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Biographical Memoirs: V.58 (1989)

Chapter: Harry Frederick Harlow

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Suggested Citation:"Harry Frederick Harlow." National Academy of Sciences. 1989. Biographical Memoirs: V.58. Washington, DC: The National Academies Press. doi: 10.17226/1645.
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HARRY FREDERICK HARLOW October 31, 1905-December 6, 1981 BY JOSEPH B. SIDOWSKI AND DONALD B. LINDSLEY HARRY HARLOW was born Harry F. Israel in Fairfield, Iowa, the third of four sons born to Lon H. anct Noble (Rock) Israel. For reasons unknown, he changed his name legally to HarIow while in college. After forty-four years of association with the University of Wisconsin (1930-1974), he became professor emeritus and retired to Tucson, Arizona. where he served as honorary research professor of the Uni- versity of Arizona. In his later years, he suffered from Par- kinsonism. He died of a brain tumor in 1981. ACADEMIC YEARS Harry HarIow, as he himself described it, was a shy, retir- ing, and callow youth when he began his college studies at Reect College in Portland, Oregon, in 1923. After one year he decidecl to follow his brother to Stanford University, where he wouIc! receive his B.A. degree in 1927, majoring in psychology. His original intent had been to major in English, but an unfavorable grade in that subject anct an exciting in- troductory course in psychology changed his mind. His po- etic nature and an ability to use the English language in a humorous manner remained, later contributing greatly to his success both as a teacher and a professional lecturer. While still an undergraduate, HarIow supported himself 219

220 BIOGRAPHICAL MEMOIRS working as an assistant to the experimental psychologist Wal- ter R. Miles, who was elected to the National Academy of Sciences in 1933. As a graduate student at Stanford, HarIow came under the tutelage of Calvin P. Stone, who was elected to the Academy in 1943. As a graduate student, Harry held a teaching assistantship uncler Paul R. Darnworth in social psychology and research assistantships under Stone in be- havioral studies on rats. His cloctoral dissertation dealt with the social facilitation of eating behavior in rats, combining elements of his ongoing experiments as an assistant. Much later, Harlow sail! that he learner! scientific methoclology and techniques from Stone, but he always consiclerec] Miles his moral and ethical mentor. He admirect Lewis W. Terman, then head of the depart- ment of psychology, and learned about theory in psychology from him. Terman had been elected to the Academy in 1928. Towarc! the ens! of Harry's second graduate year, Terman wrote to Harlow's mother of his great progress in psychology ant! his preparation for academic teaching and research. However, later when Harry was seeking an academic posi- tion, Stone, Terman, and Miles all advised him to consoler a junior college position because of a speech defect, which they thought interfered with his ability to articulate clearly and sometimes brought forth smiles when he said "wat" for rat! Despite this advice, he accepted a position as an assistant professor of psychology at the University of Wisconsin in 1930, where he regularly taught the large introductory class in psychology. With cleterminec! application, his diction and enunciation steadily improved, ant] he became one of the most effective and popular lecturers on campus. It was prob- ably with these student audiences that he developer! his un- hurriecI, clipped manner of speech that along with his cre- ative intellect and great wit ultimately made him one of the

HARRY FREDERICK HARLOW 221 most entertaining, elective, and sought after speakers in all of psychology. Hired as a comparative animal psychologist, Harlow ar- rived at the University of Wisconsin in 1930 to learn that there was no animal laboratory. However, he soon found a cramped cubicle in which to house his rats, which happened to be just below the office of the Dean of Men who didn't appreciate the odors wafting upward. As a result, Harry was displaced from that location and given a small space in the University Medical School. There he began studies of the social facilitation of feeding responses in monkeys, an extension of his doctoral research with rats. But that space, too, proved vulnerable and temporary, and his first steps into a major career dedicated to the study of nonhuman primate behavior began at a bridge party, when the wife of the chairman of the psychology department sug- gested that he study primates at the local Vilas Park Zoo. The Zoo afforded an opportunity to work with a variety of pri- mates, including an orangutan, baboons, and monkeys, ex- periences that were to prove invaluable and would lead to an unexpected turn in his career. PRIMATE LABORATORIES AND RESEARCH Harlow's first primate research facility consisted mainly of a few tables, a test tray, and test objects at the Vilas Park Zoo. In 1932 the University of Wisconsin made available to him a very small, two-story structure that had previously been the Forest Products Laboratory. It was badly in need of renova- tion. With his own meager funds and the aid of Walter Grether, Paul Settlage and other graduate students this was accomplished. The result was a usable research facility and the first real primate laboratory in Wisconsin's Department of Psychology.

222 BIOGRAPHICAL MEMOIRS Acquiring a small colony of monkeys, HarIow and his graduate students enthusiastically began developing new ant! unique ways to study primate behavior, both qualitatively and quantitatively. Using the oddity principle and matching- from-sample procedures they were able to study perceptual discrimination involving figures and patterns on visual clis- plays or objects that cliffered in color, size, shape, or texture. By introducing time clelays between stimulus presentation and opportunity to respond (method of cielayect response), they could study both learning and memory (lecay. Combin- ing different tasks in so-callect test batteries they could ex- plore and identify the nature ant! extent of "animal intelli- gence" in various species as wed as in humans. In orcler to conduct these experiments in a uniform way they designed and built a stanciard piece of equipment, known as the Wis- consin General Test Apparatus (WGTA). This device was acloptec! and used by many investigators over the years, even . . unto recent times. One of the most significant discoveries HarIow and his associates macle in their first primate laboratory dealt with the formation of learning sets, that is, the process by which animals "learn to learn." Their procedure was to present pairs of objects or patterns that cliffered in features such as size, color, and shape over a series of trials. The objects changer! every few trials, and the animal graclually learner! to abstract particular features that clifferentiatecl the correct response object from others. In this way, discrimination cues became generalizect and a learning set was establishect. Hariow and his students, as well as others, exploited this technique in the study of brain lesions ant! other experimental vari- ables. The origin and concept of the learning set idea was not sullen. From 1939 to 1940, cluring a sabbatical year, Hariow held a Carnegie fellowship at Columbia University with

HARRY FREDERICK HARLOW 223 famed anthropologist Franz Boas. While at Columbia he at- tendect a seminar by the German neurologist Kurt Goiclstein and became familiar with his theories concerning abstract and concrete intelligence and learning, which relied heavily upon performance on block-sorting tests such as the Weig! or Vigotsky tests. In these tests small wooclen blocks vary- ing in size, color, ant] shape must be sorted anct grouped ac- corcling to one or more of such categorical features and the principle of a category iclentifie(l. Accorcling to GoIclstein, only humans are capable of abstract thought. Hariow tenta- tively ctisagreed. Upon returning to Wisconsin he pursued research that eventually demonstrated—contrary to GoIcI- stein's view that monkeys could also solve WeigI ancT Vigot- sky type problems, suggesting certain levels of abstract thought and reasoning. These results, together with those from his earlier studies of odclity and matching-to-sample discriminations caused HarIow to focus on the question of methoclology. Limited by cost, upkeep, anct availability of monkeys, Har- low was forced to ignore the usual experimental procedures of the time; that is, use of naive and different animals for each condition or problem, as was the practice with cheap ant! plentiful rodents. He used the same monkeys for the study of a variety of problems. If separate groups of monkeys hac! been used to learn single, simple discriminations, he might not have discovered the concept of learning set. He fur- ther realized that subjecting monkeys to series of similar but related problems paralleled the situations in which children learn. At a time when Thornclikian trial-and-error learning was at variance with the "Ah ha!" solutions attributed by Gestalt- ists to sudden insight, HarIow presented results on multiple- problem solution to explain how animals learn-to-learn a problem-by-problem exposition of the briciges between trial-

224 BIOGRAPHICAL MEMOIRS and-error learning and insight. These results posed addi- tional difficulties for the conditioning theories of Clark Hull and Kenneth Spence, influential learning theorists at that time. Sometimes bitter arguments ensued, but HarIow's re- sults and interpretations could not be denied. His Earning set results were enthusiastically received when presented in his Presidential Address before the Midwestern Psychological Association in 1948. The subsequent wide acceptance of these results undoubtedly enhanced his reputation as a cre- ative scientist and with it his confidence in his general ap- proach to scientific investigation. Ahead of their time, these studies oriented the methods and thinking of modern cog- nitive psychologists toward natural as opposed to contrived . . . information processing. Another notable accomplishment involved investigations of newly conceived and identified curiosity and manipulation Graves, in cooperation with Robert A. Butler, Donald R. Meyer, and Harry's wife, Margaret Kuenne HarIow, a child psychologist. At a time when drives were considered to be wholly or partly physiological, HarIow and his associates es- tablished the fact that the curiosity and manipulation drives were intrinsic parts of the rhesus monkey's motivational structure. Food, water, and sex were not found sufficient or necessary to initiate behaviors resulting from curiosity and manipulation drives. Monkeys were just naturally curious and would work hard, if necessary, to satisfy their curiosity. They would, for instance, manipulate mechanical puzzles in- cessantly without the rewards deemed necessary by behav- ioral theorists of the day. Furthermore, HarIow's monkeys learned complicated tasks without being deprives! of basic necessities such as food and water. Along with the foregoing studies of a strictly behavioral and psychological nature, which had such an important bear-

