Frank Ambrose Beach, April 13, 1911--June 15, 1988 | By Donald A. Dewsbury | Biographical Memoirs

Photo by Betty Jane Nevis,
Berkeley, California
|

Frank Ambrose Beach
April 13, 1911 June 15, 1988
By Donald A. Dewsbury
|
FRANK A. BEACH WAS arguably the premier psychobiologist of
his generation, influencing the development of psychobiology in
numerous, diverse ways. Believing that learned behavior was too complex
for detailed analysis, he shifted the focus of the field toward the
study of instinctive, or as he preferred, species-specific behavioral
patterns, such as mating and parental behavior.
A
major impact was Beach's movement of the field toward increased
physiological considerations, as in research on the neural and endocrine
determinants of behavior. Along with William C. Young, he established
the field of behavioral endocrinology. Physiological analysis can
quickly become reductionist; in Beach's hands, by contrast, it was
integrative. He sought to understand behavior not only with respect to
the two-way relationships with neural and endocrine processes but in
dynamic relation to the complex environment in which animals live.
Further, Beach believed that behavior should be understood in an
evolutionary framework. The function of behavior was to permit animals
to adapt to complex and ever-changing environments. He sought an
integrative psychobiology that would transcend these levels of analysis
and focus on behavior, but it would be rooted in the study of its
physiological correlates on the one hand and its adaptive function on
the other. He thus tried to unify the physiological and comparative
factions of psychobiology.
Beach strongly valued
experimentation and was a skilled experimenter--a seat-of-the-pants,
follow-your-nose kind of experimentation, rather than one based on
sophisticated mathematical analysis or elaborate equipment. He was a
down-to-earth Midwesterner. Beach's most lasting influence stemmed from
his ability to think about the field and to write integrative articles
that would synthesize developing trends and shape their evolution toward
an improved integrated psychobiology. His sense of timing was exquisite.
Frank Ambrose Beach was born in Emporia, Kansas, on
April 13, 1911. His mother was Bertha Robinson Beach; his father, Frank
A. Beach, was a professor and head of the Department of Music at Kansas
State Teachers College in Emporia. The music building at what is now
Emporia State University is named in his honor. Beach rarely used the
"Jr." associated with his name.
Beach attended the
teacher's college with the goal of becoming an English teacher. He
received his B.S. degree in education in 1932. His course was not always
smooth. Freshman year grades were so low that Beach's parents sent him
away to Antioch College for a year to improve his academic motivation.
The strategy worked, and he returned to Emporia for his junior year,
becoming a campus leader and taking his first course in experimental
psychology. Although its effect was not immediate, this course altered
his life's direction. His instructor, James B. Stroud, had earned his
doctorate with Harvey A. Carr at the University of Chicago, and Beach
later named Stroud as the teacher who had exerted the greatest influence
on his professional development.
Unable to find a
job teaching English when he graduated at the peak of the Depression,
Beach accepted a fellowship from Stroud on the condition that Beach
would pursue an M.S. in psychology. Although his work was in clinical
psychology, Beach chose as a thesis topic the determination of whether
rats had color vision. Because the department had no animal facilities,
Beach had to establish and run them himself. He did this successfully
and received his master's degree in 1933.
Still
unable to find a steady job, Beach took Stroud's advice to explore the
possibility of further study in anthropology at the University of
Chicago. He traveled to Chicago and found that he was unable to pursue
that path because the Department of Anthropology had no stipends for
beginning students. However, a courtesy call to Carr led to a fellowship
and graduate study in psychology during 1933-34. Between the fellowship
and odd jobs, including singing in a choir, Beach was able to study for
a year in Chicago. Carr, the noted leader of the Chicago functionalist
school of psychology, proved to be an ideal teacher. Beach was not
enamored of the mathematical approach of Louis Thurstone, one of
Chicago's "star" faculty members. He had a lifelong distrust of complex
mathematical operations, believing that just because it was possible to
perform statistical operations did not mean one should do so. It was a
third faculty member, Karl Lashley, a notoriously poor classroom
teacher, who exerted a lasting influence on Beach. Lashley allowed
students to work on their own, and Beach found the laboratory
environment and the problems of physiological analysis in which Lashley
was interested to be irresistible. Outside of psychology, courses with
Paul Weiss and C. Judson Herrick also made an impact.
