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HARRY FREDERICK HARLOWOctober 31, 1905-December 6, 1981BY JOSEPH B. SIDOWSKI ANDDONALD B. LINDSLEYHARRY 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 namelegally to HarIow while in college. After forty-four years ofassociation with the University of Wisconsin (1930-1974), hebecame 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 YEARSHarry HarIow, as he himself described it, was a shy, retir-ing, and callow youth when he began his college studies atReect College in Portland, Oregon, in 1923. After one yearhe decidecl to follow his brother to Stanford University,where he wouIc! receive his B.A. degree in 1927, majoring inpsychology. 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 ahumorous manner remained, later contributing greatly to hissuccess both as a teacher and a professional lecturer.While still an undergraduate, HarIow supported himself219

220BIOGRAPHICAL MEMOIRSworking as an assistant to the experimental psychologist Wal-ter R. Miles, who was elected to the National Academy ofSciences in 1933. As a graduate student at Stanford, HarIowcame under the tutelage of Calvin P. Stone, who was electedto the Academy in 1943. As a graduate student, Harry helda teaching assistantship uncler Paul R. Darnworth in socialpsychology and research assistantships under Stone in be-havioral studies on rats. His cloctoral dissertation dealt withthe social facilitation of eating behavior in rats, combiningelements of his ongoing experiments as an assistant. Muchlater, Harlow sail! that he learner! scientific methoclology andtechniques from Stone, but he always consiclerec] Miles hismoral and ethical mentor.He admirect Lewis W. Terman, then head of the depart-ment of psychology, and learned about theory in psychologyfrom him. Terman had been elected to the Academy in 1928.Towarc! the ens! of Harry's second graduate year, Termanwrote to Harlow's mother of his great progress in psychologyant! 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 ajunior college position because of a speech defect, which theythought interfered with his ability to articulate clearly andsometimes brought forth smiles when he said "wat" for rat!Despite this advice, he accepted a position as an assistantprofessor of psychology at the University of Wisconsin in1930, where he regularly taught the large introductory classin psychology. With cleterminec! application, his diction andenunciation steadily improved, ant] he became one of themost 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 HARLOW221most entertaining, elective, and sought after speakers in allof psychology.Hired as a comparative animal psychologist, Harlow ar-rived at the University of Wisconsin in 1930 to learn thatthere was no animal laboratory. However, he soon found acramped cubicle in which to house his rats, which happenedto be just below the office of the Dean of Men who didn'tappreciate the odors wafting upward.As a result, Harry was displaced from that location andgiven a small space in the University Medical School. Therehe began studies of the social facilitation of feeding responsesin monkeys, an extension of his doctoral research with rats.But that space, too, proved vulnerable and temporary, andhis first steps into a major career dedicated to the study ofnonhuman primate behavior began at a bridge party, whenthe wife of the chairman of the psychology department sug-gested that he study primates at the local Vilas Park Zoo. TheZoo 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 anunexpected turn in his career.PRIMATE LABORATORIES AND RESEARCHHarlow's first primate research facility consisted mainly ofa few tables, a test tray, and test objects at the Vilas Park Zoo.In 1932 the University of Wisconsin made available to him avery small, two-story structure that had previously been theForest Products Laboratory. It was badly in need of renova-tion. With his own meager funds and the aid of WalterGrether, Paul Settlage and other graduate students this wasaccomplished. The result was a usable research facility andthe first real primate laboratory in Wisconsin's Departmentof Psychology.

222BIOGRAPHICAL MEMOIRSAcquiring a small colony of monkeys, HarIow and hisgraduate students enthusiastically began developing new ant!unique ways to study primate behavior, both qualitatively andquantitatively. Using the oddity principle and matching-from-sample procedures they were able to study perceptualdiscrimination involving figures and patterns on visual clis-plays or objects that cliffered in color, size, shape, or texture.By introducing time clelays between stimulus presentationand 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 toconduct these experiments in a uniform way they designedand built a stanciard piece of equipment, known as the Wis-consin General Test Apparatus (WGTA). This device wasacloptec! and used by many investigators over the years, even. .unto recent times.One of the most significant discoveries HarIow and hisassociates macle in their first primate laboratory dealt withthe formation of learning sets, that is, the process by whichanimals "learn to learn." Their procedure was to presentpairs of objects or patterns that cliffered in features such assize, color, and shape over a series of trials. The objectschanger! every few trials, and the animal graclually learner!to abstract particular features that clifferentiatecl the correctresponse object from others. In this way, discrimination cuesbecame generalizect and a learning set was establishect. Hariowand his students, as well as others, exploited this techniquein the study of brain lesions ant! other experimental vari-ables.The origin and concept of the learning set idea was notsullen. From 1939 to 1940, cluring a sabbatical year, Hariowheld a Carnegie fellowship at Columbia University with