HARRY FREDERICK HARLOW 225 ing on theoretical issues with regard to motivation, drives, and learning, HarIow and his colleagues engaged in a pro- gram of neurophysiolog~cal anct behavioral studies in an ef- fort to determine the role of the central nervous system, and especially the cerebral cortex, in conditioning, visual discrimi- nation, learning, and memory. The need for more refined behavioral tests in connection with these brain lesion-behav- ioral studies lect to the Wisconsin General Test Apparatus (WGTA) and to a great variety of test batteries and proce- clures. In pioneering investigations with Stagner (1933) ant! Sett- lage (1939), as well as in one of his own studies (1940), Har- low sought to determine whether a classical Pavlovian con- clitioned response could be establishect in the cat if, during the normal training procedure, the paw-lifting response to the unconditioned! stimulus (shock) was eliminates! or modi- fied by curare paralysis. Testing for the response to the con- ditioned stimulus (tone or light flash) was done after the cu- rare paralysis had worn off. Apparently the assumption was that everything, including the motor discharge blocked by the curare at the neuromuscularjunction, would be the same, except for the absence of the paw-lifting response to shock. After an appropriate period of training, ant! when the muscle was free of paralysis, they found that no conditioned response could be elicited. Although this appeared to be a landmark cliscovery, there were obvious flaws in the hypoth- esis, for proprioceptive feedback was also eliminated by the lack of movement caused by the curare. Furthermore, the result was subsequently shown by others to be inconclusive when it was found that curare tract a depressing effect on the central nervous system, as well as a paralyzing action at the neuromuscular junction. Hariow then abandoned this type of research, but many years later he considered that decision

226 BIOGRAPHICAL MEMOIRS to be a mistake. In hindsight he felt that he hac! been on the verge of an important discovery that was not unearthed until years later by other investigators. From about 1940 on, HarIow, his students, and associates made repeated attempts to determine the ejects of brain lesions and ablations on the ability of monkeys to make sen- sory discriminations and perform various tasks on tests de- veloped for use in the WGTA. Many of these studies resultect in important contributions, but very little of major signifi- cance evolvecI, compared with the earlier ancI later areas of investigation with which HarIow was associated. One set of studies conductecl by HarIow and Dagnon ~ ~ 943), Spaet (1943), ant! Campbell (1945) may be mentioned for its pio- neering importance in the clarification of an issue with re- garc! to the function of the prefrontal cortex in monkeys. Carlyle Jacobsen, working in the laboratory of physiologist John F. Fulton at Yale in the 1930s, had stuclied the clelayed response performance of monkeys following prefrontal cor- tex ablations and found that the monkeys couIc! not seem to determine which foodwelIs had been baiter! prior to the time- clelay introduced in the delayed response test. Jacobsen re- ported that the prefrontal cortex lesions had caused a deficit in immediate and short-term memory. HarIow and his associates had founct variability in the per- formance of their lesioned monkeys, but there was clear evi- clence that some monkeys could manage the time-delays and other discriminations that would not have been possible with severe memory deficits. Instead, they attributed the variabil- ity and the sometimes poor performance to an inability to attend to the task ant! avoic! distractions. These results, how- ever, were anteciated by the publications of Malmo ~ ~ 942) and Finan (1942), who used equipment and procedures like Ja- cobsen's except that the experimental chamber was in com- plete darkness to insure that the monkeys' attention was

HARRY FREDERICK HARLOW 227 focused only upon the stimulus panels, thus avoiding dis- tractions. These findings were later confirmed by French and Harlow ~ ~ 9621. Thus, ~acobsen's putative memory loss results could now be interpreted as due to distraction and inatten- tion rather than an inability to form, store, and retrieve mem- ories after prefrontal lobe ablations. Such results whether interpreted as attention or memory deficits, had important implications for the performance of human frontal loboto- mies, initiated in 1936 by the Portuguese neurosurgeon An- tonio Egas Moniz and continued through the 1940s and into the 1950s before being generally abandoned, despite some reported improvement in depressive and other psychopath- ological conditions. Earlier recognition of the disadvantages of such operations as revealed by animal studies like Harlow's might have forestalled the vast number of lobotomies per- formed. In 1932 Harlow moved into a two-story building that was to be his laboratory for the next twenty years. This building had less than the desirable amount of space in which to fit a small colony of monkeys, graduate students, postdoctoral vis- itors, laboratory equipment, and facilities for experimenta- tion. It also lacked the necessary office and desk space for the analysis and storage of research data. Furthermore, it was in the early stages of the Depression and financial support was in short supply everywhere. There were, of course, no fed- eral granting agencies at that time to support research and training fellowships for graduate students, as there would be later in the 1950s and beyond. These, however, were prob- lems faced by most college and university professors lucky enough to have a job. It is said, "Where there is a will, there is a way!" Harry had a will, and he found a way. He was highly motivated and had recently found a goal that would become a lifetime en- deavor: focus on the rhesus monkey (Macaca mulata) as an

228 BIOGRAPHICAL MEMOIRS experimental mode! for the study of the neural and behav- ioral aspects underlying human psychology. He soon found that not the least of his problems was the upkeep and survival of his monkeys. Over the next twenty years he developer! the experience ant] knowlecige necessary to sustain primates over- long periods of time within animal enclosures, though they enjoyed only a few summer months of the warm weather typical of their natural habitat. It was also in this laboratory that Hariow supervises! his first Ph.D. student, Abraham MasIow, who later developed the self-actualization theory of motivation and was creditect with being one of the founders of the humanistic psychology movement. In 1953, the primate laboratory operations were moved from their initial location to a renovated cheese factory sev- eral city blocks from the campus. The motivational, learning, and neurophysiological-behavioral research was continued ant! expanclect, resulting in a need for more monkeys. For- tunately, the space was now acloquate. Because of import problems, disease, and the cost of the monkeys, the decision was made to start a breeding colony of rhesus monkeys. There was virtually no information available on the care and rearing of laboratory-born monkeys. Methods were clevised through trial, error, and observation to enhance the proba- bility that the newborns wouIc! survive. Initially, forty infant rhesus monkeys were separated from their mothers and raiser! in separate cages. The result was disease-free animals that manifested bizarre ant! psychopath- ological behaviors. These abnormal behavior syndromes were attributed to the effects of early isolation and lecl to some of Harry HarIow's most fascinating and best-known re- search. The breeding, rearing, and nursery procedures proved successful overall, and a subsequent published report with A. I. Blomquist served as a guide for breeders in other animal installations, including zoos. HarIow's infant pri-

HARRY FREDERICK HARLOW 229 mate care methods were eventually adopted in many places arounct the world. The availability of laboratory-born infants led to the study of the ontogeny of learning and the clevelopment of age- sensitive learning tests, some of which showed that learning- set formations die] not develop until approximately twelve to eighteen months of age. They provided interesting data for comparison with the age-level stages of intelligence and men- tal growth established by lean Piaget, the famous Swiss chilct psychologist. But the infant rhesus macaques were to pro- cluce even better known (lata, specifically in the areas of af- fection and love. HarIow's research on affectional systems evolved primar- ily from a bizarre result of infant isolation: the inability to reproduce upon reaching maturity. This, of course, influ- encecl the supply of research animals. To remedy this situa- tion, HarIow thought of a way to provide "mothering" of a sort to the isolates] infants by developing surrogate mothers. He hac! earlier notes] the strong attachment of infant mon- keys to their diaper cloths. This led him to the Plea of a cloth- coverec} wire framework resembling a monkey mother. The concept of a surrogate mother was not new; it was HarIow's genius in creating simple experimental situations in which to use the surrogate that was novel and important. With the aid of graduate student Robert Zimmerman, surrogates were built to replace the biological mothers in attempts to "nor- malize" the behavior of the isolated infants. Some of the sur- rogates were maple of bare wire; others were covered with terrycloth. Other maternal characteristics were adcled, such as protruding rubber nipples for the supply of milk, internal temperature controls for warming or cooling, and mechani- cal arrangements for provicling gentle rocking motion. Sub- sequent studies shower! that an infant's attachment to its sur- rogate mother was clue as much to "contact comfort" as it was

230 BIOGRAPHICAL MEMOIRS to nourishment provided by feecting. Also, the warmth and rocking were found to be important factors. Very important to HarIow was the fact that he hacl now found a way to raise disease-free monkey infants in isolation. Surrogate mothers now proviclec! warmth, comfort, and sustenance in an environment that couIcl be controlled ant! modified as required by experimental research programs. The tempo and scope of the infant monkey research now increased and many studies were undertaken, the results of which often interested! psychoanalysts and challenged psy- choanalytic theory as well as traclitional learning theory. When social development of surrogate-mother-raised and biological-mother-raised monkey infants was compared, it was fount] that natural genetic mothers were significantly better at socializing the young. The importance of peer re- lationships was studiecl by raising infants together and away from aclult animals, ant! it was found that the presence of peers and play opportunities was important to the process of social development. Some of the tiara inclicated that the peer- to-peer interactions were more important than those between mother and infant. HarIow reporter! that one of the most important relationships determining normal sexual behavior as aclults was the peer play cluring infancy and childhood of these monkeys. Laboratory research extended into the abnormal as well as the normal behaviors. The bizarre behaviors resulting from isolation were user} as a basis for studying the long-term effects of isolation per se. Animals separates! from their moth- ers at birth and isolated for periods of six months or longer showed deficits in social, sexual, and other behaviors. The longer the isolation, the worse the deficit. Impregnated, sex- ually mature female isolates showoct few of the normal mother responses expected of rhesus females. In some cases these motherIess-mothers grossly abused their newborns, in-

