The Counseling Psychologist

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CHAPTER 3 Biological Psychology  CHAPTER 4 Comparative Psychology  30 Psychology as a Profession such as Atlantic Monthly, Business World, Advertising World, and The Woman’s Herald, thus making business psychology known to a broad audience of potential employers and con- sumers. Scott promoted the psychology of suggestion, argu- ing that successful advertising suggested a course of action, that is, buying the product. He wrote, “Man has been called the reasoning animal but he could with greater truthfulness be called the creature of suggestion. He is reasonable, but he is to a greater extent suggestible” (Scott, 1903, p. 59). In applying suggestion to advertising, Scott advocated two tech- niques: the direct command (e.g., “Use Peterson’s Tooth Powder”) and the return coupon. Both techniques were thought to stimulate compulsive obedience. In the subsequent theoretical debates in the advertising community on the nature of consumer behavior, other approaches displaced Scott’s views (see Kuna, 1976, 1979), but his work gave psychology considerable visibility in the world of business and paved the way for many psychologists who would follow in advertising such as Harry Hollingworth, Daniel Starch, and John B. Watson. Although business psychology can be said to have begun in the ﬁeld of advertising, it quickly branched into other prominent areas. When increased emphasis on efﬁciency led to the “scientiﬁc management” of Frederick Winslow Taylor (1911), psychologists entered that arena as well. Efﬁciency meant not only better management and more effective adver- tising but also better training of workers, improved employee selection procedures, better ways to control employee per- formance, and better understanding of human actions in work. Prominent in these areas was Harvard psychologist Hugo Münsterberg (1863–1916), who argued in his book, Psychology and Industrial Efﬁciency (1913), that the key to workplace efﬁciency was matching job and worker and that successful matches generated satisﬁed employees, quality work, and high productivity. Münsterberg promoted psy- chology as the science of human efﬁciency, noting that psychology had the tools to create the perfect match by de- termining the mental traits required for any job and the men- tal traits of workers. That his ideas were well received by a broad public is evidenced by the fact that his book was for a time on the national list of best-sellers. Psychologists began to develop mental tests to evaluate workers and jobs (ship captains, trolley car operators, sales- women), work that was to prove especially important when they were asked to oversee the selection program for the United States armed forces during World War I. Business psychology had begun in the universities, but its practice soon moved to business settings as psychologists found full- time employment, particularly as personnel ofﬁcers involved with selection, job analysis, and training. Such opportunities expanded considerably after World War I, establishing the psychologist as a key player in the world of business. The Counseling Psychologist As noted earlier, with the proliferation of types of jobs around the turn of the twentieth century, people had more oc- cupational choices than ever before. Vocational counseling, which had been a part of the business of nineteenth-century phrenologists, became even more important. The most inﬂu- ential ﬁgure in the vocational guidance movement of the early twentieth century was not a psychologist but an indi- vidual trained in engineering and law, Frank Parsons (1854–1908). He wrote his most important work in the wan- ing days of his life, a book published after he died, entitled Choosing a Vocation (1909). Parsons’s formula for success- ful guidance involved: (a) a clear understanding of the indi- vidual’s talents, limitations, and interests, (b) knowledge about diverse jobs including what was required for success in those jobs, and © matching those two kinds of information for the best vocational guidance. There were clear ties between Parsons’s approach and the matching between jobs and people that was the focus of psy- chologists in personnel work in businesses. Parsons, as part of the progressive movement of the times, emphasized the reduction of human inefﬁciency—as reﬂected in the high turnover of workers—through the application of a careful program of career planning. Vocational guidance became a mantra of progressive reformers and soon found its way into the American mainstream with the formation of the National Vocational Guidance Association in 1913. Quickly, the vocational guidance counselor was integrated into elementary and secondary schools across America, beginning a strong association between guidance and educa- tion. It also made its way into industry through personnel selection. Psychologists found the issues of person and career matching amenable to the new applied science of psychology and worked to develop reliable and valid measures of indi- vidual traits and abilities for use in guidance and selection. Guidance counseling became even more prominent in schools after the passage of the National Vocational Educa- tion Act in 1917. Following the First World War, vocational guidance centers (or “clinics,” as they were sometimes called) were established as well at colleges and universities. For example, Witmer founded a separate vocational guidance clinic at the University of Pennsylvania in 1920 that was headed by one of his doctoral graduates, Morris Viteles (1898–1996), who would later distinguish himself as an industrial psychologist. The Beginnings of the New Profession of Psychology 31 In all of these vocational guidance centers and clinics, the key component of the arsenal of the guidance specialists was mental tests, including interest tests that were developed in the 1920s, and a growing number of aptitude and ability tests that were used not only in guidance but also for selec- tion. This vocational role, both in personnel work and in guidance, remained relatively stable until after the Second World War. (See the chapters by Koppes and Baker in this volume.) The School Psychologist We have already noted that the origins of school psychology lie in the psychological clinic of Lightner Witmer. Thomas Fagan (1992) has written that: School psychology was one of many child-saving services orig- inating in the period of 1890 to 1920 …. t originated in response to compulsory schooling, which provided the stage for development of separate special educational programs for atypi- cal children. School psychology emerged in the middle of the child study movement. (p. 241) The child study work of Hall focused attention on a broad spectrum of child behavior and education. Many of Hall’s master’s and doctoral students at Clark University worked in what could be described as school psychology, includ- ing three particularly inﬂuential pioneers: Henry Herbert Goddard (1866–1957), Lewis Terman (1877–1956), and Arnold Gesell (1880–1961). Goddard was employed at the New Jersey Training School for Feebleminded Girls and Boys in Vineland when he began his research on mental retardation, searching for better tools for intellectual assessment and for methods of effective education and training of mentally handicapped children. Goddard was frustrated in his work at Vineland using the measurement tools he had learned at Clark University and from Cattell’s work. Whereas those tools seemed appropriate for assessment of children of normal intelligence, they were not useful for the children at Vineland. In a 1908 trip to Europe, Goddard learned of a new approach to intelligence testing developed by French psychologist Alfred Binet (1857–1911). Goddard translated the test for English- language use, tested it on samples of public school children as well as the students at the Vineland Training School, and published his version of the test in 1909. Its popularity as an instrument of intellectual assessment spread rapidly, culmi- nating in the version published by Terman in 1916 that be- came known as the Stanford-Binet Intelligence Test. Goddard’s role in school psychology, and more broadly in educational reform, cannot be overstated. He sought to apply the science of psychology to the questions then facing public schools, particularly regarding the educability of children labeled subnormal in intelligence. Through his research efforts, his training workshops for teachers, and the promi- nence of his ideas in American education, Goddard was instrumental in promoting special education opportunities in American schools (even though many of those efforts went beyond what he would have endorsed). More important for psychology, he established a place for psychologists in the schools as diagnosticians of mental capacity, a role that was often synonymous with the label of school psychologist in the twentieth century (Zenderland, 1998). Terman, like Goddard, also focused on intellectual assess- ment.Although Terman conducted some research on mentally handicapped children (including some work published with Goddard using subjects at Vineland), his work with children came to be more focused on gifted students, and he is arguably best known (beyond the Stanford-Binet) for the longitudinal studies of children identiﬁed as gifted, the “genius studies,” that