HARRY FREDERICK HARLOW223famed anthropologist Franz Boas. While at Columbia he at-tendect a seminar by the German neurologist Kurt Goiclsteinand became familiar with his theories concerning abstractand concrete intelligence and learning, which relied heavilyupon 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 theprinciple of a category iclentifie(l. Accorcling to GoIclstein,only humans are capable of abstract thought. Hariow tenta-tively ctisagreed. Upon returning to Wisconsin he pursuedresearch that eventually demonstrated—contrary to GoIcI-stein's view that monkeys could also solve WeigI ancT Vigot-sky type problems, suggesting certain levels of abstractthought and reasoning. These results, together with thosefrom his earlier studies of odcl*ty and matching-to-samplediscriminations caused HarIow to focus on the question ofmethoclology.Limited by cost, upkeep, anct availability of monkeys, Har-low was forced to ignore the usual experimental proceduresof the time; that is, use of naive and different animals foreach condition or problem, as was the practice with cheapant! plentiful rodents. He used the same monkeys for thestudy of a variety of problems. If separate groups of monkeyshac! been used to learn single, simple discriminations, hemight not have discovered the concept of learning set. He fur-ther realized that subjecting monkeys to series of similar butrelated problems paralleled the situations in which childrenlearn.At a time when Thornclikian trial-and-error learning wasat 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 aproblem-by-problem exposition of the briciges between trial-

224BIOGRAPHICAL MEMOIRSand-error learning and insight. These results posed addi-tional difficulties for the conditioning theories of Clark Hulland Kenneth Spence, influential learning theorists at thattime. Sometimes bitter arguments ensued, but HarIow's re-sults and interpretations could not be denied. His Earning setresults were enthusiastically received when presented in hisPresidential Address before the Midwestern PsychologicalAssociation in 1948. The subsequent wide acceptance ofthese 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, thesestudies oriented the methods and thinking of modern cog-nitive psychologists toward natural as opposed to contrived. . .information processing.Another notable accomplishment involved investigationsof newly conceived and identified curiosity and manipulationGraves, in cooperation with Robert A. Butler, Donald R.Meyer, and Harry's wife, Margaret Kuenne HarIow, a childpsychologist. At a time when drives were considered to bewholly or partly physiological, HarIow and his associates es-tablished the fact that the curiosity and manipulation driveswere intrinsic parts of the rhesus monkey's motivationalstructure. Food, water, and sex were not found sufficient ornecessary to initiate behaviors resulting from curiosity andmanipulation drives. Monkeys were just naturally curiousand 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 monkeyslearned complicated tasks without being deprives! of basicnecessities such as food and water.Along with the foregoing studies of a strictly behavioraland psychological nature, which had such an important bear-

HARRY FREDERICK HARLOW225ing 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, andespecially the cerebral cortex, in conditioning, visual discrimi-nation, learning, and memory. The need for more refinedbehavioral 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-cl*tioned response could be establishect in the cat if, duringthe normal training procedure, the paw-lifting response tothe 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 wasthat everything, including the motor discharge blocked bythe 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 themuscle was free of paralysis, they found that no conditionedresponse could be elicited. Although this appeared to be alandmark cliscovery, there were obvious flaws in the hypoth-esis, for proprioceptive feedback was also eliminated by thelack of movement caused by the curare. Furthermore, theresult was subsequently shown by others to be inconclusivewhen it was found that curare tract a depressing effect on thecentral nervous system, as well as a paralyzing action at theneuromuscular junction. Hariow then abandoned this typeof research, but many years later he considered that decision

226BIOGRAPHICAL MEMOIRSto be a mistake. In hindsight he felt that he hac! been on theverge of an important discovery that was not unearthed untilyears later by other investigators.From about 1940 on, HarIow, his students, and associatesmade repeated attempts to determine the ejects of brainlesions 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 resultectin important contributions, but very little of major signifi-cance evolvecI, compared with the earlier ancI later areas ofinvestigation with which HarIow was associated. One set ofstudies 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 physiologistJohn F. Fulton at Yale in the 1930s, had stuclied the clelayedresponse performance of monkeys following prefrontal cor-tex ablations and found that the monkeys couIc! not seem todetermine 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 deficitin 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 andother discriminations that would not have been possible withsevere memory deficits. Instead, they attributed the variabil-ity and the sometimes poor performance to an inability toattend to the task ant! avoic! distractions. These results, how-ever, were anteciated by the publications of Malmo ~ ~ 942) andFinan (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 HARLOW227focused only upon the stimulus panels, thus avoiding dis-tractions. These findings were later confirmed by French andHarlow ~ ~ 9621. Thus, ~acobsen's putative memory loss resultscould 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 whetherinterpreted as attention or memory deficits, had importantimplications 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 intothe 1950s before being generally abandoned, despite somereported improvement in depressive and other psychopath-ological conditions. Earlier recognition of the disadvantagesof such operations as revealed by animal studies like Harlow'smight have forestalled the vast number of lobotomies per-formed.In 1932 Harlow moved into a two-story building that wasto be his laboratory for the next twenty years. This buildinghad less than the desirable amount of space in which to fit asmall 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 theanalysis and storage of research data. Furthermore, it was inthe early stages of the Depression and financial support wasin short supply everywhere. There were, of course, no fed-eral granting agencies at that time to support research andtraining fellowships for graduate students, as there would belater in the 1950s and beyond. These, however, were prob-lems faced by most college and university professors luckyenough to have a job.It is said, "Where there is a will, there is a way!" Harryhad a will, and he found a way. He was highly motivated andhad recently found a goal that would become a lifetime en-deavor: focus on the rhesus monkey (Macaca mulata) as an