HARRY FREDERICK HARLOW 231 dicating the importance of early socialization in the learning of proper caretaking and mothering behaviors. "Gove created, love (destroyed ant! love regained" guiclect the affectional-systems research Harry pursued with his wife, Margaret Kuenne HarIow. Love was creates! by parental- infant, peer-peer, anc! surrogate-infant attachments. Iso- lation and separation led to its destruction. Systematic reha- bilitation with a younger monkey (a peer, social-mocleling therapist) resulted in love regained. The reactions of isolate-raised infants separated from their mothers were akin to those of human infants suffering from anaclitic depression as clescribed by British psychiatrist I. A. Bow~by. When peer-rearecT monkeys were separated for several clays similar kinds of clepressed behaviors were notect by HarIow ancI S. Suomi, anti the pattern persisted. Love was regained by rehabilitation, mainly with younger monkeys. A series of studies with infants isolated from birth for various periods of time indicates! that placement with normal, same-age mates or with mature females who expe- r~enced normal mothering was essentially unsuccessful in so- cializing the "depressed" monkeys. The aggressive and dom- inant behaviors of these animals were not changed. However, placement of "(repressed" monkeys with younger normal monkeys immediately upon release from isolation eventually led to play and socialization. This therapeutic technique, de- veloped with Suomi, was later used with some success by oth- ers in rehabilitating institutionalized human children ctiag- nosed as depressives. Harry and Margaret HarIow also collaborated in research on the activities of monkey nuclear families living in adjacent enclosures. The setting allowed for the study of infant inter- actions, fathering, ant! other relationships. HarIow pursued the research on monkey nuclear families and depression un- ti! his retirement from Wisconsin. He believed, however, that

232 BIOGRAPHICAL MEMOIRS his most significant contributions came out of his surrogate studies on love, isolation ejects, anct psychopathology. Har- low often said that when you work with monkeys, you think of human problems, anct he believed that human data gen- eralized to monkeys very nicely, if not vice versa. From a general purview of his many scientific and pro- fessional publications, it is clifficult to pinpoint a central theme. His main goal, it seems, was to study a single species, the rhesus monkey, to learn all he could about its behavior and cognitive processes, and to relate the results to humans. The first decade of his tenure at the University of Wisconsin was dedicated to finding suitable laboratory research facili- ties. Then he strove to find the best empirical methods for working with monkeys and to develop unique tests for as- sessing their sensory and perceptual abilities and the moti- vational circumstances uncler which they worked best at solv- ing graded levels of problems anti tasks. This lect to the development of elaborate and creative test batteries and the WGTA, all of which benefittecl and stimulated his many stu- clents and others throughout the worIcI. He turned briefly to the cat to investigate further the nature of the conditioner! response. During the next three decades Harry and his stu- dents and colleagues used monkeys and the WGTA tests to try to locate in each of the principal regions of the cerebral cortex the extent to which various functions were subserved, eliminated, or modified uncler the influence of anaesthesia, radiation, and ablation. Understancting the neural basis of behavior never seemed to interest him as much as unclerstancTing behavior itself. He concentrated on behavior studies with monkeys throughout the last three decades of his life, opening up new vistas with regard to the cognitive aspects of behavior ant! the social ant! affective consequences of manipulation of the environment on early development. Harry HarIow was a clecluc.tive, qual-

HARRY FREDERICK HARLOW 233 itative empiricist and phenomenologist, whose greatest clis- coveries and contributions resulted from planned serendip- ity. Serendipity even entered into his poetry, which he created quickly and freely anc! often injected into his publications and talks. OTHER SCIENTIFIC AND PROFESSIONAL ACTIVITIES By his own admission, HarIow was more of a nativist, who believer! in the inheritance of characteristics to a greater ex- tent than many of his contemporaries in psychology. Yet this personal inclination was not strongly reflected in his re- search, which in its cognitive and social aspects emphasized environmental influences. He foresaw the importance of bio- chemistry in studies of behavior, ancT in 1958 he credited a volume on the Biological and Biochemical Bases of Behavior with neurophysiologist Clinton N. Woolsey, also of the University of Wisconsin and a member of the National Academy of Sci- ences. The book resulted from a symposium they had plannecl jointly and was a pioneering example of interctisci- plinary research. A year later he colIaboratec! with N. A. Waisman, a pediatrician interested in the genetic basis of phenylketonuria in monkeys and humans. Although not directly involved in the space program, the Wisconsin Laboratory supplied one of the first monkeys sent into space. In 1954 Harry cooperates! with aerospace pioneer D. C. Simons on a series of stratospheric plastic balloon flights to study the elects on monkeys of exposure to radia- tion above 90,000 feet. (At the time, low energy, heavy nu- clear particles of primary cosmic racliation could not be re- produced with available accelerators.) At about the same time, he was involved in investigating the behavioral effects of cortical implantations of radioactive cobalt. During a two-year leave from the University from ~950 to 1952, HarIow served with the Department of the Army in

234 BIOGRAPHICAL MEMOIRS the Pentagon as chief of the Human Resources Branch. In that position, he was responsible for proposing the establish- ment of a Human Resources Research Office for the concluct of psychological research. The recommendation was imple- mented in 1951 and the office established on the campus of The George Washington University, with Meredith Crawford as its director. With Phillip Sapir of the National Institute of Mental Health, he collaborated in the cofounding of the NIMH Small Grants Program. From ~ 95 ~ to ~ 963, HarIow served as editor of the Journal of Comparative and Physiological Psychology, using that position to advance primatology. The proportion of publications (led- icated to primate behavior increased noticeably over the twelve years of his editorship. He also encouraged the pub- lication of articles on the developmental aspects of psychol- ogy and behavior, but the number of manuscripts in those areas proved disappointing. In 1964, one of the seven national Regional Primate Re- search Centers was established adjacent to the University of Wisconsin Primate Laboratory. HarIow servect as its director until 1971. Harry F. HarIow was electecl to the National Academy of Sciences in 195~ and to the American Academy of Arts and Sciences in 1961. At the 52nd meeting of the Society of Ex- perimental Psychologists in 1956, he was awarded the War- ren Medal for "a series of brilliantly conceived experiments on the behavior of monkeys, inclucling studies of motivation, learning, and problem solving." From 1958 to 1959, Hariow served as president of the American Psychological Associa- tion; in 1960, he received its Distinguished Scientific Contri- bution Awarc! for "curiosity and imagination which opened up new areas of research in animal behavior and enhanced the position of comparative psychology." President Lyndon B. Johnson presented him with the Na-

HARRY FREDERICK HARLOW 235 tional Medal of Science in 1967, and in 1973 he received the Gold Medal Awarct of the American Psychological Asso- ciation. HarIow accepted the Kittay Scientific Foundation Award in 1975 for his use of monkey models to study psy- chopathological behaviors. The Primate Laboratory at the University of Wisconsin was dedicated and named in his honor in ~ 984. Harry Frederick Harlow was an unassuming man of many talents. He was a poet and gifted writer, an excellent bridge player, and a pretty goocl locally competitive tennis player. He was generous in sharing time anti ideas with students who wisher! to set up primate laboratories elsewhere. His long walks with professional colleagues and graduate students were legendary, as were his many professional talks. Harlow married Clara Mears in 1932. Two sons, Robert and Richard, were born of this union. In 1946 the marriage was dissolved ant} each party later remarried. In ~ 948, Harry married Margaret Kuenne, a child psychologist of note, with whom he colIaboratec! on numerous research, publication, and editing activities. Two children, Pamela Ann and Jona- thon, were born of this marriage. Margaret diec! of cancer in 1971. Shortly after her death, Harry and Clara, his first wife (then a willow) remarried. Upon his retirement from the University of Wisconsin in 1974, Harry and Clara moved to Tucson, Arizona, where Harry heal an honorary appoint- ment at the University of Arizona. He ant! Clara collaborated on several publications, the most notable being a book en- titlecI, The Human Model: Primate Perspective, published in ~979. Harry died in ~98~.