began in California in 1921. His revision of the Binet test was better psychometrically than Goddard’s across all intel- lectual levels but especially so in the higher ranges. Terman, like Goddard, enhanced the role of psychologist as assessor of intellectual functioning and as designer of curricula for special-needs children, particularly gifted children. Gesell was the ﬁrst person in the United States to hold the title of “school psychologist,” according to Fagan (1992). He was hired by the Connecticut State Board of Education in 1915 to evaluate schoolchildren and make recommendations for those who needed special treatment. Gesell’s duties in the beginning of his work were research oriented, but he later came to be consumed by a caseload of 502 schoolchildren (and his duties were similar to those of contemporary school psychologists). The signiﬁcance of Gesell’s appointment was that the title “school psychologist” was associated “with services to exceptional children, especially the mentally deﬁ- cient, and it associated the functions of that title as primarily diagnostic testing for placement decisions in the newly created programs for the handicapped” (Fagan, 1987, p. 406). Although Gesell is perhaps the most prominent of the early school psychologists, he was not the only person performing those duties by 1915. Already schools were employing teach- ers in intellectual assessment roles as well as curriculum design for special children. Norma Estelle Cutts (1892–1988) played such a role as early as 1914 in the New Haven, Connecticut, schools after working with Goddard for a year at Vineland (Fagan, 1989). She was one of many individuals whom Goddard inﬂuenced to become school psychologists, 32 Psychology as a Profession most of them women who already had teaching experience. (See the chapter by Fagan in this volume.) The Clinical Psychologist At the beginning of the twentieth century, psychopathology was the domain of psychiatry and, to a lesser extent, neurol- ogy. Psychiatry, arguably the oldest of the medical specialties (excluding surgery), originated with the superintendents of mental asylums at the end of the eighteenth century. After a half century of asylum management, the superintendents formed an organization entitled the Association of Medical Superintendents of American Institutions for the Insane in 1844 and in the same year began publication of their journal, The American Journal of Insanity. The organization’s name was later changed to the American Medico-Psychological Association in 1892 and in 1921 to the American Psychiatric Association; the journal name was changed as well in 1921 to the American Journal of Psychiatry (Grob, 1994). The abnormal mind was of interest to some, perhaps many, of the early psychologists, but the domains of diagnosis and treat- ment seemed clearly within the boundaries of medicine, and few psychologists saw any need to venture there. That would soon change. Origins of any ﬁeld are rarely, if ever, unequivocal—and so it is with clinical psychology. We have already discussed the contributions of Lightner Witmer with respect to school and clinical psychology. Not only did he establish the ﬁrst psychology clinic in 1896, but as early as 1897 he had de- scribed a training program for psychologists to work in a ﬁeld that he had named “clinical psychology,” a ﬁeld that would draw from the knowledge base in medicine, education, and psychology (particularly child psychology). An expanded description of this ﬁeld and a rationale for its further devel- opment appeared in the inaugural issue of his journal, The Psychological Clinic (Witmer, 1907), a journal that largely published reports of the cases seen in Witmer’s clinic. Witmer was clearly interested in the difﬁculties that chil- dren exhibited in the classroom and believed that psychologi- cal science could offer solutions to behavioral problems of perception, learning, motivation, and emotion. He champi- oned the need for accurate diagnosis based on psychological and medical tests (the latter were performed by associated physicians). Slowly others began to share his vision, and, by 1914, there were psychology clinics at 19 universities. Witmer’s focus was on children (and chieﬂy on problems that impeded learning). Others soon broadened the scope of clini- cal psychology. But, the duties of these early clinical psychol- ogists remained focused on diagnosis and recommendations for treatment, with limited roles in actual treatment until after World War II. Psychotherapy, a book published in 1909 by Hugo Münsterberg, represents an early psychology-based contri- bution to the clinical intervention literature. It was a non- Freudian textbook grounded in a theory of psychophysical parallelism, which argued that all psychical processes had a parallel brain process. His volume argued for the scientiﬁc study of the processes of psychotherapy and viewed psy- chotherapy as a clinical endeavor separate from “psychiatry.” Other inﬂuences came from physicians cognizant of the potential contributions of psychology. Morton Prince (1854–1929) was a neurologist interested in the problems of psychopathology and one who recognized the importance of psychology in the study and treatment of psychological dis- orders. His most famous book, The Dissociation of a Person- ality (1908), was a lengthy and insightful description of a case of multiple personality. His contributions to clinical psy- chology were considerable and include his founding of the Journal of Abnormal Psychology in 1906, which published the early work on experimental psychopathology, and his establishment of the Psychological Clinic at Harvard Univer- sity in 1926, which he housed in the Department of Philoso- phy (where psychology was located) rather than in Harvard’s medical school. Another physician, William Healy (1869–1963), headed the Juvenile Psychopathic Institute, which opened in Chicago in 1909. Healy had studied with William James and had also been inﬂuenced by the work of Goddard at Vineland. His institute was to be both a research facility, investigating the causes of juvenile delinquency, and a treatment facility. He hired psychologist Grace Fernald (1879–1950) to work with him, and when she left, he replaced her with another psychol- ogist, Augusta Bronner (1881–1966), whom he would later marry. Both Fernald and Bronner used the title “clinical psy- chologist” and played important roles in research, diagnosis, and treatment. Other juvenile courts and corrections facilities began to hire psychologists for similar roles (Levine & Levine, 1992). Other stimulants to the development of clinical psychol- ogy before World War I included the work on mental assess- ment by Goddard and other advances in mental testing; the ﬁve addresses given by Sigmund Freud (1856–1939) at Clark University in 1909 that fostered considerable interest in psy- choanalysis in America but more broadly in the nature of causation in mental illness; the mental hygiene movement begun around 1908 by former mental patient Clifford Beers (1876–1943) and psychiatrist Adolf Meyer (1866–1950), a movement that sought to understand the early causes of men- tal illness and how conditions might be changed (in families World War I and the Growth of Psychological Practice 33 and society) to minimize psychological problems; and the popularity of the Emmanuel Movement begun by a Wundt doctoral student, Elwood Worcester (1862–1940), in his Boston church, a movement that spread across the United States emphasizing the alliance of medicine and psychology in treating mental disorders, a movement credited with the emergence of psychotherapy in America (Caplan, 1998). All of these forces brought psychology into greater contact with issues of mental pathology and afforded new jobs for psychologists, largely as mental testers. As the demand for these diagnostic services grew, clinical psychologists peti- tioned the APA in 1915 for a certiﬁcation program for quali- ﬁed psychologists in consulting roles, a measure that was seen to protect the public and to preserve the jobs of consulting psychologists. When the APA declined to provide such certi- ﬁcation, several psychologists, including J. E. Wallace Wallin (1876–1969) and Leta S. Hollingworth (1886–1939), formed in 1917 a new, short-lived organization entitled the American Association of Clinical Psychologists (AACP), arguably the ﬁrst association of professional psychologists. The member- ship totaled only about 45 psychologists in its ﬁrst year, some in university settings, some in applied jobs. The association was a clear statement of another of psychology’s applied specialists coming of age: the clinical psychologist. (See the chapter by Routh and Reisman in this volume.) WORLD WAR I AND THE GROWTH OF PSYCHOLOGICAL PRACTICE The foundations for the modern practice of psychology were well in place before the beginning of the First World War. Psychologists could be found working in schools, businesses, hospitals, and social and clinical service agencies. The num- ber of such individuals was still relatively small, particularly in comparison to their colleagues in colleges and universities. Two world wars would dramatically reverse that ratio. The ﬁrst would promote the rapid development of the practice specialties; the second would open the ﬂoodgates for psy- chological practice, including psychologists as independent