228BIOGRAPHICAL MEMOIRSexperimental mode! for the study of the neural and behav-ioral aspects underlying human psychology. He soon foundthat not the least of his problems was the upkeep and survivalof his monkeys. Over the next twenty years he developer! theexperience ant] knowlecige necessary to sustain primates over-long periods of time within animal enclosures, though theyenjoyed only a few summer months of the warm weathertypical of their natural habitat. It was also in this laboratorythat Hariow supervises! his first Ph.D. student, AbrahamMasIow, who later developed the self-actualization theory ofmotivation and was creditect with being one of the foundersof the humanistic psychology movement.In 1953, the primate laboratory operations were movedfrom their initial location to a renovated cheese factory sev-eral city blocks from the campus. The motivational, learning,and neurophysiological-behavioral research was continuedant! expanclect, resulting in a need for more monkeys. For-tunately, the space was now acloquate. Because of importproblems, disease, and the cost of the monkeys, the decisionwas made to start a breeding colony of rhesus monkeys.There was virtually no information available on the care andrearing of laboratory-born monkeys. Methods were clevisedthrough trial, error, and observation to enhance the proba-bility that the newborns wouIc! survive.Initially, forty infant rhesus monkeys were separated fromtheir mothers and raiser! in separate cages. The result wasdisease-free animals that manifested bizarre ant! psychopath-ological behaviors. These abnormal behavior syndromeswere attributed to the effects of early isolation and lecl tosome of Harry HarIow's most fascinating and best-known re-search. The breeding, rearing, and nursery proceduresproved successful overall, and a subsequent published reportwith A. I. Blomquist served as a guide for breeders in otheranimal installations, including zoos. HarIow's infant pri-

HARRY FREDERICK HARLOW229mate care methods were eventually adopted in many placesarounct the world.The availability of laboratory-born infants led to the studyof 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 toeighteen months of age. They provided interesting data forcomparison with the age-level stages of intelligence and men-tal growth established by lean Piaget, the famous Swiss chilctpsychologist. 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 toreproduce 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 asort 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. Theconcept of a surrogate mother was not new; it was HarIow'sgenius in creating simple experimental situations in which touse the surrogate that was novel and important. With the aidof graduate student Robert Zimmerman, surrogates werebuilt 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 withterrycloth. Other maternal characteristics were adcled, suchas protruding rubber nipples for the supply of milk, internaltemperature 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

230BIOGRAPHICAL MEMOIRSto nourishment provided by feecting. Also, the warmth androcking were found to be important factors.Very important to HarIow was the fact that he hacl nowfound a way to raise disease-free monkey infants in isolation.Surrogate mothers now proviclec! warmth, comfort, andsustenance in an environment that couIcl be controlled ant!modified as required by experimental research programs.The tempo and scope of the infant monkey research nowincreased and many studies were undertaken, the results ofwhich often interested! psychoanalysts and challenged psy-choanalytic theory as well as tracl*tional learning theory.When social development of surrogate-mother-raised andbiological-mother-raised monkey infants was compared, itwas fount] that natural genetic mothers were significantlybetter at socializing the young. The importance of peer re-lationships was studiecl by raising infants together and awayfrom aclult animals, ant! it was found that the presence ofpeers and play opportunities was important to the process ofsocial development. Some of the tiara inclicated that the peer-to-peer interactions were more important than those betweenmother and infant. HarIow reporter! that one of the mostimportant relationships determining normal sexual behavioras aclults was the peer play cluring infancy and childhood ofthese monkeys.Laboratory research extended into the abnormal as wellas the normal behaviors. The bizarre behaviors resultingfrom isolation were user} as a basis for studying the long-termeffects of isolation per se. Animals separates! from their moth-ers at birth and isolated for periods of six months or longershowed deficits in social, sexual, and other behaviors. Thelonger the isolation, the worse the deficit. Impregnated, sex-ually mature female isolates showoct few of the normalmother responses expected of rhesus females. In some casesthese motherIess-mothers grossly abused their newborns, in-