As he experienced financial difficulties and a job
teaching high school English finally opened up, Beach took off a year to
teach junior and senior English at Yates Center, Kansas. It was an
active year, during which he directed plays, supervised the student
newspaper, and performed other routine duties of a high school English
teacher. He even conducted psychological research on the rate of
learning of a stylus maze by his students.
The
lure of psychology was great, however, and a university fellowship
enabled Beach to return to Chicago after one year of teaching. By now,
Lashley had moved to Harvard, but Carr was receptive to a dissertation
proposal for a study in Lashley's field--on the effects of lesions to
the cerebral cortex on maternal behavior in rats. With Lashley gone,
Beach again had to rely on his own skills and on fellow students for
help, but he was able to complete his Ph.D. candidacy examination and
his dissertation research in one year, only his second at Chicago. Among
his important associates at Chicago were David Krech and Leon
Pennington. Beach sought employment while he wrote the dissertation and
completed his language requirements. Lashley came to the rescue,
offering Beach an assistantship in neuropsychology at Harvard.
In March 1936 Beach married Anna Beth Odenweller, a
fellow Kansan, who had been studying at the Goodman School of Theater at
the Chicago Art Institute and whom Beach met in the choir. They would
have two children, Frank A. Beach III born in 1937 and Susan Elizabeth
Beach born in 1942.
At Harvard during 1936-37,
Beach studied the effects of similar brain lesions on another
instinctive behavior in rats, copulation. He would come to spend more
time during his career studying copulatory behavior than any other
problem. He later called this a halcyon year, with much time available
for research but little time for language study. While at Harvard, he
formed lifelong friendships with Donald Hebb, Edwin Ghiselli, George
Drew, and Andre Ray.
After a year at Harvard,
Lashley "kicked him out of the nest," recommending Beach for a position
as an assistant curator in the Department of Experimental Biology of the
American Museum of Natural History in New York. The museum position
allowed Beach an opportunity--both time and facilities--for full-time
research. The chance to interact closely with diverse biologists in
fields such as mammalogy, herpetology, and ornithology had a broadening
effect and helped shape the expansive biological perspective that so
characterized his later work. The only problem was in getting along with
department chair G. Kingsley Noble.
By 1940 Beach
had finally learned sufficient German and French to pass his language
exams at Chicago, and he returned to the university for his oral
examination. Carr came out of retirement to conduct the examination, and
Beach received his Ph.D. degree in 1940.
A
colleague at Harvard had suggested that the loss of copulatory behavior
that followed the cortical lesions Beach had given to rats might be due
to indirect effects of the lesions on the endocrine system. To learn
more about endocrinology, Beach audited a course with Robert Gaunt at
New York University. When he found that there was little on
endocrinological effects on behavior, he began library research for a
term paper that he eventually developed into his first book, Hormones
and Behavior, published in 1948.
When Noble
died in December 1940, museum director Roy Chapman Andrews was prepared
to shut down the Department of Experimental Biology. Beach, however,
lobbied various scientists around the country to intervene, with the
result that Beach became the new chairman with the rank of full curator.
He changed the name to the Department of Animal Behavior. The department
provided a home for numerous very active comparative psychologists for
many years. It was during this period, however, that Beach acquired his
lifelong aversion to academic administration.
In
1946 Beach left the museum for a position in the Department of
Psychology at Yale University. He later claimed that he had done a poor
job at that point in his career in teaching lecture classes, but he was
allowed to teach mainly smaller seminars and to supervise graduate
students. In 1952 Beach was named a Sterling professor of psychology at
Yale.
Beach valued science and believed himself
responsible for communicating his results to his colleagues, but during
most of his career he was not especially active in popularizing science.