practitioners of psychotherapy. It can be argued that American psychologists were un- prepared for World War I. On April 6, 1917, two days after America’s entry into the war, much of the leadership ofAmer- ican psychology—at least those located on the East Coast— were attending the annual meeting of E. B. Titchener’s “experimentalists” at Harvard University (see Boring, 1938, 1967). In attendance was Robert M. Yerkes (1876–1956), who was the current president of the APA. Yerkes chaired a discussion about psychology’s role in the war that led to an emergency meeting of the APA Council called for the end of April. At that meeting, Yerkes established a dozen com- mittees that were charged with pursuing various roles for psychologists within the war effort. Only two of those really materialized. One involved a testing program of nearly two million military recruits, headed by Yerkes, that developed group intelligence tests, namely the Army Alpha and Army Beta. The second program was headed by Walter Dill Scott, who used his experience in developing job selection tests to assess the job skills of more than three million military per- sonnel, a task accomplished by his staff’s development of more than 100 separate selection instruments in a little more than 12 months. After the war, Scott was awarded the Distin- guished Service Medal by the U.S. Army for this monumen- tally successful program. He was the only psychologist to be so honored in World War I (Napoli, 1981). The exact number of American psychologists who partic- ipated in the war is not known, but the ﬁgure is likely be- tween 250 and 300, counting those who served as consultants as well as those in uniform. Toward the end of the war, some were stationed at the 40 U.S. Army hospitals, where their assignments brought them into direct contact with issues of psychopathology. One example was Harry Hollingworth (1880–1956), a faculty member on leave from Barnard College who, as a captain in the army, was working at the army hospital in Plattsburgh, New York, examining approxi- mately 1,200 soldiers suffering from “shell shock” and other psychological disorders. Based on those experiences, Hollingworth wrote a book entitled The Psychology of Func- tional Neuroses (1920). Although Hollingworth was not led into clinical psychology by his wartime experiences, other psychologists were. All of the activities of psychologists during the war are far beyond the scope of this chapter. What is important to em- phasize, though, is that the war efforts by psychologists had important implications for the public and for the discipline of psychology. The work of psychologists, especially in selec- tion, was seen by the government and the public as a program of considerable success. Such favorable press brought many consulting opportunities to psychologists after the war, and psychologists were quick to take advantage of such applied opportunities. For example, Scott founded The Scott Com- pany, a consulting ﬁrm of psychologists based in Pittsburgh, to do contract work for businesses and government agencies. Further, the war work convinced psychologists of the value of their science, that is, that they had something signif- icant to offer in the public sector that was grounded in fact, not myth. This newly gained disciplinary awareness for psy- chologists, the public’s perception of the value of psychology as demonstrated by success in the war work, the growing 34 Psychology as a Profession economic prosperity of America in the 1920s, and the rapid social changes in American society after the war were all fac- tors that led to the further development of the profession of psychology. THE 1920s: THE DECADE OF POPULAR PSYCHOLOGY American historians have written of the public euphoria in the United States that followed World War I. American forces had helped to win the war in Europe. There was general economic prosperity, and a growing belief in the American dream that anything was possible, with hard work. Writing for the American public in 1925, psychologist John B. Watson (1878–1958) promoted this nurturistic optimism: Give me a dozen healthy infants, well-formed, and my own speciﬁed world to bring them up in and I’ll guarantee to take any one at random and train him to become any type of specialist I might select—doctor, lawyer, artist, merchant-chief and, yes, even beggar-man and thief, regardless of his talents, penchants, tendencies, abilities, vocations, and race of his ancestors. (Watson, 1925, p. 82) Americans seemed delirious with the potential for psychol- ogy to improve their lives. The ﬁrst popular psychology magazines (four of them) began publication in the decade. Countless self-help books were published, and newspapers carried daily columns of psychological advice. Touting the value of psychology for the public, journalist Albert Wiggam (1928) wrote: Men and women never needed psychology so much as they need it to-day. . . . You cannot achieve these things [effectiveness and happiness] in the fullest measure without the new knowledge of your own mind and personality that the psychologists have given us. (p. 13) Public demand for psychological services grew rapidly, and consequently, many individuals, with little or no training in psychology, offered their services to the public as psychologists. Consulting psychologists were especially concerned about such pseudopractitioners and petitioned the APA to create a certiﬁcation program to identify psychologists qualiﬁed to consult with the public. Initially, the APA balked at the idea but relented in 1924, when it established such a program. Four years later, after fewer than 30 psychologists had received certiﬁcation, the program was abandoned (Sokal, 1982b). There was no mechanism for enforcement of such a program, and the public seemed incapable of making distinctions between qualiﬁed psychologists and unqualiﬁed ones, or at least was uninterested in doing so. Nevertheless, psychology of all kinds prospered—and the professional opportunities in business, school, clinical, and counseling psychology grew at a rapid rate. STRUGGLES FOR PROFESSIONAL IDENTITY As early as 1915, consulting psychologists had petitioned the APA to recognize the growth of applied psychology by com- mitting some program time at the annual meeting for discus- sion of professional issues. But APA leadership had balked, citing the APA’s sole stated objective as an organization that existed for the advancement of psychology as a science. When the American Association of Clinical Psychologists (AACP) had been founded in 1917, there was concern within the APA that the group would lead to a rupture in organized psychology. In negotiations between the two groups, the AACP agreed to dissolve in 1919 and reorganize as the Clini- cal Section of the APA. The Clinical Section identiﬁed three goals: “promoting better working relationships within clinical and within allied ﬁelds, developing professional standards for practitioners, and encouraging research and publication on topics in clinical psychology” (Napoli, 1981, p. 26). Two years later, in 1921, the APA created a second section on consulting psychology, and the short-lived certiﬁcation pro- gram would stem from the efforts of this group. The consulting/ clinical psychologists recommended two additional APA sec- tions, one on educational psychology and the other on indus- trial psychology, but those two requests were denied. As the professional opportunities for psychologists grew and as problems in professional practice occurred, these psy- chologists made additional requests of the APA. They called on the APA to develop a code of professional ethics. They sought help in protecting the label “psychologist.” They called for changes in graduate training that included additional ap- plied psychology experiences, including internships (which had begun as early as 1908 but were still uncommon, see Routh, 2000). And, they asked that psychology departments hire more faculty who had signiﬁcant practical experience. Except for some minimal gestures toward the applied group, theAPAlargely ignored those requests that were important for the professionalization of psychology, reminding the group of its mantra that the APA was a scientiﬁc association. Throughout the 1920s, more than a dozen applied psy- chology groups were formed, most of them state associations. The largest of those was the New York State Association of Consulting Psychologists, which had begun in 1921. By 1930, it was clear to the professional psychologists that the Postwar Growth of the Practice of Psychology 35 APA was not going to support their efforts. In that year New York University psychologist Douglas Fryer led a reorgani- zation of the New York group, renamed it the Association of Consulting Psychologists (ACP), and extended its geograph- ical boundaries for membership to include the entire United States. The ACP, thus, became the ﬁrst “national” association for professional psychologists. In 1933, the ACP published its code of professional ethics, the ﬁrst such document for psy- chologists. And, in 1937, it began publication of the Journal of Consulting Psychology, arguably the ﬁrst professional psychology journal. ACP struggled to establish itself as the national associa- tion for professional psychologists; however, it was dominated by New York psychologists. In 1935, a plan was initiated to broaden