HARRY FREDERICK HARLOW231dicating the importance of early socialization in the learningof proper caretaking and mothering behaviors."Gove created, love (destroyed ant! love regained" guiclectthe 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-moclelingtherapist) resulted in love regained.The reactions of isolate-raised infants separated fromtheir mothers were akin to those of human infants sufferingfrom anacl*tic depression as clescribed by British psychiatristI. A. Bow~by. When peer-rearecT monkeys were separated forseveral clays similar kinds of clepressed behaviors were notectby HarIow ancI S. Suomi, anti the pattern persisted.Love was regained by rehabilitation, mainly with youngermonkeys. A series of studies with infants isolated from birthfor various periods of time indicates! that placement withnormal, 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 normalmonkeys immediately upon release from isolation eventuallyled 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 researchon the activities of monkey nuclear families living in adjacentenclosures. The setting allowed for the study of infant inter-actions, fathering, ant! other relationships. HarIow pursuedthe research on monkey nuclear families and depression un-ti! his retirement from Wisconsin. He believed, however, that

232BIOGRAPHICAL MEMOIRShis most significant contributions came out of his surrogatestudies on love, isolation ejects, anct psychopathology. Har-low often said that when you work with monkeys, you thinkof 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 centraltheme. His main goal, it seems, was to study a single species,the rhesus monkey, to learn all he could about its behaviorand cognitive processes, and to relate the results to humans.The first decade of his tenure at the University of Wisconsinwas dedicated to finding suitable laboratory research facili-ties. Then he strove to find the best empirical methods forworking with monkeys and to develop unique tests for as-sessing their sensory and perceptual abilities and the moti-vational circ*mstances uncler which they worked best at solv-ing graded levels of problems anti tasks. This lect to thedevelopment of elaborate and creative test batteries and theWGTA, all of which benefittecl and stimulated his many stu-clents and others throughout the worIcI. He turned briefly tothe 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 totry to locate in each of the principal regions of the cerebralcortex 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 seemedto interest him as much as unclerstancTing behavior itself. Heconcentrated on behavior studies with monkeys throughoutthe last three decades of his life, opening up new vistas withregard to the cognitive aspects of behavior ant! the social ant!affective consequences of manipulation of the environmenton early development. Harry HarIow was a clecluc.tive, qual-

HARRY FREDERICK HARLOW233itative empiricist and phenomenologist, whose greatest clis-coveries and contributions resulted from planned serendip-ity. Serendipity even entered into his poetry, which he createdquickly and freely anc! often injected into his publicationsand talks.OTHER SCIENTIFIC AND PROFESSIONAL ACTIVITIESBy his own admission, HarIow was more of a nativist, whobeliever! in the inheritance of characteristics to a greater ex-tent than many of his contemporaries in psychology. Yet thispersonal inclination was not strongly reflected in his re-search, which in its cognitive and social aspects emphasizedenvironmental influences. He foresaw the importance of bio-chemistry in studies of behavior, ancT in 1958 he credited avolume on the Biological and Biochemical Bases of Behavior withneurophysiologist Clinton N. Woolsey, also of the Universityof Wisconsin and a member of the National Academy of Sci-ences. The book resulted from a symposium they hadplannecl 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 ofphenylketonuria in monkeys and humans.Although not directly involved in the space program, theWisconsin Laboratory supplied one of the first monkeys sentinto space. In 1954 Harry cooperates! with aerospace pioneerD. C. Simons on a series of stratospheric plastic balloonflights 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 sametime, he was involved in investigating the behavioral effectsof cortical implantations of radioactive cobalt.During a two-year leave from the University from ~950 to1952, HarIow served with the Department of the Army in

234BIOGRAPHICAL MEMOIRSthe Pentagon as chief of the Human Resources Branch. Inthat position, he was responsible for proposing the establish-ment of a Human Resources Research Office for the concluctof psychological research. The recommendation was imple-mented in 1951 and the office established on the campus ofThe George Washington University, with Meredith Crawfordas its director. With Phillip Sapir of the National Institute ofMental Health, he collaborated in the cofounding of theNIMH Small Grants Program.From ~ 95 ~ to ~ 963, HarIow served as editor of the Journalof Comparative and Physiological Psychology, using that positionto advance primatology. The proportion of publications (led-icated to primate behavior increased noticeably over thetwelve 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 thoseareas proved disappointing.In 1964, one of the seven national Regional Primate Re-search Centers was established adjacent to the University ofWisconsin Primate Laboratory. HarIow servect as its directoruntil 1971.Harry F. HarIow was electecl to the National Academy ofSciences in 195~ and to the American Academy of Arts andSciences 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 experimentson the behavior of monkeys, inclucling studies of motivation,learning, and problem solving." From 1958 to 1959, Hariowserved as president of the American Psychological Associa-tion; in 1960, he received its Distinguished Scientific Contri-bution Awarc! for "curiosity and imagination which openedup new areas of research in animal behavior and enhancedthe position of comparative psychology."President Lyndon B. Johnson presented him with the Na-