236 BIOGRAPHICAL MEMOIRS SELECTED BIBLIOGRAPHY 1932 Social facilitation of feeding in the albino rat. I. Genet. Psychol., 41:211-21. With H. Uehling and A. H. Maslow. Comparative behavior of pri- mates: I. Delayed reaction tests on primates from the femur to the orangoutan. J. Comp. Psychol., 13 :313 - 43. With A. H. Maslow. Comparative behavior of primates: II. Delayed reaction tests on primates at Bronx Park Zoo. l. Comp. Psychol., 14:97-107. Comparative behavior of primates: III. Complicated delayed re- action tests on primates. I. Comp. Psychol., 14:241-52. With R. H. Israel. Comparative behavior of primates: IV. Delayed reaction tests on subnormal humans. {. Comp. Psychol., 14:253-62. With R. Stagner. Psychology of feelings and emotions: I. Theory of feelings. Psychol. Rev., 39:570 - 89. Food preferences of the albino rat. I. Genet. Psychol., 41 :430-38. 1933 With H. Yudin. Comparative behavior of primates: V. Delayed re- actions in primates in horizontal and vertical planes. I. Comp. Psychol., 16:143-47. With H. Yudin. Social facilitation of feeding in the monkey and its relation to attitudes of ascendance and submission. J. Comp. Psychol.,16:171-86. With R. Stagner. Effect of complete striate muscle paralysis upon the learning process. I. Exp. Psychol., 16:283-94. With R. Stagner. Psychology of feelings and emotions: II. Theory of emotions. Psychol. Rev., 40: 184 -195 and 368 -80. 1934 With P. Settlage. Comparative behavior of primates: VII. Capacity of monkeys to solve patterned string tests. J. Comp. Psychol., 18:423-35. 1936 With P. Settlage. Concerning the sensory pathway in the condi- tioned reflex. J. Comp. Psychol., 22:279-82.

HARRY FREDERICK HARLOW 237 With P. Settlage. The effect of application of anesthetic agents on circumscribed motor and sensory areas of the cortex. I. Psy- chol., 2: 193-200. The neurophysiological correlates of learning and intelligence. Psychol. Bull., 33 :479-534. 1937 Experimental analysis of the role of the original stimulus in con- ditioned responses in monkeys. Psychol. Rec., 1 :62-68. 1938 With I. Bromer. A test-apparatus for monkeys. Psychol. Rec. 2:434-36. 1939 With P. Settlage. The effect of curarization of the fore part of the body upon the retention of conditioned responses in cats. I. Comp. Psychol., 27:45-48. Recovery of pattern discrimination in monkeys following unilateral occipital lobectomy. J. Comp. Psychol., 27:467-89. With I. Bromer. Comparative behavior of primates: VIII. The ca- pacity of platyrrhine monkeys to solve delayed reaction tests. J. Comp. Psychol., 28:299-304. Forward conditioning, backward conditioning, and pseudo- conditioning in the goldfish. I. Genet. Psychol., 55:49-58. 1940 With F. Toltzien. Formation of pseudo-conditioned responses in the cat. J. Genet. Psychol., 23:367-75. The effects of incomplete curare paralysis upon formation and elicitation of conditioned responses in cats. I. Genet. Psychol., 56:273-82. 1942 With J. Bromer. Acquisition of new responses during inactivation of the motor, premotor, and somesthetic cortex in the monkey. I. Genet. Psychol., 26:299-313. Response by rhesus monkeys to stimuli having multiple sign values. In: Studies in Personality, ed. Q. McNemar and M. Merrill, pp. 105-23. New York: McGraw-Hill Book Co.

238 BIOGRAPHICAL MEMOIRS Animal behavior. In: Fields of Psychology, ed. R. H. Seashore, pp. 171-96. New York: Holt, Rinehart & Winston. 1943 With T. Spaet. Solution by rhesus monkeys of multiple sign prob- lems utilizing the oddity technique. J. Comp. Psychol., 35:119- 32. With M. L. Young. Generalization by rhesus monkeys of a problem involving the Weigl principle using the oddity method. I. Comp. Psychol., 36:201-16. Solution by rhesus monkeys of a problem involving the Weigl prin- ciple using the matchin~-from-samnle method. T. ComD. Psv- chol., 36:217-27. O 1 J - - 1- -' Physiological psychology. Part II. Physiological correlates of behav- ior. In: Annual Review of Physiology, ed. i. M. Luck, vol. 5, pp. 465-78. Palo Alto, Calif.: Annual Reviews Inc. With J. Dagnon. Problem solution by monkeys following bilateral removal of the prefrontal areas: I. The discrimination and dis- crimination-reversal problems. i. Exp. Psychol., 32:351-56. With T. Spaet. Problem solution by monkeys following bilateral removal of the prefrontal areas: II. Delayed reaction problems involving use of the matching-from-sample method. }. Exp. Psychol., 32:424-34. With T. Johnson. Problem solution by monkeys following bilateral removal of the prefrontal areas: III. Test of initiation of behav- ior. I. Exp. Psychol., 32:495-500. With T. Spaet. Problem solution by monkeys following bilateral removal of the prefrontal areas: IV. Responses to stimuli having multiple sign values. J. Exp. Psychol., 33:500-7. 1944 Studies in discrimination learning by monkeys: I. The learning of discrimination series and the reversal of discrimination series. I. Genet. Psychol., 30:3-12. Studies in discrimination learning by monkeys: II. Discrimination learning without primary reinforcement. I. Genet. Psychol., 30:13-21. With M. M. Simpson. Solution by rhesus monkeys of a nonspatial delayed response to the color or form attribute of a single stim-

HARRY FREDERICK HARLOW 239 ulus (Weigl principle delayed reaction). J. Comp. Psychol., 37:211-20. 1945 With R. J. Campbell. Problem solution by monkeys following bilat- eral removal of the prefrontal areas: V. Spatial delayed reac- tions. I. Exp. Psychol., 35:110-26. With S. Poch. Discrimination generalization by macaque monkeys to unidimensional and multidimensional stimuli. I. Comp. Psy- chol., 35:353-65. Studies in discrimination learning by monkeys: III. Factors influ- encing the facility of solution of discrimination problems by rhesus monkeys. l. Genet. Psychol., 32:213-27. Studies in discrimination learning by monkeys: IV. Relative diffi- culty of discrimination between stimulus-objects and between comparable patterns with homogeneous and with heteroge- neous grounds. I. Genet. Psychol., 32:317-21. Studies in discrimination learning by monkeys: V. Initial perform- ance by experimentally naive monkeys on stimulus-object and pattern discriminations. i. Genet. Psychol., 33:3-10. Studies in discrimination learning by monkeys: VI. Discrimination between stimuli differing in both color and form, only in color, and only in form. I. Genet. Psychol., 33:225-35. 1946 With M. Noer. Discrimination of ambivalent cue stimuli by ma- caque monkeys. J. Genet. Psychol., 34:165-77. With M. Zable. The performance of rhesus monkeys on series of object-quality and positional discriminations and discrimination reversals. J. Comp. Psychol., 39:13-23. 1947 With P. H. Settlage. Effect of extirpation of frontal areas on learn- ing performance of monkeys. Assoc. Res. Nerv. Ment. Dis., 27:446-59. With E. M. Moss. The role of reward in discrimination learning in monkeys. J. Comp. Physiol. Psychol., 40:333-42. With P. H. Settlage. An effective and nontraumatic method of han- dling monkeys. Science, 106:300.

240 BIOGRAPHICAL MEMOIRS 1948 With G. Andrew. Performance of macaque monkeys on a test of the concept of generalized triangularity. Comp. Psychol. Monogr., 19: No. 1, Serial No. 100. With L. Grandine. Generalization of the characteristics of a single learned stimulus by monkeys. I. Comp. Physiol. Psychol., 41 :327-38. Studying animal behavior. In: Methods of Psychology, ed. T. G. An- drews, pp. 319-47. New York: {ohn Wiley & Sons. With E. M. Moss. Problem solution by monkeys following extensive unilateral decortication and prefrontal lobotomy of the contra- lateral side. J. Psychol., 25:223-26. With P. Settlage and M. Zable. Problem solution by monkeys fol- lowing bilateral removal of the prefrontal areas: VI. Perform- ance on tests requiring contradictory reactions to similar and to identical stimuli. J. Exp. Psychol., 38:50-65. 1949 Physiological psychology. In: Annual Review of Physiology, ed. J. M. Luck, pp. 269-96. Palo Alto, Calif.: Annual Reviews Inc. The formation of learning sets. Psychol. Rev., 56:51-65. With M. K. Harlow. Learning to think. Sci. Am., 181:36-39. With D. R. Meyer. The development of transfer of response to patterning by monkeys. I. Comp. Physiol. Psychol., 42:454-62. 1950 With R. T. Davis and P. H. Settlage. Performance of normal and brain-operated monkeys on mechanical puzzles with and with- out food incentive. J. Genet. Psychol., 77:305-11. Learning and satiation of response in intrinsically motivated com- plex puzzle performance by monkeys. J. Comp. Physiol. Psy- chol., 43:289-94. Performance of catarrhine monkeys on a series of discrimination reversal problems. I. Comp. Physiol. Psychol., 43:321-39. Analysis of discrimination learning by monkeys. J. Exp. Psychol., 40:26-39. With M. K. Harlow and D. R. Meyer. Learning motivated by a ma- nipulation drive. J. Exp. Psychol., 40:228-34.