the ACP membership by creating a federation of so- cieties. All the existing state associations were invited to join as well as the Clinical Section of the APA. Eventually the fed- eration plan was abandoned, and it was decided to create a wholly new organization, the American Association for Applied Psychology (AAAP), which began in 1938. The ACP and the Clinical Section of the APA both disbanded and became part of AAAP. The ACP journal was continued by the AAAP—as its ofﬁcial organ. The AAAP began with four sections: clinical, consulting, educational, and industrial psychology. Fryer served as the ﬁrst president of AAAP and was followed in later years by such important applied psychologists as Walter Van Dyke Bingham (1880–1952) and Carl Rogers (1902–1987). The AAAP’s success was manifested largely through its sections in which psychologists with similar needs could work to- gether on issues of common concern. Each section wrote its own by-laws, elected its own ofﬁcers, created its own com- mittees, and planned its own program at the annual meeting of the AAAP. Even though most of the AAAP members retained their mem- berships in the older APA, many identiﬁed more strongly with the new organization than with APA because AAAP provided the professional identity, the collegial relations, and the professional assistance that APA had been unwilling to offer. (Benjamin, 1997, p. 728) Although the AAAP was quite successful in serving the needs of professional psychologists, the organization lasted only slightly more than seven years. Its demise had nothing to do with the service it was providing for the growing profes- sion of psychology. With the United States at war in 1942, there was federal government pressure on the various psychological organizations to come together with one voice for the national good. Negotiations among several groups (including the Society for the Psychological Study of Social Issues, or SPSSI, and the Psychometric Society) but princi- pally steered by the two heavyweights, the APA and the AAAP, led to the establishment of a “new” American Psychological Association. The new APA began with 18 charter divisions, a model borrowed from the sectional structure of the AAAP; a new journal that was intended to be a journal of “professional psy- chology,” the American Psychologist (Benjamin, 1996); and a new central ofﬁce in Washington, D.C. (Capshew, 1999). The new APA also had a new statement of objectives which read: “to advance psychology as a science, as a profession, and as a means of promoting human welfare” (Wolﬂe, 1946/1997, p. 721). The “professional” goal had come, of course, from the AAAP, and the “human welfare” goal from the SPSSI. The APA looked and sounded like a new kind of organization, one that had ﬁnally acknowledged the presence of the profession of psychology. However, professional psy- chologists would soon learn that they had little real support (or power) within the new association. It would be almost 30 years before that situation changed in any dramatic way. POSTWAR GROWTH OF THE PRACTICE OF PSYCHOLOGY Whereas American psychologists were caught napping by the First World War, they did not repeat that mistake for the sec- ond one. Both the APA and the AAAP had committees in place by 1939 to plan for psychology’s role should the United States enter the war. As noted earlier, in the ﬁrst war psychol- ogists worked largely in two areas: examination of recruits and personnel selection. However, in the Second World War, the involvement of psychologists was substantially more diverse—and it included recruitment, selection, training, equipment design, propaganda, surveying attitudes in the United States and abroad, examining and testing prisoners of war, morale studies, intelligence work, and personality stud- ies, including an analysis of Adolf Hitler (Capshew, 1999; Hoffman, 1992). The verdict on psychologists’ performance in the war was an incredibly favorable one. The legacy of that performance was a growth in scientiﬁc and professional opportunities for psychologists that was unprecedented in psychology’s history. The profession beneﬁted particularly, and no group beneﬁted more than clinical psychology. Clinical Psychology Early in the war, the federal government began planning to meet the mental health needs of returning veterans, which were judged to be substantial. Perhaps the government hoped 36 Psychology as a Profession to avoid the hard feelings among veterans that occurred as a result of their poor treatment following the First World War, ill feeling that had led to a massive march on Washington, D.C. It was evident in 1942 that psychiatrists were too few in number to provide the necessary clinical services, so the federal government mandated that the United States Public Health Service (USPHS) and the Veterans Administration (VA) signiﬁcantly expand the pool of mental health profes- sionals. That translated into increasing the availability of clinical psychologists. The USPHS and VA worked with the new APA to expand doctoral training programs in clinical psychology and to identify programs of acceptable quality. The latter goal led to the formation of the APA’s accreditation program for clinical psychology programs in 1946 and for counseling psychology programs in 1952. The former goal initiated a series of meet- ings with department heads of doctoral psychology programs who had extant clinical psychology programs or were inter- ested in developing such programs. The USPHS promised funding to university graduate programs to support clinical psychology students, and the VA promised funding for prac- tica and internship training (Moore, 1992). Because the GI bill had been altered to include beneﬁts for graduate study, money was also available from that program to support doctoral training for veterans, and many chose to pursue advanced study in psychology, with much of that interest directed toward clinical psychology. Although an accreditation process was already in place within the APA as of 1946, there was no agreed-upon model for clinical training. Discussions of such models dated to the 1890s with a proposal from Witmer, which was followed by subsequent curriculum and training proposals by APA’s Clin- ical Section in 1918–1919 in a series of articles in the Jour- nal of Applied Psychology, by the ACP, and by the AAAP. As a leader in the AAAP, clinical psychologist David Shakow (1901–1981) was the key ﬁgure in drafting a model curricu- lum for clinical training. He developed a proposal for the AAAP in 1941 that shaped all subsequent discussions, lead- ing to the report of the Committee on Training in Clinical Psychology (CTCP), an APA committee founded in 1946 with Shakow as chair and funded by the VA and the USPHS. The committee’s formidable charge was to (a) formulate a recommended program for training in clinical psychology, (b) formulate standards for institutions giving training in clinical psychology, including both universities and internship and other practice facilities; © study and visit institu- tions giving instruction in clinical psychology and make a detailed report on each institution. (Baker & Benjamin, 2000, p. 244) Shakow and his committee published their report in 1947 (American Psychological Association, 1947). Two years later it became the framework for the most famous report in the history of professional training in psychology, the “Boulder Report.” That report was the result of the joint work of 73 individuals from psychology and related ﬁelds who came together in Boulder, Colorado, for two weeks in the summer of 1949 to produce a model of clinical training in psychology that became known as the “Boulder model” or “scientist- practitioner model” (Raimy, 1950). The architects of this model argued that it was both possible and desirable to train clinical psychologists as competent practitioners and scien- tists, a view that continues to be debated today. Not only was there a new formal model for clinical train- ing, but there was a new model for the clinical psychologist as practitioner (one that involved training as a psychotherapist, a role for psychologists that was strongly supported by the fed- eral government). Clinical psychologists would break from their tradition in psychometrics to focus on the delivery of psychotherapy. In 1948, the federal government established the National Institute of Mental Health, which gave further impetus to both the training in and practice of clinical psy- chology (VandenBos, Cummings, & DeLeon, 1992). The turf disputes with psychiatry had been minor skirmishes before the war, but bigger battles were about to break out as psychol- ogists began to be true competitors of psychiatrists. As the numbers of psychologists who worked as practi- tioners grew, the pressures for certiﬁcation, licensing, and even insurance reimbursement for clients again surfaced within the profession. Connecticut was the ﬁrst state to enact a psychologist certiﬁcation law in 1945. Over the next 30 years, professional psychologists worked state by state to get state legislatures to pass laws creating psychology licens- ing boards. These efforts were largely the responsibility of state psychological associations, although by 1970 the APA began providing some coordination and consultation. In the mid-1950s, the Board of Professional Affairs was created by the APA, with the mission to establish