen der menschlichen Erblehre. Gottingen: Muster-Schmidt
Verlag. 560 pp.
1956
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Representative terms from entire chapter:
human genetics