HARRY FREDERICK HARLOW 1951 241 With D. R. Meyer and P. H. Settlage. A survey of delayed response performance by normal and brain-damaged monkeys. I. Comp. Physiol. Psychol., 44: 17-25. With A. I. Riopelle, P. H. Settlage, and H. W. Ades. Performance of normal and operated monkeys on visual learning tests. i. Comp. Physiol. Psychol., 44:283-89. With D. R. Meyer and P. H. Settlage. The effects of large cortical lesions on the solution of oddity problems by monkeys. }. Comp. Physiol. Psychol., 44:320-26. With D. R. Meyer and H. W. Ades. Retention of delayed responses and proficiency in oddity problems by monkeys with preoccip- ital ablations. Am. I. Psychol., 64:391-96. Thinking. In: Theoretical Foundations of Psychology, ed. H. Helson, pp. 452-505. New York: D. Van Nostrand. Levels of integration along the phylogenetic scale: Learning aspect. In: Social Psychology at Crossroads, ed. J. R. Roher, pp. 121-41. New York: Harper and Bros. Primate learning. In: Comparative Psychology, 3d ea., ed. C. P. Stone, pp. 183-238. New York: Prentice-Hall. Learning theories. In: Current Trends in Psychological Theory, ed. W. Dennis, pp. 57-84. Pittsburgh: University of Pittsburgh Press. 1952 With R. Weiner. The effect of nembutal upon learned perform- ances of the rhesus monkey. I. Genet. Psychol., 46:43-50. With i. M. Warren. Discrimination learning by normal and brain operated monkeys. T. Genet. Psychol., 81:45-52. With D. R. Meyer. Effects of multiple variables on delayed response performance by monkeys. {. Genet. Psychol., 81:53-61. Learning. In: Annual Review of Psychology, ed. C. P. Stone. on. 29- 54. Palo Alto, Calif.: Annual Reviews Inc. With D. R. Meyer. Paired-comparisons scales for monkey rewards. J. Comp. Physiol. Psychol., 45:73-79. With J. M. Warren. Learned discrimination performance by mon- keys after prolonged postoperative recovery from large cortical lesions. i. Comp. Physiol. Psychol., 45:119-26. Functional organization of the brain in relation to mentation and

242 BIOGRAPHICAL MEMOIRS behavior. In: The Biology of Mental Health and Disease, pp. 244- 53. New York: Hoeber and Co. With R. T. Davis, P. H. Settlage, and D. R. Meyer. Analysis of fron- tal and posterior association syndromes in brain-damaged mon- keys. J. Comp. Physiol. Psychol., 45:419-29. With I. M. Warren. Formation and transfer of discrimination learn- ing sets. J. Comp. Physiol. Psychol., 45:482-89. With R. W. Leary, P. H. Settlage, and D. D. Greenwood. Perform- ance on double-alternation problems by normal and brain- injured monkeys. J. Comp. Physiol. Psychol., 45:576-84. 1953 Mice, monkeys, men, and motives. Psychol. Rev., 60:23-32. Motivation as a factor in the acquisition of new responses. In: Cur- rent Theory and Research in Motivation: A Symposium, ed. M. R. ~ones, pp. 24 - 49. Lincoln, Neb.: University of Nebraska Press. Higher functions of the nervous system. In: Annual Review of Phys- iology, ed. V. E. Hall, vol. 15, pp. 493-514. Palo Alto, Calif.: Annual Reviews Inc. With I. C. Fay and I. D. Miller. Incentive size, food deprivation, and food preference. J. Comp. Physiol. Psychol., 46:13-15. 1954 Motivational forces underlying learning. In: Learning Theory, Per- sonality Theory, and Clinical Research (The Kentucky Sympo- sium), pp. 36-53. New York: John Wiley & Sons. With G. E. McClearn. Object discrimination learned by monkeys on the basis of manipulation motives. i. Comp. Physiol. Psy- chol., 47:73-76. With R. A. Butler. Persistence of visual exploration in monkeys. I. Comp. Physiol. Psychol., 47:258-63. With G. E. McClearn. The effect of spatial contiguity on discrimi- nation learning by rhesus monkeys. J. Comp. Physiol. Psychol., 47:391-94. 1955 With N. C. Blazek. Persistence of performance differences on dis- criminations of varying difficulty. J. Comp. Physiol. Psychol., 48:86-89. With L. E. Moon. Analysis of oddity learning by rhesus monkeys. I. Comp. Physiol. Psychol., 48:188-94.

HARRY FREDERICK HARLOW 243 With I. F. Hall and I. M. Warren. The effects of reserpine (serpasil) on the delayed response in monkeys. J. Psychol., 40:159-61. With K. A. Schiltz and P. H. Settlage. Effect of cortical implantation of radioactive cobalt on learned behavior of rhesus monkeys. I. Comp. Physiol. Psychol., 48:432-36. With G. M. French. Locomotor reaction decrement in normal and brain-damaged rhesus monkeys. J. Comp. Physiol. Psychol., 48:496-501. With W. A. Mason and R. R. Rueping. The development of manip- ulatory responsiveness in the infant rhesus monkey. J. Comp. Physiol. Psychol., 52:555-58. The brain and learned behavior. Comput. Autom., 4:6-14. With L. E. Moon and C. P. Bogumill. Some effects of periodic x- radiation. Science, 122: 1-2. 1956 With R. A. Butler. The effects of auditory distraction on the per- formance of monkeys. J. Genet. Psychol., 54: 15 -20. With R. A. Butler. Discontinuous pursuit performance by rhesus monkeys. J. Genet. Psychol., 54:21-30. With L. E. Moon. The effects of repeated doses of total-body-x- radiation on motivation and learning in rhesus monkeys. J. Comp. Physiol. Psychol., 49:60-65. With A. M. Schrier. Effect of amount of incentive on discrimination learning by monkeys. J. Comp. Physiol. Psychol., 49: 117-22. With A. M. Schrier and D. G. Simons. Exposure of primates to cosmic radiation above 90,000 feet. I. Comp. Physiol. Psychol., 49: 195-200. With N. C. Blazek and G. E. McClean. Manipulatory motivation in the infant rhesus monkey. I. Comp. Physiol. Psychol., 49:444- 48. With W. A. Mason and N. C. Blazek. Learning capacities of the infant rhesus monkey. J. Comp. Physiol. Psychol., 49:449-53. Current and future advances in physiological psychology. Am. Psy- chol., 11:273-77. 1957 With L. H. Hicks, Jr. Discrimination learning theory: Uniprocess vs. duoprocess. Psychol. Rev., 64:104-9. Experimental analysis of behavior. Am. Psychol., 12:485-90.

244 BIOGRAPHICAL MEMOIRS With R. A. Butler. Discrimination learning and learning sets to vi- sual exploration incentives. I. Genet. Psychol., 57:257-85. With I. E. Farber and L. l. West. Brainwashing, conditioning, and DDD (debility, dependency, and dread). Sociometry, 20:271-85. With I. M. Warren, R. W. Leary, and G. M. French. Function of the association cortex in monkeys. Brit. I. An. Beh., 4: 132-38. With A. M. Schrier. Direct manipulation of the relevant cue and difficulty of discrimination. J. Comp. Physiol. Psychol., 50:576- 80. 1958 With W. A. Mason. Learned approach by infant rhesus monkeys to the sucking situation. Psychol. Rep., 4:79-82. With W. A. Mason. Formation of conditioned responses in infant monkeys. J. Comp. Physiol. Psychol., 51:68-70. With W. A. Mason. Performance of infant rhesus monkeys on a spatial discrimination problem. l. Comp. Physiol. Psychol., 51: 71-74. With A. M. Schrier. Effect of reserpine on avoidance of humans by rhesus monkeys. J. Genet. Psychol., 59:149-55. The evolution of learning. In: Behavior and Evolution, ed. A. Roe and G. Simpson, pp. 269-90. New Haven: Yale University Press. With R. R. Zimmermann. The development of affectional re- sponses in infant monkeys. Proc. Am. Philos. Soc., 102:501-9. Ed. Harry Harlow and C. N. Woolsey. Behavioral contributions to interdisciplinary research. In: Biological and Biochemical Bases of Behavior, pp. 3-23. Madison: University of Wisconsin Press. The nature of love. Am. Psychol., 13:763 -85. 1959 Learning set and error factor theory. In: Psychology: A Study of a Science, ed. S. Koch, pp. 492-538. New York: McGraw-Hill Book Co. With W. A. Mason. Initial responses of infant rhesus monkeys to solid foods. Psychol. Rep., 5:193-99. Basic social capacity of primates. Hum. Biol., 31:40-53. With M. Levine and B. Levinson. Trials per problem as a variable in the acquisition of discrimination learning set. I. Comp. Phys- iol. Psychol., 52 :396-98.