HARRY FREDERICK HARLOW235tional Medal of Science in 1967, and in 1973 he received theGold Medal Awarct of the American Psychological Asso-ciation. HarIow accepted the Kittay Scientific FoundationAward in 1975 for his use of monkey models to study psy-chopathological behaviors. The Primate Laboratory at theUniversity of Wisconsin was dedicated and named in hishonor in ~ 984.Harry Frederick Harlow was an unassuming man of manytalents. He was a poet and gifted writer, an excellent bridgeplayer, and a pretty goocl locally competitive tennis player.He was generous in sharing time anti ideas with students whowisher! to set up primate laboratories elsewhere. His longwalks with professional colleagues and graduate studentswere legendary, as were his many professional talks.Harlow married Clara Mears in 1932. Two sons, Robertand Richard, were born of this union. In 1946 the marriagewas dissolved ant} each party later remarried. In ~ 948, Harrymarried Margaret Kuenne, a child psychologist of note, withwhom 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 in1971. Shortly after her death, Harry and Clara, his first wife(then a willow) remarried. Upon his retirement from theUniversity of Wisconsin in 1974, Harry and Clara moved toTucson, Arizona, where Harry heal an honorary appoint-ment at the University of Arizona. He ant! Clara collaboratedon several publications, the most notable being a book en-titlecI, The Human Model: Primate Perspective, published in~979. Harry died in ~98~.

236BIOGRAPHICAL MEMOIRSSELECTED BIBLIOGRAPHY1932Social 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 tothe orangoutan. J. Comp. Psychol., 13 :313 - 43.With A. H. Maslow. Comparative behavior of primates: II. Delayedreaction 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. Delayedreaction tests on subnormal humans. {. Comp. Psychol.,14:253-62.With R. Stagner. Psychology of feelings and emotions: I. Theoryof feelings. Psychol. Rev., 39:570 - 89.Food preferences of the albino rat. I. Genet. Psychol., 41 :430-38.1933With 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 itsrelation to attitudes of ascendance and submission. J. Comp.Psychol.,16:171-86.With R. Stagner. Effect of complete striate muscle paralysis uponthe learning process. I. Exp. Psychol., 16:283-94.With R. Stagner. Psychology of feelings and emotions: II. Theoryof emotions. Psychol. Rev., 40: 184 -195 and 368 -80.1934With P. Settlage. Comparative behavior of primates: VII. Capacityof monkeys to solve patterned string tests. J. Comp. Psychol.,18:423-35.1936With P. Settlage. Concerning the sensory pathway in the condi-tioned reflex. J. Comp. Psychol., 22:279-82.

HARRY FREDERICK HARLOW237With P. Settlage. The effect of application of anesthetic agents oncirc*mscribed motor and sensory areas of the cortex. I. Psy-chol., 2: 193-200.The neurophysiological correlates of learning and intelligence.Psychol. Bull., 33 :479-534.1937Experimental analysis of the role of the original stimulus in con-ditioned responses in monkeys. Psychol. Rec., 1 :62-68.1938With I. Bromer. A test-apparatus for monkeys. Psychol. Rec.2:434-36.1939With P. Settlage. The effect of curarization of the fore part of thebody upon the retention of conditioned responses in cats. I.Comp. Psychol., 27:45-48.Recovery of pattern discrimination in monkeys following unilateraloccipital 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., 55:49-58.1940With F. Toltzien. Formation of pseudo-conditioned responses inthe cat. J. Genet. Psychol., 23:367-75.The effects of incomplete curare paralysis upon formation andelicitation of conditioned responses in cats. I. Genet. Psychol.,56:273-82.1942With J. Bromer. Acquisition of new responses during inactivationof 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.

238BIOGRAPHICAL MEMOIRSAnimal behavior. In: Fields of Psychology, ed. R. H. Seashore, pp.171-96. New York: Holt, Rinehart & Winston.1943With 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 probleminvolving 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.O1J- -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 bilateralremoval 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 bilateralremoval of the prefrontal areas: II. Delayed reaction problemsinvolving use of the matching-from-sample method. }. Exp.Psychol., 32:424-34.With T. Johnson. Problem solution by monkeys following bilateralremoval 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 bilateralremoval of the prefrontal areas: IV. Responses to stimuli havingmultiple sign values. J. Exp. Psychol., 33:500-7.1944Studies in discrimination learning by monkeys: I. The learning ofdiscrimination series and the reversal of discrimination series.I. Genet. Psychol., 30:3-12.Studies in discrimination learning by monkeys: II. Discriminationlearning without primary reinforcement. I. Genet. Psychol.,30:13-21.With M. M. Simpson. Solution by rhesus monkeys of a nonspatialdelayed response to the color or form attribute of a single stim-

HARRY FREDERICK HARLOW239ulus (Weigl principle delayed reaction). J. Comp. Psychol.,37:211-20.1945With 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 monkeysto 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 byrhesus monkeys. l. Genet. Psychol., 32:213-27.Studies in discrimination learning by monkeys: IV. Relative diffi-culty of discrimination between stimulus-objects and betweencomparable patterns with hom*ogeneous 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 andpattern discriminations. i. Genet. Psychol., 33:3-10.Studies in discrimination learning by monkeys: VI. Discriminationbetween stimuli differing in both color and form, only in color,and only in form. I. Genet. Psychol., 33:225-35.1946With 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 ofobject-quality and positional discriminations and discriminationreversals. J. Comp. Psychol., 39:13-23.1947With 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 inmonkeys. J. Comp. Physiol. Psychol., 40:333-42.With P. H. Settlage. An effective and nontraumatic method of han-dling monkeys. Science, 106:300.