An exception appeared during the early Yale years with a series of
articles that dealt with learned behavioral patterns, which he published
in Natural History magazine. His rhetorical strategies in these
articles focused on the importance of hard-nosed science in
understanding even the most complex behavior.
During his tenure at Yale, Beach became progressively
more active in the affairs of various scientific organizations. He
regarded this as a responsibility of scientists, but as a finite one. He
was elected president of the Eastern Psychological Association and was
selected as a charter member of the psychobiology panel of the National
Science Foundation. In 1955 he became a member of the National Research
Council's Committee for the Study of Problems of Sex, and two years
later was made chairman. Years later, he would pride himself in having
closed down the committee. Federal funding had come to dwarf that of the
committee, and Beach believed that committees often search for functions
long after their useful purpose has been achieved. He was determined not
to let this happen. He also served on the Publications Board and Policy
and Planning Board of the American Psychological Association, the
Advisory Board of the Marine Studios and Marine Research Laboratory in
St. Augustine, Florida, and on the Board of Scientific Directions of the
Roscoe B. Jackson Memorial Laboratory in Bar Harbor, Maine.
Beach also was invited to several prestigious
lectureships, including the William James Lectures at Harvard, the Smith
College Lectures, and the Jake Gimbel Lectures on the Psychology of Sex
at the University of California, Berkeley, and Stanford University.
While at Yale, Beach became more interested in human
sexual behavior and teamed up with anthropologist Clellan S. Ford to
write a book, Patterns of Sexual Behavior, which surveyed that
field in broad perspective.
Prior to 1948 Beach
worked with rats, hamsters, cats, and pigeons, but he then instituted a
program of work with dogs that would last much of the rest of his
career. Beach valued his Yale years for the opportunity to work with
bright students and colleagues under ideal conditions and to experience
life in a first-rate university.
Beach spent the
1957-58 year as a fellow at the Center for Advanced Study in the
Behavioral Sciences in Stanford, California. His primary endeavor while
there was to write a textbook on comparative psychology, but after
writing numerous chapters he finally gave up in frustration.
During his stay in Stanford, he was approached
regarding the possibility of a permanent move to the University of
California, Berkeley. He accepted the position partially because he had
come to learn that with each such move he had become acquainted with a
new set of colleagues from whom he could learn and enrich his own
research program. He also was attracted to the warmer climate and the
easy-going atmosphere. Beach accepted the position on the conditions
that he would determine his own teaching assignments, have ample
research space, be given a full-time secretary, and never be asked to
serve as department chairman.
Beach continued to
flourish in the Berkeley climate. He liked the California graduate
students and the fact that the department was not dominated by learning
theory as had been Yale in the heyday of Clark Hull. A major
accomplishment in Berkeley was Beach's founding of the Field Station for
Behavioral Research on a beautiful site overlooking the campus and San
Francisco Bay. There Beach could continue his program of research on
dogs in a more open environment. The site later became the location of a
major program of research on hyenas in which Beach participated on a
part-time basis after his retirement.
In 1961 and
1962 Beach hosted conferences under the sponsorships of the Committee
for Research in Problems of Sex, the National Science Foundation, and
the National Institute of Mental Health. These conferences brought
together students of sexual behavior with very diverse perspectives and
resulted in the edited volume Sex and Behavior in 1965.
On the occasion of his sixty-fifth birthday, his
former students and postdoctoral fellows held a working conference in
Berkeley in his honor. Beach regarded it as a highlight of his academic
life. This conference resulted in another book, Sex and Behavior:
Status and Prospectus.2
Beach was
a splendid mentor for graduate students and postdoctoral fellows.
Indeed, he provided a role model for many. He knew exactly when to turn
a sympathetic ear and when to deliver a swift kick in the pants. His
work with students was so strong that he received the American
Psychological Foundation's award for distinguished teaching in
biopsychology in 1985. He was cordial, yet maintained distance; graduate
students did not call him Frank. Beach thought the roles of father
figure and buddy incompatible.
Through much of his
career, he limited his teaching to graduate students and small groups.