HARRY FREDERICK HARLOW 245 With R. R. Zimmermann. Affectional responses in the infant mon- key. Science, 130:421-32. Love in infant monkeys. Sci. Am., 200:68-74. The development of learning in the rhesus monkey. Am. Sci., 47:459-79. With M. Levine. Learning sets with one- and twelve-trial oddity problems. Am. J. Psychol., 72:253-357. With H. A. Waisman, H. L. Wang, and R. R. Sponholz. Experi- mental phenylketonuria in the monkey. Proc. Soc. Exp. Biol. Med., 101:864-65. With W. A. Mason and R. R. Rueping. The development of manip- ulatory responsiveness in the infant rhesus monkey. I. Comp. Physiol. Psychol., 52:555-58. 1960 With M. K. Harlow, R. R. Rueping, and W. A. Mason. Performance of infant rhesus monkeys on discrimination learning, delayed response, and discrimination learning set. l. Comp. Physiol. Psychol., 53: 113 -21. Affectional behavior in the infant monkey. In: The Central Nervous System and Behavior, ed. M. A. B. Brazier, pp.307-57. New York: Josiah Macy, fir. Foundation. Primary affectional patterns in primates. Am. I. Orthopsychiatry, 30:676-84. With H. A. Waisman, H. L. Wang, and G. Palmer. Phenylketonuria in infant monkeys. Nature, 188: 1124-25. With K. Akert, O. S. Orth, and K. A. Schiltz. Learned behavior of rhesus monkeys following neonatal bilateral prefrontal lobot- omy. Science, 132: 1944-45. 1961 With A. I. Blomquist. The infant rhesus monkey program at the University of Wisconsin Primate Laboratory. Proc. An. Care Panel, 11:57-64. With L. R. Cooper. Note on a rebus monkey's use of a stick as a weapon. Psychol. Rep., 8:418. With M. Levine and T. Pontrelli. Supplementary report: The ef- fects of problem length on transfer during learning-set per- formance. I. Exp. Psychol., 61: 192. The development of affectional patterns in infant monkeys. In:

246 BIOGRAPHICAL MEMOIRS Determinants of Infant Behavior, ed. B. M. Foss, pp. 75-97. Lon- don: Methuen. With M. K. Harlow. A study of animal affection. Nat. Hist., 70~101:48-55. With A. I. Riopelle. Stimulus and reward displays in discrimination learning. J. Genet. Psychol., 98:183-86. With W. A. Mason. The effects of age and previous training on patterned-strings performance of rhesus monkeys. I. Comp. Physiol. Psychol., 54:704-9. 1962 Effects of radiation on the central nervous system and on behav- ior—general survey. In: Response of the Nervous System to Ionizing Radiation, ed. T. i. Haley and R. S. Snider, pp. 627-44. New York: Academic Press. Development of affection in primates. In: Roots of Behavior: Genetics, Instinct, and Socialization in Animal Behavior, ed. E. Bliss, pp. 157- 66. New York: Harper (Hoeber). The heterosexual affectional system in monkeys. Am. Psychol., 17:1-9. With T. E. Cadell and H. A. Waisman. PEG changes in experimen- tal phenylketonuria. Electroencephalogr. Clin. Neurophysiol., 14:540-43. With M. K. Harlow. The effect of rearing conditions on behavior. Bull. Menninger Clin., 26:213-24. With M. K. Harlow. Principles of primate learning. In: Lessons from Animal Behavior for the Clinician, ed. S. A. Barnett, pp. 37-48. London: National Spastics Society. Development of the second and third affectional systems in ma- caque monkeys. In: Research Approaches to Psychiatric Problems, ed. T. T. Tourlentes, S. L. Pollack, and H. E. Himwich, pp.209- 29. New York: Grune & Stratton. With G. M. French. Variability of delayed-reaction performance in normal and brain-damaged rhesus monkeys. l. Neurophysiol., 25: 585-99. With i. M. Lockhart. The influence of spatial configuration and percentage of reinforcement upon oddity learning. I. Comp. Physiol. Psychol., 55:495-501. With M. K. Harlow. Social deprivation in monkeys. Sci. Am., 207:136-46.

HARRY FREDERICK HARLOW 247 With I. E. King. Effect of ratio of trial 1 reward to nonreward on the discrimination learning of macaque monkeys. J. Comp. Physiol. Psychol., 55:872-75. With B. Seay and E. Hansen. Mother-infant separation in monkeys. J. Child Psychol. Psychiatry, 3:123-32. The effects of early experience on affectional behavior in monkeys. In: Biological Influences in Mental Health, pp. 27-33. Fifth annual research conference. Michigan Department of Mental Health. 1963 With H. A. Cross and H. I. Fletcher. Ejects of prior experience with test stimuli on learning-set performance of monkeys. J. Comp. Physiol. Psychol., 56:204-7. With G. M. Sterritt and E. Goodenough. Learning set develop- ment: Trials to criterion vs. six trials per problem. Psychol. Rep., 13:267-71. An experimentalist views the emotions. In: The Expression of Emotion in Man, ed. P. H. Knapp, pp. 254-65. New York: International Universities Press. With M. K. Harlow and E. W. Hansen. The maternal affectional system of rhesus monkeys. In: Maternal Behavior in Mammals, ed. H. L. Rheingold, pp. 254-81. New York: John Wiley & Sons. The maternal affectional system. In: Determinants of InfantBehav- iourII, ed. B. M. Foss, pp. 3-33. London: Methuen. 1964 With K. Akert and K. A. Schiltz. The effects of bilateral prefrontal lesions on learned behavior of neonatal, infant, and preadoles- cent monkeys. In: The Frontal Granular Cortex and Behavior, ed. J. M. Warren and K. Akert, pp. 126-48. New York: McGraw- Hill Book Co. Early social deprivation and later behavior in the monkey. In: Un- fin~shed Tasks in the Behavioral Sciences, ed. A. Abrams, H. H. Gar- ner, and I. E. P. Tomal, pp. 154-73. Baltimore: Williams & Wil- kins. A behavioral approach to psychoanalytic theory. Sci. Psychoanal., 7:93-1 13. With B. Seay and B. K. Alexander. Maternal behavior of socially

248 BIOGRAPHICAL MEMOIRS deprived rhesus monkeys. J. Abnorm. Soc. Psychol., 69:345- 54. With G. L. Rowland and G. A. Griffin. The effect of total social deprivation on the development of monkey behavior. In: Recent Research on Schizophrenia, Psychiatric Research Report 19, ed. P. Solomon and B. C. Glueck, pp. 116-35. Washington, D.C.: American Psychiatric Association. 1965 With H. A. Waisman. Experimental phenylketonuria in infant monkeys. Science, 147:685-95. With H. A. Cross. Prolonged and progressive effects of partial iso- lation on the behavior of macaque monkeys. }. Exp. Res. Pers., 1 :39-49. With M. K. Harlow. The effects of early social deprivation on pri- mates. In: Desafferentation Experimentale Et Clinique, ed. I. de Aju- riaguerra, pp. 67-77. Geneva, Switzerland: Georg & Cie S.A. With R. O. Dodsworth and M. K. Harlow. Total social isolation in monkeys. Proc. Natl. Acad. Sci. USA, 54:90-97. Ed. Harry Harlow, A. M. Schrier, and F. Stollnitz. Behavior of Non- human Primates, vol. I and II. New York: Academic Press. With M. K. Harlow. The affectional systems. In: Behavior of Non- human Primates, vol. II, ed. Harry Harlow, A. M. Schrier, and F. Stollnitz, pp. 287-334. New York: Academic Press. With B. K. Alexander. Social behavior of juvenile rhesus monkeys subjected to different rearing conditions during the first six months of life. Zool. J. Physiol., 71:489-508. With R. L. Raisler. Learned behavior following lesions of posterior association cortex in infant, immature, and preadolescent mon- keys. J. Comp. Physiol. Psychol., 60:167-74. Sexual behavior in the rhesus monkey. In: Sex and Behavior, ed.F. A. Beach, pp. 234-65. New York: John Wiley & Sons. With B. Seay. Maternal separation in the rhesus monkey. }. Nerv. Ment. Dis., 140:434-41. With G. Griffin. Induced mental and social deficits in rhesus mon- keys. In: The Biosocial Basis of Mental Retardation, ed. S. F. Osler and R. E. Cooke, pp. 87-106. Baltimore: The Johns Hopkins Press. Total social isolation: Effects on macaque behavior. Science, 148:666.

HARRY FREDERICK HARLOW 1966 249 With M. K. Harlow. Affection in primates. Discovery, 27:11-17. With M. K. Harlow, R. O. Dodsworth and G. L. Arling. Maternal behavior of rhesus monkeys deprived of mothering and peer association in infancy. Proc. Am. Philos. Soc., 110:58-66. With G. D. Mitchell, E. I. Raymond, and G. C. Ruppenthal. Long- term effects of total social isolation upon behavior of rhesus monkeys. Psychol. Rep., 18:567-80. The primate socialization motives. Trans. Stud. Coll. Physicians Philadelphia, 33:224-37. With E. W. Hansen and R. O. Dodsworth. Reactions of rhesus monkeys to familiar and unfamiliar peers. }. Comp. Physiol. Psychol., 61:274-79. With W. D. Joslyn, M. G. Senko and A. Donn. Behavioral a~r~ct.s of reproduction in primates. J. Anim. Sci., 25:49-65. - or ~ rig With M. K. Harlow. Effect de la privation precoce de contacts so- ciaux chez les primates. Rev. Med. Psychosom. Psychol. Med., 8:1-24. With G. A. Griffin. Effects of three months of total social depriva- tion on social adjustment and learning in the rhesus monkey. Child Dev., 37:533-47. With M. K. Harlow. Learning to love. Am. Sci., 54:234-72. With G. D. Mitchell, G. C. Ruppenthal, and E. I. Raymond. Long- term effects of multiparous and primiparous monkey mother rearing. Child Dev., 37:781-91. With B. Seay. Mothering in motherless mother monkeys. Br. I. Soc. Psychiatry, 1:63-69. 1967 With M. K. Harlow. Reifungs-faktoren im Sozialen Verhalten. Psyche: Z. Psychoanal. Anwendung, 21:193-210. With A. S. Chamove and G. D. Mitchell. Sex differences in the in- fant-directed behavior of preadolescent rhesus monkeys. Child Dev., 38:329-35. With G. D. Mitchell, G. A. Griffin, and G. W. M011er. Repeated ma- ternal separation in the monkey. Bull. Psychon. Soc.,8: 197-98. With M. K. Harlow. The young monkeys. Psychol. Today, 1:41-47. With G. L. Arling. Effects of social deprivation on maternal behav- ior of rhesus monkeys. J. Comp. Physiol. Psychol., 64:371-77.