240BIOGRAPHICAL MEMOIRS1948With G. Andrew. Performance of macaque monkeys on a test ofthe concept of generalized triangularity. Comp. Psychol.Monogr., 19: No. 1, Serial No. 100.With L. Grandine. Generalization of the characteristics of a singlelearned 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 extensiveunilateral 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 toidentical stimuli. J. Exp. Psychol., 38:50-65.1949Physiological 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 topatterning by monkeys. I. Comp. Physiol. Psychol., 42:454-62.1950With R. T. Davis and P. H. Settlage. Performance of normal andbrain-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 discriminationreversal 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 HARLOW1951241With D. R. Meyer and P. H. Settlage. A survey of delayed responseperformance by normal and brain-damaged monkeys. I. Comp.Physiol. Psychol., 44: 17-25.With A. I. Riopelle, P. H. Settlage, and H. W. Ades. Performanceof 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 corticallesions 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 responsesand 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.1952With 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 brainoperated monkeys. T. Genet. Psychol., 81:45-52.With D. R. Meyer. Effects of multiple variables on delayed responseperformance 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 corticallesions. i. Comp. Physiol. Psychol., 45:119-26.Functional organization of the brain in relation to mentation and

242BIOGRAPHICAL MEMOIRSbehavior. 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., 45:576-84.1953Mice, 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., 46:13-15.1954Motivational 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 monkeyson 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.,47:391-94.1955With 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 HARLOW243With 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 implantationof radioactive cobalt on learned behavior of rhesus monkeys. I.Comp. Physiol. Psychol., 48:432-36.With G. M. French. Locomotor reaction decrement in normal andbrain-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: 1-2.1956With 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 rhesusmonkeys. 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 discriminationlearning by monkeys. J. Comp. Physiol. Psychol., 49: 117-22.With A. M. Schrier and D. G. Simons. Exposure of primates tocosmic radiation above 90,000 feet. I. Comp. Physiol. Psychol.,49: 195-200.With N. C. Blazek and G. E. McClean. Manipulatory motivation inthe infant rhesus monkey. I. Comp. Physiol. Psychol., 49:444-48.With W. A. Mason and N. C. Blazek. Learning capacities of theinfant rhesus monkey. J. Comp. Physiol. Psychol., 49:449-53.Current and future advances in physiological psychology. Am. Psy-chol., 11:273-77.1957With L. H. Hicks, Jr. Discrimination learning theory: Uniprocessvs. duoprocess. Psychol. Rev., 64:104-9.Experimental analysis of behavior. Am. Psychol., 12:485-90.

244BIOGRAPHICAL MEMOIRSWith 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, andDDD (debility, dependency, and dread). Sociometry, 20:271-85.With I. M. Warren, R. W. Leary, and G. M. French. Function of theassociation cortex in monkeys. Brit. I. An. Beh., 4: 132-38.With A. M. Schrier. Direct manipulation of the relevant cue anddifficulty of discrimination. J. Comp. Physiol. Psychol., 50:576-80.1958With W. A. Mason. Learned approach by infant rhesus monkeys tothe sucking situation. Psychol. Rep., 4:79-82.With W. A. Mason. Formation of conditioned responses in infantmonkeys. J. Comp. Physiol. Psychol., 51:68-70.With W. A. Mason. Performance of infant rhesus monkeys on aspatial discrimination problem. l. Comp. Physiol. Psychol., 51:71-74.With A. M. Schrier. Effect of reserpine on avoidance of humans byrhesus monkeys. J. Genet. Psychol., 59:149-55.The evolution of learning. In: Behavior and Evolution, ed. A. Roeand G. Simpson, pp. 269-90. New Haven: Yale UniversityPress.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 tointerdisciplinary research. In: Biological and Biochemical Bases ofBehavior, pp. 3-23. Madison: University of Wisconsin Press.The nature of love. Am. Psychol., 13:763 -85.1959Learning set and error factor theory. In: Psychology: A Study of aScience, ed. S. Koch, pp. 492-538. New York: McGraw-HillBook Co.With W. A. Mason. Initial responses of infant rhesus monkeys tosolid 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 variablein the acquisition of discrimination learning set. I. Comp. Phys-iol. Psychol., 52 :396-98.