standards for profes- sional practice, foster the application of psychological knowledge, and maintain satisfactory relations with other professions (American Psychological Association, 1957). The struggles for equality were not only in the legislatures but also with insurance companies and employers. Employer- paid health insurance had emerged as an employee beneﬁt dur- ing World War II. During the 1950s and 1960s, labor unions sought to achieve such coverage and expand it (and to include psychotherapy services). After years of urging by practition- ers, the APA created an Ad Hoc Committee on Insurance and Related Social Developments in 1963 to meet with insurance industry ofﬁcials in order to get psychologists included in Postwar Growth of the Practice of Psychology 37 reimbursement plans (and for such reimbursement to be at parity with that of psychiatrists). Leonard Small, Rogers Wright, Milton Theaman, and Nicholas Cummings were central in this undertaking. The committee also created model “freedom-of-choice” legislation, which individual state psy- chological associations could try to get adopted in their state (Cummings, 1979). Legislative language was such that if an insurance company reimbursed a psychiatrist for a particular service, it must also provide such reimbursement when the same service is provided by a qualiﬁed licensed psychologist. Later, professional psychologists would use the courts as well in their struggle for equality. It took, for example, a lawsuit ﬁled by the APA against the American Psychoana- lytic Institute to establish the right of psychologists to be trained in psychoanalytic centers controlled by the institute (DeAngelis, 1989). The APA Practice Directorate, which was formed in the early 1990s by combining the Ofﬁce of Professional Practice and the Ofﬁce of Professional Affairs, evaluates cases and develops selected ones that further the independent practice of psychology. The directorate’s efforts are partly funded by a yearly special assessment to all mem- bers who engage in practice activities. Divisions of the APA in which at least 50% of its members contribute to the as- sessment are identiﬁed as “practice Divisions.” Efforts outside of the APA also contributed to the devel- opment of standards of excellence for practitioners. The American Board of Examiners in Professional Psychology (ABEPP) was created in 1947 “to award diplomas for ad- vanced competency in the ﬁeld” (Riess, 1992, p. 769). Later the term “Examiners” was omitted, and at least ﬁve separate specialty boards exist today under the aegis of the parent organization. Following the conference in Boulder, several other con- ferences were held to establish training guidelines for a clinical as well as for other professional subspecialties (see Cohen, 1992), but the 1973 “Vail Conference” (also in Colorado) gave credence to the burgeoning programs offer- ing doctor of psychology (PsyD) degrees from universities as well as from freestanding schools (Korman, 1974). The history of the establishment of professional schools and the PsyD degree has been well documented by Peterson (1992) and Stricker and Cummings (1992). As of June 2001, there were 53 such schools accredited by the APA. Professional schools now graduate over 50% of all clinical students. Counseling Psychology As a profession, counseling psychology changed consider- ably following the war. Vocational guidance remained a duty, but that work would soon shift primarily to guidance counselors within secondary schools. And, the selection du- ties that had occupied many in vocational guidance became more exclusively the property of industrial psychologists. In place of these activities, “psychotherapy” came to counseling psychology, initially through the writings and teachings of Carl Rogers, who trained many counseling psychologists after the war in “non-directive” counseling and therapy techniques. The 1950s proved to be a decade of crisis for counseling psychologists. It was a crisis of identity, or at least role con- fusion. Counseling psychologists who previously garnered most of their identity as vocational counselors had been called on in increasing numbers to provide a range of services to military veterans both in hospital settings and community service centers. Rehabilitation took on a broader meaning, and in addition to vocational planning, counselors were working on general issues of adjustment with service person- nel seeking to integrate into the general society. Likewise, the role of student personnel workers in higher education began to focus more broadly on student adjustment. Changes for the counseling profession in the 1950s were evidenced by several clear markers. “Counseling psychol- ogy” became the appellation of choice at the Northwestern Conference of 1951, a meeting speciﬁcally organized to explore changes in the ﬁeld and to make plans for the future. Out of that conference came several initiatives that affected Division 17, the APA, and the VA. In 1952, Division 17 changed its name from “Counseling and Guidance” to “Counseling Psychology.” The Veterans Administration established two new psychological job descrip- tions: Counseling Psychologist (Vocational) and Counseling Psychologist. In that same year, the APA began accrediting doctoral programs in counseling psychology, partly in re- sponse to a doctoral training curriculum recommended by a Division 17 committee (APA, 1952). The ﬁnal identifying component of a profession was added in 1954 with the establishment of a new publication, the Journal of Counseling Psychology. It might seem that counseling psychology had arrived as a profession. Such professionals had an organizational home, a journal, doctoral training programs, and jobs. There were, however, continued difﬁculties in deﬁning the ﬁeld that led to a Division 17 Committee on Deﬁnition report in 1956 (Amer- ican Psychological Association, 1956) and a “crisis” report on counseling psychology as a profession, written in 1960. This latter report was initiated by the APA’s Education and Training (E&T) Board, which appointed a three-person committee to prepare a report on the status of counseling psychology as a professional specialty (Berg, Pepinsky, & Shoben, 1980). 38 Psychology as a Profession The leadership of Division 17 was not pleased with the unilateral actions of the E&T Board. When the E&T report appeared, the division commissioned its own three-person committee, which drafted a much more optimistic report on the status of counseling psychology arguing that the profes- sion was thriving, even if graduate programs were not. This 1961 report found that, The rate of growth of counseling psychology has been normal despite limited ﬁnancial support for the development of graduate programs and the support of graduate students. . . . The social demand for well prepared counseling psychologists is great and continues to increase. The Division of Counseling Psychology has a deep professional obligation to meet this social need. (Tyler, Tiedeman, & Wrenn, 1980, p. 124) Part of the dissatisfaction within counseling psychology was caused by its comparison with clinical psychology, a profession that was growing at a fantastic rate. By that yard- stick, any ﬁeld would have looked to be in trouble. There was concern from many in counseling that the ﬁeld should clearly distinguish itself from clinical psychology, whereas others suggested merging the training of the two ﬁelds while main- taining differences in the nature of practice. Traditional work in vocational guidance had been modi- ﬁed by the experiences of counseling psychologists in the VA and student personnel work in higher education. What emerged was a new specialty area that had as its focus the adjustment of the individual to the demands of everyday life, whether those demands were vocational, educational, or interpersonal. The emphasis on developmental processes of average individuals facing day-to-day life was seen as a clear contrast to the emphasis on psychopathology that was the bread and butter of the clinical psychologist. Industrial Psychology Other practice specialties also beneﬁted from psychologists’ record of accomplishment during the war. Historian Donald Napoli (1981) wrote this about the postwar growth of indus- trial psychology: The military had given psychologists a chance to prove the effectiveness of selection, classiﬁcation, and aptitude testing, and psychologists met the challenge successfully. Civilian em- ployers also offered new opportunities, which grew largely from the labor shortage produced by wartime mobilization. Business managers, beset by high rates of absenteeism and job turnover, took unprecedented interest in hiring the right worker and keeping him contented on the job. Management turned to psychologists . . . and the amount of psychological testing quickly increased. Surveys show that in 1939 only 14% of busi- nesses were using such tests; in 1947 the proportion rose to 50%, and in 1952, 75%. (p. 138) Another area of substantial development for the industrial psychologist that grew out of the wartime work was the ﬁeld of human factors or engineering psychology. The military, in particular, continued to employ psychologists in its research on human–machine interactions, but businesses as well began to employ psychologists to design irons, telephones, arc welders, vending machines, chemical reﬁneries, and