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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

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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

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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.

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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

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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 demonstratedcontrary 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-

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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-

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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

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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

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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

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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-

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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

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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.

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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.

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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.

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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-

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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

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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 Aect, 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 .

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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.

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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-

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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.

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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).