HARRY FREDERICK HARLOW245With 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 oddityproblems. 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., 52:555-58.1960With M. K. Harlow, R. R. Rueping, and W. A. Mason. Performanceof infant rhesus monkeys on discrimination learning, delayedresponse, and discrimination learning set. l. Comp. Physiol.Psychol., 53: 113 -21.Affectional behavior in the infant monkey. In: The Central NervousSystem 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. Phenylketonuriain infant monkeys. Nature, 188: 1124-25.With K. Akert, O. S. Orth, and K. A. Schiltz. Learned behavior ofrhesus monkeys following neonatal bilateral prefrontal lobot-omy. Science, 132: 1944-45.1961With A. I. Blomquist. The infant rhesus monkey program at theUniversity of Wisconsin Primate Laboratory. Proc. An. CarePanel, 11:57-64.With L. R. Cooper. Note on a rebus monkey's use of a stick as aweapon. 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:

246BIOGRAPHICAL MEMOIRSDeterminants 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 discriminationlearning. J. Genet. Psychol., 98:183-86.With W. A. Mason. The effects of age and previous training onpatterned-strings performance of rhesus monkeys. I. Comp.Physiol. Psychol., 54:704-9.1962Effects of radiation on the central nervous system and on behav-ior—general survey. In: Response of the Nervous System to IonizingRadiation, ed. T. i. Haley and R. S. Snider, pp. 627-44. NewYork: 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 fromAnimal 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 innormal and brain-damaged rhesus monkeys. l. Neurophysiol.,25: 585-99.With i. M. Lockhart. The influence of spatial configuration andpercentage 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 HARLOW247With I. E. King. Effect of ratio of trial 1 reward to nonreward onthe 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 annualresearch conference. Michigan Department of Mental Health.1963With H. A. Cross and H. I. Fletcher. Ejects of prior experiencewith 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 Emotionin Man, ed. P. H. Knapp, pp. 254-65. New York: InternationalUniversities Press.With M. K. Harlow and E. W. Hansen. The maternal affectionalsystem 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.1964With K. Akert and K. A. Schiltz. The effects of bilateral prefrontallesions 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

248BIOGRAPHICAL MEMOIRSdeprived rhesus monkeys. J. Abnorm. Soc. Psychol., 69:345-54.With G. L. Rowland and G. A. Griffin. The effect of total socialdeprivation on the development of monkey behavior. In: RecentResearch on Schizophrenia, Psychiatric Research Report 19, ed. P.Solomon and B. C. Glueck, pp. 116-35. Washington, D.C.:American Psychiatric Association.1965With H. A. Waisman. Experimental phenylketonuria in infantmonkeys. 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 inmonkeys. 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, andF. Stollnitz, pp. 287-334. New York: Academic Press.With B. K. Alexander. Social behavior of juvenile rhesus monkeyssubjected to different rearing conditions during the first sixmonths of life. Zool. J. Physiol., 71:489-508.With R. L. Raisler. Learned behavior following lesions of posteriorassociation 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. Oslerand R. E. Cooke, pp. 87-106. Baltimore: The Johns HopkinsPress.Total social isolation: Effects on macaque behavior. Science,148:666.

HARRY FREDERICK HARLOW1966249With M. K. Harlow. Affection in primates. Discovery, 27:11-17.With M. K. Harlow, R. O. Dodsworth and G. L. Arling. Maternalbehavior of rhesus monkeys deprived of mothering and peerassociation 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 rhesusmonkeys. Psychol. Rep., 18:567-80.The primate socialization motives. Trans. Stud. Coll. PhysiciansPhiladelphia, 33:224-37.With E. W. Hansen and R. O. Dodsworth. Reactions of rhesusmonkeys 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.sof reproduction in primates. J. Anim. Sci., 25:49-65.- or ~rigWith 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 motherrearing. Child Dev., 37:781-91.With B. Seay. Mothering in motherless mother monkeys. Br. I. Soc.Psychiatry, 1:63-69.1967With 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. ChildDev., 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.

250BIOGRAPHICAL MEMOIRS1968With A. I. Blomquist, C. I. Thompson, K. A. Schiltz, and M. K.Harlow. Effects of induction age and size of frontal lobe lesionson learning in rhesus monkeys. In: The Neuropsychology of De-velopment: A Symposium, ed. R. L. Isaacson, pp. 79-120. NewYork: 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 DellaRicerca Sull Apprendimento E La Memory, ed. D. Bovet, F. Bovet-Nitti, and S. Oliverio, pp. 139-56. Rome: Accademia Nazionaledej Lincei.With G. W. .M011er and G. D. Mitchell. Factors affecting agonisticcommunication in rhesus monkeys (Macaca mulatta). Behav-iour, 31:339-57.A primate. Science, 165:274.1969With M. K. Harlow. Effects of various mother-infant relationshipson 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: UnfinishedBusiness, ed. R. B. McCleod, pp. 21-30. Washington, D.C.:American Psychological Association.Age-mate or peer affectional system. In: Advances in the Study ofBehavior, 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 HARLOW251With 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 surfacesamong infant rhesus monkeys. Bull. Psychon. Sci., 17:279-80.With K. A. Schiltz and M. K. Harlow. Effects of social isolation onthe learning performance of rhesus monkeys. In: Proceedings ofthe 2nd International Congress of Pr~matology, ed. C. R. Carpenter,vol. 1, pp. 178-85. Basel/New York: Karger.1970With 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 ofdepression in monkeys. Mainly Monkeys, 1:6-12.With A. C. Deets. Nipple preferences in nursing singleton- andtwin-reared rhesus monkey infants. Dev. Psychol., 2: 159-62.With C. I. Thompson, A. J. Blomquist, and K. A. Schiltz. Learningin rhesus monkeys after varying amounts of prefrontal lobedestruction during infancy and adolescence. Brain Res., 18:343-53.With S. I. Suomi. Induction and treatment of psychiatric states inmonkeys. 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 ofcombined frontal and temporal lesions on learned behaviors inrhesus monkeys. Proc. Natl. Acad. Sci. USA., 66:577-82.With A. S. Chamove and H. A. Waisman. Abnormal social behaviorin phenylketonuric monkeys. J. Abnorm. Psychol., 76:62-68.With A. C. Deets. S. D. Singh, and A. I. Blomquist. Effects of bilat-