the like. Human factors remained an important part of industrial psy- chology into the 1960s but gradually separated, a transition begun in the late 1950s when APA’s Division 21 (Engineering Psychology) and the Human Factors Society were founded. It was replaced by psychologists interested in applying social psychological theories to the problems of organizations, lead- ing to the growth of the “O” half of the I-O psychologist. Prior to the war, most industrial psychologists served as consultants to businesses, thus working part-time as profes- sionals. After the war, however, that pattern changed dramati- cally. Businesses offered full-time employment opportunities, and consequently graduate programs began to train the I-O practitioners to ﬁll those jobs. School Psychology Unlike the other three practice specialties, the Second World War had much less impact on the practice of school psychol- ogy. Such practice has always been more circumscribed, as the label would imply. Furthermore, whereas the doctoral degree has been assumed to be the minimal level of training necessary for professional practice in the other three special- ties, historically most school psychologists have practiced with a master’s degree or specialty credential. Further, in the ﬁrst half of the twentieth century, school psychologists came from many different educational backgrounds, sometimes with little training in psychology. Fagan (1990) has divided the history of school psychology into “Hybrid years” (1890–1969) and “Thoroughbred years” (1970 to present). The Hybrid years describe a period when school psychology was “a blend of many kinds of educa- tional and psychological practitioners loosely mobilized around a dominant role of psychoeducational assessment for special class placement” (p. 913). That role still exists in the Thoroughbred years, but the practitioner is more narrowly deﬁned as a school psychologist, typically someone who has a master’s or doctoral degree in school psychology from a nationally accredited program. A “Professional” Journal within APA 39 The ﬁrst master’s degree training program for school psy- chologists was initiated at New York University in 1928 and the ﬁrst doctoral training program at the University of Illinois in 1953. The APA did not begin accrediting doctoral pro- grams in school psychology until 1971, and only accredits at the doctoral level. Master’s degree programs are accredited by the National Association of School Psychologists (NASP), an organization founded in 1969. Like the other practice specialties, there have been sig- niﬁcant postwar changes for school psychologists as they, too, have struggled to ﬁnd their identity as a profession (see the report of the Thayer Conference, Cutts, 1955). Still, the Thoroughbred years have been ones of tremendous growth in training and practice for the ﬁeld. Psychological services in the schools have increased dramatically since the 1970s, in part stimulated by America’s baby boom, but also by federal legislation on education, particularly laws on special educa- tion such as the landmark Public Law 94-142, enacted in 1974, which mandated education for all children regardless of handicap. A “PROFESSIONAL” JOURNAL WITHIN APA When the new APA was formed in 1946, a new “profes- sional” journal was established, the American Psychologist. Initially, many articles on professional training and profes- sional job opportunities were published in the AP. After 10 years, the AP was serving a broader associationwide role, and the statement about “the professional journal” of psy- chology was quietly removed in 1957. It would be 12 more years before practitioners got back a “professional” journal from the APA. In 1966, Donald K. Freedheim was asked by George W. Albee, who was then president of the Division of Clinical Psychology (12), to edit the newsletter of the division, which was a mimeographed publication. A magazine-like format, with a new logo, was developed. The format lent itself to having pictures, which enhanced the readability of the publication, but also helped to identify authors at conventions. With this new professional- looking publication, the editor invited contributions from members of other service divisions (e.g., school, indus- trial, counseling), as they were facing similar issues of standards of practice, training, and licensing that were of concern to the clinical members. Submissions from across the spectrum of professional fields in psychology grew, and there were clearly important issues that all the spe- cialties shared. The APA had a ﬁne stable of scientiﬁc journals at the time but no publication that was appropriate for the sorts of material important to the practicing community—policy issues, case histories, training and internship opportunities, and so forth. It was apparent that a truly professional practice–oriented journal was needed. The APA had just received a large grant from the National Science Foundation to develop new, innovative publications on an experi- mental basis. “The Clinical Psychologist” was about to be transformed into an “experimental publication” called Pro- fessional Psychology, with an editorial board made up of members from across subspecialty ﬁelds. The inaugural issue, fall 1969, contained “The Clinical Psychologist,” and the cover of the journal retained the logo that had been de- veloped for the newsletter. By the second issue of the quar- terly, “The Clinical Psychologist” was pulled out to be pub- lished separately. The transition from newsletter to journal was not always smooth. “The Clinical Psychologist” had carried book reviews, but none of the APA journals did. All APA-published reviews were in Contemporary Psychology. The PP editor believed it important to retain reviews for both the conve- nience of the readership and the clear fact that few practice- type publications would be reviewed in CP. After much discussion, the review section was allowed in the new jour- nal, a major exception by the APA Publications and Commu- nications Board (P&C Board). A similar, though less crucial, matter came up regarding authors’ pictures, which had sel- dom appeared in APA journals, except in the American Psy- chologist. Not only were pictures maintained in Professional Psychology, but they started appearing in other APA journals as well, beginning with Contemporary Psychology. Another conﬂict emerged over the size of the publication, as the APA Journals Ofﬁce wanted it to be in the standard 7-by-10-inch format then instituted for all other APA journals (except the American Psychologist, which was larger). The newsletter had been in a 6-by-9-inch format, and the PP editor believed that its successor should retain its distinct (and convenient) size, in part to distinguish it from the scientiﬁc journals. With the editor threatening to withdraw from the publication, the smaller format prevailed—at least for the seven-year term of the editor. After a year of being in experimental status, and submissions growing monthly, the quarterly was made an “ofﬁcial” APA publication—and the editor allowed to serve on the Council of Editors. In 1983, the title of the journal expanded to Professional Psychology: Research and Practice. PP is currently pub- lished six times a year. During the editorial term of Patrick H. DeLeon (1995–2000), with Gary R. VandenBos serving as the managing editor, Professional Psychology made an even greater effort to address the interests of the practice commu- nity. After conducting three reader surveys during the ﬁrst 40 Psychology as a Profession year (one of which involved an innovative nationwide tele- phone conference call hookup involving over 50 sub- scribers), PP readers made crystal clear that they wanted articles that “provide practical advice and concrete sugges- tions that could be implemented in everyday practice set- tings, rather than merely placing the new ﬁndings within the context of the existing published literature (and then com- menting upon needed future research)” (DeLeon & Vanden- Bos, 2000, p. 595). PP ’s coverage included managed care, prescription privi- leges for psychologists, telehealth care, expanding roles for psychologists within the public policy (including legislative and administrative) arena, and behavioral health-service de- livery within primary care. Each of these issues has become of major concern to the profession and to the nation’s overall health-delivery system during the past decade. The reader- ship numbers (individual and institutional) steadily increased to approximately 8,000, making Professional Psychology the second most popular subscribed to APA journal. In retrospect, the concerted effort to promulgate APA Practice Directorate efforts and relevant federal public health initiatives (e.g., those of the U.S. Surgeon General), although perhaps highly unusual for an APA journal, have had an impact in educating the ﬁeld regarding the changes evolving within their practice environments. Also, efforts to engage women and ethnic minorities in the editorial process (and thereby enrich the breadth of coverage) were particularly successful. The overall percentage of ethnic minority mem- bers in the APA at the time was 5.38%; in sharp contrast, in 1999 three of the ﬁve Professional Psychology associate edi- tors were female, and two associate editors were members of ethnic minorities. Further, 34.7% of the editorial