In a remarkable reversal of his earlier career pattern, Beach became
motivated for undergraduate teaching late in his career. He taught large
undergraduate classes in comparative psychology and introduced an
experimental class in human sexuality. The latter effort led him to edit
another book, Human Sexuality in Four Perspectives (1976). He was
not satisfied with his first efforts at these courses, but he felt he
improved with experience.
Beach formally retired
in 1978, but he remained active in research. His first wife having died
in 1971, Beach married Noel Gaustad, who was especially important to him
as his health declined. Beach died in Berkeley in 1988 at the age of
seventy-seven. In the hospital the week before his death, Beach was
still reading literature and working with a co-author on one more
article.
Among his honors were honorary doctorates
from McGill University, Williams College, and Emporia State University;
the Warren Medal of the Society of Experimental Psychologists; and the
Distinguished Scientific Contribution Award of the American
Psychological Association. He was elected to the National Academy of
Sciences (at age thirty-eight), the American Philosophical Society, and
the American Academy of Arts and Sciences. The Frank A. Beach Award and
Lectureship were established in 1990 as a means of encouraging and
rewarding young researchers in behavioral endocrinology. The lectures
are published annually in Hormones and Behavior.
Frank Beach was an extrovert--at ease with people and
able to get along well with almost everyone. He could live hard at
times. He knew how to party. He was as comfortable at the poker table as
he was in the laboratory. At the same time, he had an exquisite
sensitivity for the English language and its use, along with a deep
respect for learning of all sorts and for the culture in which he lived.
He was possessed of great wit, eager to deflate pomposity, and was not
afraid to ask questions others would regard as naïve. When he left
the room, however, he understood fully what had transpired. Beach knew
well how to use his rich sense of humor in the interest of making and
dramatizing salient points of a serious academic nature.
Beach was a conservative in many senses of the word.
His academic values were old-fashioned, as were his political views. He
was proud that he did not let me spend all of the funds in my National
Science Foundation stipend; we returned some money to the federal
government. This is unusual! He did not support the activism rampant on
the Berkeley campus during the 1960s.
Beach's primary
contributions lay more in programmatic accomplishments and in directing
the field than in any single discovery. Nevertheless, his achievements
in some areas can be summarized in approximate topical and chronological
order.
In his early work, Beach explored the effects of
various interventions on instinctive behavior, primarily in rats. He
started out with brain lesions and such behavioral patterns as
copulation, parental behavior, and activity patterns. Themes that would
be prevalent in his work emerged early. The first study of the effects
of hormones on mating behavior and the first developmental study came in
1941; the first study of stimulus control of mating was in 1942.
Beach's first synthetic review dealt with the central
nervous mechanisms of reproductive behavior in vertebrates and came in
1942; others would follow. He concluded that both hormones and sensory
input were needed to act on neural mechanisms for the display of
copulatory behavior. Among his views was the belief that hormones have a
more important effect in the lower vertebrates; the sexual activity of
higher vertebrates is more dependent on cortical mechanisms. He also
believed that the activity of males is less hormone-bound and more
dependent on cortex than is that of females. Beach considered the
stimulus arousal of behavior to be multi-sensory, including olfactory,
tactile, auditory, and visual. None is critical; rather, they sum in the
brain to activate a sexual arousal mechanism essential for the
initiation of behavior. The arousal mechanism, he thought, is
independent of a hypothetical sub-cortical copulatory mechanism that is
responsible for the execution of the behavior once an arousal threshold
is reached--as a result of hormonal and sensory activity--and the
behavior occurs. Because hormones and sensory input from different
modalities sum, Beach proposed that activity in one component could
compensate for the inactivity elsewhere in the initiation of behavior.
Beach viewed the development of behavior as a
complex interaction of genes and environment, but he thought that early
play and other early experience were more important in primates than in
the lower vertebrates. He argued that once there has been a proper study
of the impact of genes and the environment the very necessity of a
category of "instinctive behavior" would become unnecessary. These
themes of the complex development and control of behavior, which emerged
early in his career, were explored throughout his tenure as a leading
psychobiologist.