250 BIOGRAPHICAL MEMOIRS 1968 With A. I. Blomquist, C. I. Thompson, K. A. Schiltz, and M. K. Harlow. Effects of induction age and size of frontal lobe lesions on learning in rhesus monkeys. In: The Neuropsychology of De- velopment: A Symposium, ed. R. L. Isaacson, pp. 79-120. New York: John Wiley & Sons. With G. R. Kerr, A. S. Chamove, and H. A. Waisman. Fetal PKU: The effect of maternal hyperphenylalaninemia during preg- nancy in the rhesus monkey (Macaca mulatto). Pediatrics,42:27- 36. Learning and memory in primates. In: Attuali Orientamenti Della Ricerca Sull Apprendimento E La Memory, ed. D. Bovet, F. Bovet- Nitti, and S. Oliverio, pp. 139-56. Rome: Accademia Nazionale dej Lincei. With G. W. .M011er and G. D. Mitchell. Factors affecting agonistic communication in rhesus monkeys (Macaca mulatta). Behav- iour, 31:339-57. A primate. Science, 165:274. 1969 With M. K. Harlow. Effects of various mother-infant relationships on rhesus monkey behaviors. In: Determinants of Infant Behav- iour, ed. B. M. Foss, pp. 15-36. London: Methuen. With C. I. Thompson and I. S. Schwartzbaum. Development of so- cial fear after amygdalectomy in infant rhesus monkeys. Phys- iol. Behav., 4:249-54. William James and instinct theory. In: William fames: Unfinished Business, ed. R. B. McCleod, pp. 21-30. Washington, D.C.: American Psychological Association. Age-mate or peer affectional system. In: Advances in the Study of Behavior, ed. D. S. Lehrman, R. A. Hinde, and E. Shaw, vol. 2, pp. 333-83. New York: Academic Press. With G. R. Kerr, A. S. Chamove, and H. A. Waisman. The devel- opment of infant monkeys fed low phenylalanine diets. Pediatr. Res., 3:305-12. With S. J. Suomi. Apparatus conceptualization for psychopathol- ogical research in monkeys. Behav. Res. Methods Instrum., 1 :247-50. The anatomy of humour. Impact Sci. Soc., 19:225 - 39.

HARRY FREDERICK HARLOW 251 With G. R. Kerr and A. S. Chamove. Environmental deprivation: Its eject on the growth of infant monkeys. }. Pediatr., 75:833- 37. With C. S. Furchner. Preference for various surrogate surfaces among infant rhesus monkeys. Bull. Psychon. Sci., 17:279-80. With K. A. Schiltz and M. K. Harlow. Effects of social isolation on the learning performance of rhesus monkeys. In: Proceedings of the 2nd International Congress of Pr~matology, ed. C. R. Carpenter, vol. 1, pp. 178-85. Basel/New York: Karger. 1970 With S. l. Suomi. The nature of love simplified. Am. Psychol., 25: 161-68. With S. I. Suomi and I. K. Lewis. Effect of bilateral frontal lobec- tomy on social preferences of rhesus monkeys. J. Comp. Phys- iol. Psychol., 70:448-453. With S. J. Suomi and W. T. McKinney. Experimental production of depression in monkeys. Mainly Monkeys, 1:6-12. With A. C. Deets. Nipple preferences in nursing singleton- and twin-reared rhesus monkey infants. Dev. Psychol., 2: 159-62. With C. I. Thompson, A. J. Blomquist, and K. A. Schiltz. Learning in rhesus monkeys after varying amounts of prefrontal lobe destruction during infancy and adolescence. Brain Res., 18: 343-53. With S. I. Suomi. Induction and treatment of psychiatric states in monkeys. Proc. Natl. Acad. Sci. USA., 66:241. With S. J. Suomi. Induced psychopathology in monkeys. Eng. Sci., 33:8-14. With A. S. Chamove. Exaggeration of self-aggression following al- cohol ingestion in rhesus monkeys. J. Abnorm. Psychol., 75:207-9. With l. W. Davenport and A. S. Chamove. The semiautomatic Wis- consin general test apparatus. Behav. Res. Methods Instrum., 2: 135-38. With K. A. Schiltz, A. I. Blomquist, and C. I. Thompson. Effects of combined frontal and temporal lesions on learned behaviors in rhesus monkeys. Proc. Natl. Acad. Sci. USA., 66:577-82. With A. S. Chamove and H. A. Waisman. Abnormal social behavior in phenylketonuric monkeys. J. Abnorm. Psychol., 76:62-68. With A. C. Deets. S. D. Singh, and A. I. Blomquist. Effects of bilat-

252 BIOGRAPHICAL MEMOIRS eral lesions of the frontal granular cortex on the social behavior of rhesus monkeys. I. Comp. Physiol. Psychol., 72:452-61. With M. K. Harlow. Developmental aspects of emotional behavior. In: Physiological Correlates of Emotion, ed. P. Black, pp. 37-58. New York: Academic Press. With S. I. Suomi and C. l. Domek. Effect of repetitive infant-infant separation of young monkeys. I. Abnorm. Psychol., 76: 161-72. With S. I. Suomi and G. P. Sackett. Development of sex preference in rhesus monkeys. Dev. Psychol., 3:326-36. With A. C. Deets and A. I. Blomquist. Effects of intertrial interval and trial 1 reward during acquisition of an object discrimination learning set in monkeys. I. Comp. Physiol. Psychol., 73:501-5. 1971 With W. T. McKinney and S. I. Suomi. Depression in primates. Am. I. Psychiatry, 1 27: 1 3 1 3 -20. With S. I. Suomi. Abnormal social behavior in young monkeys. In: Exceptional Infant: Studies in Abnormalities, ed. I. Hellmuth, vol. 2, pp. 483-529. New York: Brunner Mazel. With I. L. McGaugh and R. F. Thompson. Psychology. San Fran- cisco: Albion Publishing Co. With A. I. Blomquist and A. C. Deets. Effects of manipulating in- centive visibility during the baiting phase of delayed-response problems. Learn. Motiv., 2:67-74. With S. l. Suomi. Social recovery of isolation-reared monkeys. Proc. Natl. Acad. Sci. USA, 68:1534-38. With L. A. Rosenblum. Maturational variables influencing sexual posturing in rhesus monkeys. Arch. Sex. Behav., 1:73-78. Early problem learning and early social learning. In: The Second Western Symposium on Learning: Early Learning, ed. M. E. Meyer, pp. 41-75. Bellingham: Western Washington State College. With M. K. Harlow and S. J. Suomi. From thought to therapy: Les- sons from a primate laboratory. Am. Sci., 59:538-49. With S. I. Suomi and S. D. Kimball. Behavioral effects of prolonged partial social isolation in the rhesus monkey. Psychol. Rep., 29:1171-77. With l. P. Cluck. The effects of deprived and enriched rearing con- ditions on later learning: A review. In: Cognitive Process of Non- human Primates, ed. L. E. Jarrard, pp. 103 - 19. New York: Aca- demic Press. With M. K. Harlow, K. A. Schiltz, and D. I. Mohr. The eject of

HARRY FREDERICK HARLOW 253 early adverse and enriched environments on the learning abil- ity of rhesus monkeys. In: Cognitive Processes of Nonhuman Pri- mates, ed. L. E. Jarrard, pp.121 - 48. New York: Academic Press. With S. I. Suomi. Production of depressive behaviors in young monkeys. I. Autism Child. Schizophren., 1 :246-55. With M. K. Harlow. Psychopathology in monkeys. In: Experimental Psychopathology, ed. H. D. Kimmel, pp. 203-29. New York: Aca- demic Press. With C. I. Thompson, A. J. Blomquist, and K. A. Schiltz. Recovery of function following prefrontal lobe damage in rhesus mon- keys. Brain Res., 35:37-48. With W. T. McKinney, Jr., R. G. Eising, E. C. Moran, and S. J. Suomi. Effects of reserpine on the social behavior of rhesus monkeys. Dis. Nerv. Sys., 32:735-41. With W. T. McKinney, Jr. Nonhuman primates and psychoses. J. Autism Child. Schizophren., 1 :368-75. 1972 With W. T. McKinney, fir. and S. I. Suomi. Vertical-chamber con- finement of juvenile-age rhesus monkeys. Arch. Gen. Psychia- try, 26-223-28. With S. I. Suomi. Social rehabilitation of isolate-reared monkeys. Dev. Psychol., 6:487-96. With M. K. Harlow, E. W. Hansen, and S. J. Suomi. Infantile sex- uality in monkeys. Arch. Sex. Behav., 2: 1-7. Love created love destroyed love regained. In: Modeles Anamaux Du Comportement Humain, No. 198, pp. 13 - 60. Paris: Editions du Centre National de la Recherche Scientifique. With S. J. Suomi. Depressive behavior in young monkeys subjected to vertical chamber confinement. I. Comp. Physiol. Psychol., 180:11-18. With W. T. McKinney and S. J. Suomi. Repetitive peer separations of juvenile-age rhesus monkeys. Arch. Gen. Psychiatry,27:200- 4. With I. P. Gluck and S. i. Suomi. Generalization of behavioral data between nonhuman and human animals. Am. Psychol., 27: 709-16. With M. K. Harlow. The language of love. In: Communication and A§ect, ed. T. Alloway, L. Krames, and P. Pliner, pp. 1-18. New York: Academic Press. With I. B. Sidowski and S. I. Suomi. Enhancing social attachment .