board were female and 14.7% were ethnic minorities. This was a signiﬁ- cantly higher percentage of both categories of members than almost any other APA journal. ELECTRONIC PUBLISHING INITIATIVES In January 1995, the APA Web site was available to its mem- bers and the public. The total “hits” in the ﬁrst three months was 22,474—a ﬁgure that today (in 2001) is reached every 30 minutes. Usage increased in every quarter of 1995, reach- ing 413,207 hits in the fourth quarter of that year; quarterly hits in 2001 ran at 90 million. It is interesting to note that many people from a vast array of ﬁelds turn to theAPAfor information on how to reference elec- tronic documents. The APAhas a special “style page” on elec- tronic citations. A million people access this specialized page on a relatively narrow topic every year. Nonpsychologist (and nonstudent) use of the APA Web site remains strong. Almost 35% (or some four million annual users) of the APA Web site are not psychologists or students studying psychology. In 1997, APA president-elect Martin Seligman proposed the establishment of an electronic journal called Treatment, to be published jointly with the American Psychiatric Associ- ation. For political reasons the “other APA” withdrew from the venture because of fear that psychologists might claim that reading the copublished journal would qualify them for prescribing medication. The American Psychological Associ- ation then decided to embark on the e-journal alone, which is now titled Prevention and Treatment. By the summer of 2001, under Seligman’s editorship, the journal had 20,000 regular readers, with each article being “hit” an average of 35,000 times within the ﬁrst year of release. Publications from the Practice Directorate, the e-journal, Web-based com- munications, videotapes, and over 70 new books each year constitute communications from the APA that are directed toward the practicing professional psychologist. COMMUNITY INVOLVEMENT During the 1990s, the APA became increasingly involved in several highly visible community activities that contributed to society’s appreciation of the role of psychological services. In 1991, during the Gulf War, the APA joined with the American Red Cross in forming a network of psychologists to provide mental health services to families of members of the armed services. Since then the network has been activated following natural disasters, airline crashes, and terrorist attacks in Oklahoma City (1996) and in New York City and Washington, D.C., in September 2001. In another effort, the Practice Directorate forged a part- nership with Music Television (MTV) to develop a youth antiviolence initiative titled “Warning Signs,” to help the na- tion’s youth in identifying early signs of violent behavior and to emphasize the need to get help should they see any of them (Peterson & Newman, 2000). The campaign ofﬁcially kicked off with a youth forum held in Los Angeles on April 22, 1999. The 30-minute documentary, coproduced by MTV and the APA, was the highest rated prosocial special in MTV’s history, with 3.9 million youth watching the ﬁlm. In that year, there were over 600 follow-up psychologist-led “Community Youth Forums on Violence” held across the nation, with more than 58,000 youths attending. In March 2000, the Practice Directorate launched “Warning Signs for Parents” as a logical follow-up; by the end of the year, nearly 150,000 copies of the accompanying publication had been distributed. The Prescriptive Authority (RxP-) Agenda 41 TWO ASSOCIATIONAL DEVELOPMENTS The essence of a profession is daily involvement with patients or clients. The context in which this interaction occurs, including the very important issue of reimbursement for services rendered, falls within the jurisdiction of public policy (e.g., the political process). For psychology to become an active participant within primary care (or to expand its scope of practice to include prescriptive authority) requires institutional collective knowledge of the evolving “bigger picture” and ongoing interrelationships existing within soci- ety and the generic health care arena. Historically, profes- sional psychology has, at most, seen itself as solely one of the mental health disciplines and has not concerned itself with broader public policy or public health issues (DeLeon, VandenBos, Sammons, & Frank, 1998). These two program- matic initiatives have signiﬁ

bundet

cantly changed that perspective. The APA Congressional Science Fellowship Program In 1974, Pam Flattau served as the ﬁrst APA Congressional Science Fellow, under the program established in conjunction with the American Association for the Advancement of Science (AAAS). Over a quarter of a century later, approxi- mately 125 colleagues have had the opportunity to serve on Capitol Hill (or in the administration) as APA Fellows, Robert Wood Johnson Health Policy Fellows, or in other sim- ilar national programs. In this capacity, they experienced personal involvement in the public policy process. Initially, the APA focused only on providing the experience for recent doctoral graduates; as the program matured, however, a concerted effort was made to attract more senior fellows. The APA Fellows have included individuals from almost every psychological specialty area, including several who also pos- sessed degrees in law (Fowler, 1996). Over the years, a number of psychologists have gravitated to positions of high-level public policy responsibility. During President Lyndon Johnson’s era of the “Great Society,” John Gardner served as secretary of the Department of Health, Education, and Welfare. Psychologists have served as depart- mental assistant secretaries, subject to Senate conﬁrmation; director of one of the National Institutes of Health (NIH), as well as of other federal research institutes; head of the federal Bureau of Prisons; commanders of federal health care facili- ties; and as Chief State Mental Health ofﬁcials. In the 107th Congress (2001–2002), three psychologists were elected to U.S. House of Representatives, and 12 psychologists served in the various state legislatures during that same time. With ﬁrsthand experience in the public policy process, psy- chologists have been inﬂuential in the gradual modiﬁcation of statutes and implementation of regulations that recognize psychology’s expertise. In the clinical arena, psychology’s expertise is now independently recognized throughout the judicial system and under all federal and private reimburse- ment systems. Psychology’s professional graduate students are supported under almost every federal training and service delivery initiative. The underlying unanswered question remains, however: Has professional psychology matured sufﬁciently to establish its own programmatic agenda via the public policy process (VandenBos, DeLeon, & Belar, 1991)? APAGS In 1988, the APA Council of Representatives formally estab- lished the American Psychological Association of Graduate Students (APAGS). Over the years, psychology has contin- ued to be one of, if not the, most popular undergraduate majors. By 2001, the APA membership (and afﬁliate) num- bers had grown to 155,000, with the APAGS possessing 59,700 members. An APAGS representative attends the open portions of the APA board of directors meetings (and another individual is seated on the ﬂoor of the Council of Representatives as a nonvoting member). Increasingly, as with other professions, the student voice is being heard. Several divisions, state asso- ciations, and council caucuses provide the APAGS with a voting seat on their boards of directors. Student participation brings to the APA governance delib- erations a unique focus upon the “here and now” practical consequences. The APAGS’s presence constantly reminds those within the APA governance that their deliberations do have very real consequences on future generations of profes- sional psychologists. THE PRESCRIPTIVE AUTHORITY (RxP-) AGENDA In November 1984, Senator Daniel K. Inouye addressed the annual meeting of the Hawaii Psychological Association and in closing suggested to them an entirely new legislative agenda that he proposed would ﬁt nicely into their conven- tion theme “Psychology in the 80’s: Transcending Traditional Boundaries” (e.g., seeking prescriptive authority in order to better serve their patients). After his challenge, the executive committee of the Hawaii Psychological Association agreed to pursue legislation that would study the “feasibility of allow- ing licensed psychologists to administer and prescribe med- ication in the treatment of nervous, mental and organic brain diseases.” At that time there was little enthusiasm for the 42 Psychology as a Profession proposal within the psychological community and extreme opposition within the local psychiatric community (DeLeon, Fox, & Graham, 1991). This, however, was to be the begin- ning of psychology’s prescriptive authority (RxP-) quest. In 1989, the APA Board of Professional Affairs (BPA) held a special retreat to explore the issues surrounding psy- chology obtaining RxP- authority. It concluded by strongly endorsing immediate research and study regarding the feasi- bility and the appropriate curricula in psychopharmacology so that psychologists might provide broader service to the public and more effectively meet the psychological and mental health needs of society. Further, the BPA also recom- mended that