Although much of Beach's work was with laboratory rats
and dogs, he published a study of the pouchless marsupial Marmosa
cinera in 1939 and added hamsters, pigeons, alligators, pigeons, and
other species over the years. It is not clear who first labeled Beach
"the conscience of comparative psychology," but he was noted for urging
a more comparative focus than had most psychologists studying animals,
especially in his 1950 article "The Snark Was a Boojum." Beach called on
psychologists to expand the range of species they study and the range of
behavioral patterns and problem areas they investigate. Beach wrote
various articles advocating the study of instinctive behavior, or what
was more often termed species-specific behavior.
It was shortly after World War II that the European
ethologists, especially later Nobel Prize winners Konrad Lorenz and
Nikolaas Tinbergen, began to become truly visible in North America.
Beach was a leader in calling attention to this approach and to
encouraging productive interaction between North American comparative
psychologists and European ethologists. He served on the first editorial
board to the ethological journal Behaviour.
Beach believed in the careful description of behavior
in objective terms that could be understood and used by different
investigators. He hesitated to label behavior as sexual or aggressive
because so many motor patterns could appear in different functional
contexts. The description of what occurred was to be objective and kept
separate from the functional interpretation given the behavior by the
observer.
During the 1950s and 1960s, Beach and
his students conducted a long series of studies in rats of the
determinants of the complex pattern of mounts, intromissions, and
ejaculations, which are displayed during copulation. Many of these were
related to a conceptual model carrying forward the theme of independent
sexual arousal and copulatory mechanisms. By manipulating various
temporal parameters and stimulus situations, they probed the manner in
which these mechanisms worked first to bring the animal to the threshold
of sexual activity and then to bring the hypothetical copulatory
mechanism to the point of ejaculation. After that, attention was
addressed to the problems of the recovery of sexual motivation, both
during and between multiple-ejaculation sessions.
As noted, Beach tended early in his career to believe
male behavior more complex than female behavior. In a changing cultural
climate, he later realized the previously unrecognized complexity of
female behavior. In 1976 he wrote the definitive article delineating the
differences among receptivity (to the male), attractivity (to the male),
and proceptivity (the active solicitation of the male) in female sexual
behavior. As always, his careful descriptions and his timing were well
tuned.
Throughout this whole period, Beach's interest in
hormone-behavior interactions was growing. His early book Hormones
and Behavior, along with some writings of W. C. Young, had first
crystallized the area. Beach continued to do research and to write
synthetic articles outlining and promulgating what came to be known as
behavioral endocrinology. But Beach understood that an emerging
discipline needs the accouterments. In 1975 he provided an integrative
article in American Scientist. Six years later he wrote a
scholarly history of the field. In 1979 Beach, along with Julian
Davidson and Richard Whalen, founded the field's first journal,
Hormones and Behavior.
One of the important
developments in psychobiology during the 1960s was the realization of
the important effects of early hormone action on later behavior. Beach
with others conducted much research on such early hormone effects. The
dominant view was that, whereas the effects of hormones in adults were
activational, the effects early in life were in the organization of
neural tissue. Beach was skeptical of these conclusions and wrote
several articles questioning the evidence. Although the conclusions
appear to have been largely correct, Beach's skepticism helped focus
research and thinking on the clarification of the concepts and on the
development of sufficient evidence to warrant general conclusions.
It is important to remember that for Beach
hormone-behavior interactions were a two-way street. Hormones not only
affect behavior, they also are affected by behavior. Beach devoted much
effort to writing a textbook on behavioral endocrinology, but like the
textbook on comparative psychology, it lay incomplete at his death.
In his series of studies of dogs, conducted at the
Berkeley field station, Beach could fully explore the hormonal and
environmental interactions in the regulation of sexual behavior in dogs.
Here he could breed his own animals and study them year round out of
doors. In this context he found that individuals have distinct, if
somewhat idiosyncratic, preferences for mating partners. When
copulating, dogs achieve a lock, or mechanical tie, that makes
separation of the male from the female mechanically difficult. Always
gifted with the ability to turn a phrase and create a colorful title, he
summarized the work in his 1969 article "Locks and Beagles." The work
continued for many years, and Beach later embraced with characteristic
enthusiasm the research by his successors at the field station on
hormonal and developmental factors in hyena behavior.