254 BIOGRAPHICAL MEMOIRS through fear. A study of infant monkeys. Bull. Psychon. Soc., 29:323. With A. S. Chamove and H. I. Eysenck. Personality in monkeys: Factor analysis of rhesus. Q. I. Exp. Psychol., 24:496-504. 1973 With W. T. McKinney, tr., and S. i. Suomi. Methods and models in primate personality research. In: Individual Differences in Chil- dren, ed. [. C. Westman, pp. 265 - 87. New York: John Wiley & Sons. With A. I. Blomquist and A. C. Deets. Effects of list-length and first-trial reward upon concurrent discrimination performance. Learn. Motiv., 4:28 - 39. With M. A. Novak. Psychopathological perspectives. Perspec. Biol. Med., 16:461-78. With L. D. Young, S. J. Suomi, and W. T. McKinney, Jr. Early stress and later response to separation in rhesus monkeys. Am. I. Psy- chiatry, 130:400-5. With D. M. Baysinger and P. E. Plubell. A variable-temperature surrogate-mother for studying attachment in infant monkeys. Behav. Res. Methods Instrum., 5:269-72. With K. A. Schiltz, C. I. Thompson, D. J. Mohr, and A. J. Blom- quist. Learning in monkeys after combined lesions in frontal and anterior temporal lobes. I. Comp. Physiol. Psychol., 83:271-77. With l. P. Cluck and K. A. Schiltz. Differential effect of early en- richment and deprivation on learning in the rhesus monkey (Macaca mulatta). J. Comp. Physiol. Psychol., 84:598-604. With A. S. Chamove. Avoidance learning in phenylketonuric mon- keys. J. Comp. Physiol. Psychol., 84:605-12. With A. S. Chamove and L. A. Rosenblum. Monkeys (Macaca mu- latta) raised only with peers. A pilot study. Anim. Behav., 21 :316-25. With W. T. McKinney, [r., and S. I. Suomi. New models of separa- tion and depression in rhesus monkeys. In: Separation and Depression, Clinical and Research Aspects, ed. I. P. Scott and E. C. Senay, No. 94, pp. 53-66. Washington, D.C.: American Asso- ciation for the Advancement of Science. With S. l. Suomi and M. L. Collins. Effects of permanent separa- tion from mother on infant monkeys. Dev. Psychol.; 9:376-84.

HARRY FREDERICK HARLOW 255 With A. S. Chan~ove and G. R. Kerr. Learning in monkeys fed ele- vated amino acid diets. l. Med. Primatol., 2:223-35. With P. E. Plubell and C. M. Baysinger. Induction of psychological death in rhesus monkeys. }. Autism Child. Schizophr., 3:299- 307. 1974 Induction and alleviation of depressive states in monkeys. In: Eth- ology and Psychiatry, ed. N. F. White, pp. 197-208. Toronto: Uni- versity of Toronto Press. Les affectivity. In: L 'Attachement, ed. R. Zazzo, pp. 58-72. Paris: Delachaux et Niestle. With H. E. Lauersdorf. Sex differences in passion and play. Per- spec. Biol. Med., 17:348-60. With G. C. Ruppenthal, M. K. Harlow, C. D. Eisele, and S. I. Suomi. Development of peer interactions of monkeys reared in a nuclear-family environment. Child Dev., 45:670-82. Maternal and peer affectional deprivation in primates. In: Experi- mental Behaviour: A Basis for the Study of Mental Disturbance, ed. J. Cullen, pp. 85-98. Dublin: Irish University Press. With S. J. Suomi. Induced depression in monkeys. Behav. Biol., 12:273-96. With A. C. Deets. Adoption of single and multiple infants by rhe- sus monkey mothers. Primates, 15: 193 -204. With S. S. Suomi and M. A. Novak. Reversal of social deficits pro- duced by isolation rearing in monkeys. }. Hum. Evol., 3:527- 34. 1975 With S. I. Suomi. Generalization of behavior from monkey to man. In: Psychology, ed. G. Lindzey, C. Hall, and R. F. Thompson, on. 34-35. New York: Worth. A ' ~ ~ With S. }. Suomi. Ejects of differential removal from group on social development of rhesus monkeys. J. Child Psychol. Psy- chiatry, 16: 149-64. Ethology. In: Comprehensive Textbook of Psychiatry, ed. A. M. Free- dom, H. K. Kaplan, and B. I. Sadock, pp. 317-36. Baltimore: Williams & Wilkins. With P. M. Nealis, A. Carpentier, and S. I. Suomi. Dynamic stimu-

256 BIOGRAPHICAL MEMOIRS lus display for the WGTA. Behav. Res. Methods Instrum., 7:291-93. With S. J. Suomi. Experienceas tempranas y psicopatologia indu- cida en monos rhesus. Revista Latinoamer. Psicol., 7:205-29. With C. E. Mears. Play: Early and eternal. Proc. Natl. Acad. Sci. USA, 72:1878-82. Lust, latency and love Simian secrets of successful sex. I. Sex. Res., 11:79-90. With M. A. Novak. Social recovery of monkeys isolated for the first year of life: 1. Rehabilitation and therapy. Dev. Psychol., 1 1 :453-65. With I. S. Meyer, M. A. Novak, and R. E. Bowman. Behavioral and hormonal effects of attachment object separation in surrogate- peer-reared and mother-reared infant rhesus monkeys. Dev. Psychol., 8:425-36. With S. }. Suomi, C. D. Eisele, and S. A. Grady. Depressive behav- ior in adult monkeys following separation from family environ- ment. I. Abnorm. Psychol., 84:576-78. With S. I. Suomi. The role and reason of peer relationships in rhe- sus monkeys. In: Friendship and Peer Relations, ed. M. Lewis and L. A. Rosenblum, pp. 153 - 85. New York: John Wiley & Sons. With W. T. McKinney and S. J. Suomi. Experimental psychopath- ology in nonhuman primates. In: New Psychiatric Frontiers, Amer- ican Handbook of Psychiatry, ed. D. A. Hamburg and H. K. Bro- die, vol. 6, 2 ea., pp. 310-34. New York: Basic Books. Monkeys, men, mice, and motives. In: Psychological Research: The Inside Story, ed. M. H. Siegel and H. P. Zeigler, pp. 3-22. New York: Harper & Row. 1976 With S. I. Suomi, M. L. Collins, and G. C. Ruppenthal. Effects of maternal and peer separations on young monkeys. }. Child Psy- chol. Psychiatry, 17:101 - 12. With G. C. Ruppenthal, G. L. Arling, G. P. Sackett, and S. J. Suomi. A 10-year perspective of motherless-mother monkey behavior. I. Abnorm. Psychol., 85:341-49. With S. J. Suomi. The facts and functions of fear. In: Emotions and Anxiety: New Concepts, Methods, and Applications, ed. M. Zucker- man and C. D. Spielberger, pp. 3-34. Hillsdale, N.J.: Lawrence Erlbaum Associates.

HARRY FREDERICK HARLOW 257 With S. I. Suomi and R. DeLizio. Social rehabilitation of separa- tion-induced depressive disorders in monkeys. Am. J. Psychia- try, 133:1279-85. 1977 With C. Mears. The power and passion of play. New Sci., 73:336- 38. With P. M. Nealis and S. I. Suomi. The effects of stimulus move- ment on discrimination learning by rhesus monkeys. Bull. Psy- chon. Soc., 10:161-64. With S. l. Suomi. Production and alleviation of depressive behav- iors in monkeys. In: Psychopathology: Experimental Models, ed. I. Maser and M. E. P. Seligman, pp. 131-73. San Francisco: W. H. Freeman. Birth of the surrogate mother. In: Discovery Processes in Modern Biol- ogy, ed. W. R. Klemm, pp. 133-50. Huntington, N.Y.: R. E. Krieger. With S. l. Suomi. Early separation and behavioral maturation. In: Genetics, Environment and Intelligence, ed. A. Oliverio, pp. 197- 214. Amsterdam: Elsevier. 1978 With S. I. Suomi. Early experience and social development in rhe- sus monkeys. In: Social and Personality Development, ed. M. E. Lamb, pp. 252 - 71. New York: Holt, Rinehart & Winston. With C. E. Mears. The nature of complex, unlearned responses. In: The Development of Affect, ed. M. Lewis and L. A. R. Rosen- blum, pp. 257-74. New York: Plenum Press. 1979 With C. Mears. The Human Model: Primate Perspectives. Washington, D.C.: V. H. Winston & Sons (Halsted Press).

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