252BIOGRAPHICAL MEMOIRSeral lesions of the frontal granular cortex on the social behaviorof 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-infantseparation of young monkeys. I. Abnorm. Psychol., 76: 161-72.With S. I. Suomi and G. P. Sackett. Development of sex preferencein rhesus monkeys. Dev. Psychol., 3:326-36.With A. C. Deets and A. I. Blomquist. Effects of intertrial intervaland trial 1 reward during acquisition of an object discriminationlearning set in monkeys. I. Comp. Physiol. Psychol., 73:501-5.1971With 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-responseproblems. 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 sexualposturing in rhesus monkeys. Arch. Sex. Behav., 1:73-78.Early problem learning and early social learning. In: The SecondWestern 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 prolongedpartial 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 HARLOW253early 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 youngmonkeys. I. Autism Child. Schizophren., 1 :246-55.With M. K. Harlow. Psychopathology in monkeys. In: ExperimentalPsychopathology, ed. H. D. Kimmel, pp. 203-29. New York: Aca-demic Press.With C. I. Thompson, A. J. Blomquist, and K. A. Schiltz. Recoveryof 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 rhesusmonkeys. Dis. Nerv. Sys., 32:735-41.With W. T. McKinney, Jr. Nonhuman primates and psychoses. J.Autism Child. Schizophren., 1 :368-75.1972With 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 AnamauxDu Comportement Humain, No. 198, pp. 13 - 60. Paris: Editionsdu Centre National de la Recherche Scientifique.With S. J. Suomi. Depressive behavior in young monkeys subjectedto vertical chamber confinement. I. Comp. Physiol. Psychol.,180:11-18.With W. T. McKinney and S. J. Suomi. Repetitive peer separationsof juvenile-age rhesus monkeys. Arch. Gen. Psychiatry,27:200-4.With I. P. Gluck and S. i. Suomi. Generalization of behavioral databetween nonhuman and human animals. Am. Psychol., 27:709-16.With M. K. Harlow. The language of love. In: Communication andA§ect, ed. T. Alloway, L. Krames, and P. Pliner, pp. 1-18. NewYork: Academic Press.With I. B. Sidowski and S. I. Suomi. Enhancing social attachment.

254BIOGRAPHICAL MEMOIRSthrough 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., 24:496-504.1973With W. T. McKinney, tr., and S. i. Suomi. Methods and models inprimate 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 andfirst-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 stressand later response to separation in rhesus monkeys. Am. I. Psy-chiatry, 130:400-5.With D. M. Baysinger and P. E. Plubell. A variable-temperaturesurrogate-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 frontaland 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 andDepression, 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 HARLOW255With 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 psychologicaldeath in rhesus monkeys. }. Autism Child. Schizophr., 3:299-307.1974Induction 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 ina 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., 3:527-34.1975With 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 onsocial 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-

256BIOGRAPHICAL MEMOIRSlus 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 firstyear of life: 1. Rehabilitation and therapy. Dev. Psychol.,1 1 :453-65.With I. S. Meyer, M. A. Novak, and R. E. Bowman. Behavioral andhormonal 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 andL. 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: TheInside Story, ed. M. H. Siegel and H. P. Zeigler, pp. 3-22. NewYork: Harper & Row.1976With S. I. Suomi, M. L. Collins, and G. C. Ruppenthal. Effects ofmaternal 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 andAnxiety: New Concepts, Methods, and Applications, ed. M. Zucker-man and C. D. Spielberger, pp. 3-34. Hillsdale, N.J.: LawrenceErlbaum Associates.

HARRY FREDERICK HARLOW257With S. I. Suomi and R. DeLizio. Social rehabilitation of separa-tion-induced depressive disorders in monkeys. Am. J. Psychia-try, 133:1279-85.1977With 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.1978With 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.1979With C. Mears. The Human Model: Primate Perspectives. Washington,D.C.: V. H. Winston & Sons (Halsted Press).

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