The topic of human sexuality
emerged early in Beach's career. It is worth noting that investigation
of this topic required more courage in the climate of the 1940s than it
did in the 1990s. Beach returned to questions of human sexuality
repeatedly throughout his career. Many psychologists studying nonhuman
animals study them in order to understand humans and often to enable
social interventions. For Beach, by contrast, an interest in humans was
more a matter of intellectual challenge. Humans are complex, and
unraveling their behavior presents a special challenge for which all
methods and disciplines are important. His 1977 book Human Sexuality
in Four Perspectives dealt with developmental, sociological,
physiological, and evolutionary perspectives. Beach believed that the
way to understand humans lay not in generalizing from animals to humans
but rather by studying humans the same way as other species and looking
for similarities and differences.
He was
especially cautious in generalizing across species. There are many
descriptions of homosexual and masturbatory activity in nonhuman
animals, for example. Beach stressed, however, that the male-male or
female-female mounting that can be seen in laboratory rats or monkeys
were quite different from the culturally complex processes of gender
preferences in humans. Beach cautioned that "surface similitude by
itself does not justify theoretical inferences."3 He stressed
the importance of understanding underlying mechanisms and only
generalizing across patterns with true functional and causal similarity.
In conclusion, Frank Beach was a firm believer in
basic research for the sake of knowledge, with practical application a
secondary concern. Science is not technology. He recognized, however,
that if one accepts the support of society in these endeavors, one has
the responsibility to conduct the work with integrity and to record the
results in the public record. Students were taught that experiments are
not complete until they are reported for the scientific public. Science
is serious--but it should still be fun.
Beach
believed in careful and precise behavioral measurement, but the mere
gathering of facts through what he called "ant-like industry" was of
little value unless integrated into a theoretical concept. His work
shows the effects of attention to the forest and the trees. Beach
believed that much of the research in journals was not worth doing
because it lacked a clear focus. He believed that if research is not
worth doing, it is not worth doing well and that the last thing most
scientists seem to understand are the fundamental questions with which
they are dealing.
Beach was a true believer in the
progress of science. He conducted many experiments and developed various
theoretical models. He genuinely believed, as should all scientists,
that his work was the best effort possible at the time but that it was
likely to be surpassed by later research. It was important to him that
his successors understand that what he did was reasonable in the context
of his time and of what he could have been expected to know when he
conducted the research. It did not matter that his work would be
superseded; indeed, he encouraged it. As long as his work moved the
field in the right direction, the work itself could fade. He wanted to
contribute to scientific progress and to help shape the field in a way
he felt would maximize long-term scientific understanding.
He succeeded--and he had fun doing so.
I THANK STEPHEN E. Glickman, Benjamin D. Sachs, and Irving Zucker for comments on an earlier draft of
this biographical memoir.
1 There are numerous sources on Beach. He wrote
three autobiographical chapters: Frank A. Beach. In A History of
Psychology in Autobiography, vol. 7, ed. G. Lindzey, pp. 31-58.
Englewood Cliffs, N.J.: Prentice-Hall, 1974; Confessions of an imposter.
In Pioneers in Neuroendocrinology, vol. 2, eds. J. Meites, B. T.
Donovan, and S. M. McCann, pp. 19-35. New York: Plenum, 1978; Conceptual
issues in behavioral endocrinology. In Autobiographies in
Experimental Psychology, ed. R. Gandelman, pp. 1-17. Hillsdale,
N.J.: Erlbaum, 1985.
A number of
obituaries are useful: S. E. Glickman and I. Zucker. Frank A. Beach
(1911-1988). Am. Psychol. 44(1989):1234-35; D. A. Dewsbury. Frank
Ambrose Beach: 1911-1988. Am. J. Psychol. 102(1989):414-20; B. D.
Sachs. In Memoriam: Frank Ambrose Beach. Psychobiology
16(1988):312-14.
Also useful is a
set of recollections by former students and colleagues: In Memoriam:
Frank A. Beach (April 13, 1911-June 15, 1988). Horm. Behav.
22(1988):419-43.
There is also an especially insightful
interview: J. D. Fleming and D. Maxey. The drive of the pure researcher:
Pursuit of intellectual orgasm. Psychol. Today 8(1975):68-77.
A list of Beach's publications and pre- and
postdoctoral students is available in Sex and Behavior: Status and
Prospectus, eds. T. E. McGill, D. A. Dewsbury, and B. D. Sachs. New
York: Plenum, 1978.
A selection of Beach's
academic papers can be found in the Archives of the History of American
Psychology at the University of Akron (Ohio).
2 T. E. McGill, D. A. Dewsbury, and B. D.
Sachs, eds. Sex and Behavior: Status and Prospectus. New York:
Plenum, 1978.
3 F. A. Beach. Cross-species
comparisons and the human heritage. Arch. Sex. Behav. (1976):469.
- 1937
- The neural basis of
innate behavior. I. Effects of cortical lesions upon the maternal
behavior pattern in the rat. J. Comp. Psychol. 24:393-436.
- 1942
- Central nervous
mechanisms involved in the reproductive behavior of vertebrates.
Psychol. Bull. 39:200-226.
- Analysis of the
stimuli adequate to elicit mating behavior in the sexually inexperienced
male rat. J. Comp. Physiol. Psychol. 33:163-207.
- 1944
- Relative effects of
androgen upon the mating behavior of male rats subjected to forebrain
injury or castration. J. Exp. Zool. 97:249-95.
- 1945
- Current conceptions of play in animals.
Am. Nat. 79:523-41.
- 1947
- Evolutionary changes in the physiological control of mating
behavior in mammals. Psychol. Rev. 54:297-315.
- 1948
- Hormones and Behavior
. New York:
Hoeber.
- 1950
- The snark
was a boojum. Am. Psychol. 5:115-24.
- 1951
- With C. S. Ford. Patterns of Sexual
Behavior. New York: Harper.
- 1954
- With J. Jaynes. Effects of early experience upon the
behavior of animals. Psychol. Bull. 51:239-63.
- 1955
- The descent of instinct. Psychol.
Rev. 62:401-10.
- 1956
- With L. Jordan. Sexual exhaustion and recovery in the male
rat. Q. J. Exp. Psychol. 8:121-33.
- 1965
- Sex and Behavior
. New York:
Wiley.
- 1966
- The
perpetuation and evolution of biological science. Am. Psychol.
21:943-49.
- 1967
- Cerebral
and hormonal control of reflexive mechanisms involved in copulatory
behavior. Physiol. Rev. 47:289-316.
- With B. J.
LeBoeuf. Coital behavior in dogs. I. Preferential mating in the bitch.
Anim. Behav. 15:546-58.
- 1969
- Locks and beagles. Am. Psychol. 24:971-89.
- 1970
- Coital behavior in
dogs. VI. Long-term effects of castration on mating in the male. J.
Comp. Physiol. Psychol. 70:1-32.
- 1971
- Hormonal factors controlling the differentiation,
development and display of copulatory behavior in the ramstergig and
related species. In Biopsychology of Development, eds. L. Aronson
and E. Tobach, pp. 249-96. New York: Academic Press.
- 1975
- Hormonal modification of sexually
dimorphic behavior. Psychoneuroendocrinology 1:3-23.
- Behavioral endocrinology: An emerging discipline. Am.
Sci. 63:178-87.
- 1976
- Sexual attractivity, proceptivity, and receptivity in female
mammals. Horm. Behav. 7:105-38.
- Cross-species
comparisons and the human heritage. Arch. Sexual. Behav.
5:469-85.
- 1977
- Human
Sexuality in Four Perspectives.
Baltimore: Johns Hopkins University
Press.
- 1981
- Historical
origins of modern research on hormones and behavior. Horm. Behav.
15:325-76.
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