focused attention on the responsibility of prepar- ing the profession to address current and future needs of the public for psychologically managed psychopharmacological interventions be made APA’s highest priority. Interestingly, in the 1970s, the APA board of directors had appointed a special committee to review this very matter. The recommendation at that time was that psychology not pursue prescription privi- leges, primarily since the ﬁeld was doing so well without that authority! (DeLeon, Sammons, & Fox, 2000). At the APA annual convention in Boston in 1990, the mo- tion to establish an ad hoc Task Force on Psychopharmacol- ogy was approved by a vote of 118 to 2. Their report back to council in 1992 concluded that practitioners with combined training in psychopharmacology and psychosocial treatments could be viewed as a new form of health care professional, expected to bring to health care delivery the best of both psy- chological and pharmacological knowledge. Further, the pro- posed new provider possessed the potential to dramatically improve patient care and make important new advances in treatment (Smyer et al., 1993). On June 17, 1994, APA president Bob Resnick was for- mally recognized during the graduation ceremonies at the Walter Reed Army Medical Center for the ﬁrst two Depart- ment of Defense (DoD) Psychopharmacology Fellows, Navy Commander John Sexton and Lt. Commander Morgan Sammons. This program had been directed by the Fiscal Year 1989 Appropriations bill for the Department of Defense (P.L. 100–463) (U.S. Department of Defense, 1988) and would ultimately graduate 10 fellows. Upon their graduation, each of these courageous individuals became active within the practitioner community, demonstrating to their col- leagues that psychologists can indeed readily learn to provide high-quality psychopharmacological care. Several of the graduates have become particularly involved in providing consultation to evolving postdoctoral psychopharmacology training programs. All of the external evaluations of the clinical care was provided by the DoD Fellows (ACNP, Summer, 2000). At its August 1995 meeting in New York City, the APA Council of Representatives formally endorsed prescriptive privileges for appropriately trained psychologists and called for the development of model legislation and a model train- ing curriculum. The follow year in Toronto, the council adopted both a model prescription bill and a model training curriculum. Those seeking this responsibility should possess at least 300 contact hours of didactic instruction and have supervised clinical experience with at least 100 patients requiring psychotropic medication. In 1997, the APAGS adopted a “resolution of support” for the APA position. And, that same year, at the Chicago convention, the council autho- rized the APA College of Professional Psychology to develop an examination in psychopharmacology suitable for use by state and provincial licensing boards. This exam became available in the spring of 2000. As of the summer of 2001, approximately 50 individuals had taken the examination, which covers 10 predetermined distinct knowledge areas. By late 2001, the APA Practice Directorate reported that RxP- bills had been introduced in 13 states and that the APA Council had demonstrated its support for the agenda by allo- cating contingency funding totaling $86,400 over 5 ﬁscal years. In its February 2001 reexamination of the top priorities for APA’s future, the APA Council of Representatives had placed advocacy for prescription privileges as number six of 21 ranked priorities for the association. While no comprehen- sive bill has yet passed, the U.S. territory of Guam has passed legislation authorizing appropriately trained psychologists to prescribe in the context of a collaborative practice arrange- ment with a physician. During the spring of 2001, a psycholo- gists’ prescriptive authority bill only very narrowly missed passage in New Mexico, successfully making it through two House committees, the full House, and a Senate commit- tee. Further, we would note that a reading of an amendment to the Indiana Psychology Practice Act, which passed in 1993, indicates that psychologists participating in a federal government–sponsored training or treatment program may prescribe. Thirty-one state psychological associations cur- rently have prescription privileges task forces engaged in some phase of the RxP- agenda. Patrick H. DeLeon has had the pleasure of serving as the commencement speaker for three postdoctoral masters’ psychopharmacology graduations (in Louisiana, Texas, and Florida). By the summer of 2001, co- horts of psychopharmacology classes had also graduated in Georgia (two separate classes), Hawaii, and New Mexico, with additional cohorts enrolled in several different states.The Prescribing Psychologists’Register (PPR) also reports having graduated a signiﬁcant number of students. Psychology’s RxP- agenda is steadily advancing (DeLeon, Robinson- Kurpius, & Sexton, 2001; DeLeon & Wiggins, 1996). References 43 THE TWENTY-FIRST CENTURY Unquestionably, the psychological practice environment of the twenty-ﬁrst century will be dramatically different than it is today. The speciﬁcs of change are, of course, unpre- dictable. However, at least one major trend is clear. Our nation’s health care system is just beginning to appreciate the applicability of technology, particularly computer and telecommunications technology, to the delivery of clinical services. The Institute of Medicine (IOM), which has served as a highly respected health policy “think tank” for adminis- trations and the Congress since its inception in 1970, reports that Health care delivery has been relatively untouched by the revo- lution in information technology that has been transforming nearly every other aspect of society. The majority of patient and clinician encounters take place for purposes of exchanging clin- ical information. . . . Yet it is estimated that only a small fraction of physicians offer e-mail interaction, a simple and convenient tool for efﬁcient communication, to their patients. (Institute of Medicine, 2001, p. 15) The number of Americans who use the Internet to retrieve health-related information is estimated to be about 70 mil- lion. Currently, over half of American homes possess com- puters, and while information presently doubles every 5 years, it will soon double every 17 days, with trafﬁc on the Web already doubling every 100 days (Jerome et al., 2000). And, at the same time, the IOM further reports that the lag between the discovery of more efﬁcacious forms of treatment and their incorporation into routine patient care is unnec- essarily long, in the range of about 15 to 20 years. Even then, adherence of clinical practice to the evidence is highly uneven. The era of the “educated consumer” is upon us. How con- sumer expectations and the unprecedented explosion in communications technology will affect the delivery of psy- chological care is yet to be determined. Highly complex issues such as reimbursement for virtual therapy environments, automated diagnostic testing protocols, ensuring psychologi- cally based enriched living and long-term care environments for senior citizens and the chronically ill, not to mention ﬁnancial support for clinical graduate students, will all be debated in the public policy (e.g., political) arena. Professional psychology must become active participants in this critical— and ongoing—dialogue, in order to ensure the future of pro- fessional psychology, research in applied psychology, basic psychological research, and the public welfare in terms of health care and social services. REFERENCES American Psychological Association, Committee on Training in Clinical Psychology. (1947). Recommended graduate training program in clinical psychology. 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ZOLA 47 THE MIND 47 THE BRAIN 48 SENSORY PROCESSES 51 Color Vision 51 Pitch Detection 52 LEARNING AND MEMORY 53 MOTIVATION AND EMOTION 56 Emotion 56 Motivation 57 COGNITIVE NEUROSCIENCE 59 CONCLUSION 62 REFERENCES 62 The great questions of philosophy, the mind–body problem and the nature of knowledge, were also the questions that drove early developments in the pathways to modern psy- chology. This is especially true of biological or physiological psychology. Wilhelm Wundt, who founded experimental psy- chology, titled his major work Foundations of Physiological Psychology (1874/1908). William James, the other major ﬁg- ure in the development of modern psychology, devoted a third of his inﬂuential text Principles of Psychology (1890) to the brain and nervous system. Both Wundt and James studied medicine and philosophy, and both considered themselves physiologists. Their goal was not to reduce psychology to physiology but rather to apply the scientiﬁc methods of phys- iology to the study of the mind. The other driving force in early biological psychology was the study of the brain and nervous system. The major topics in modern biological psychology are sen- soryprocesses, learning andmemory,motivation and emotion, and most recently cognition—in short, behavioral and cogni- tive neuroscience. A number of other areas began as part of physiological psychology and have spun off to become ﬁelds in their own right. We treat the major topics in biological psy- chology separately in the text that follows. But ﬁrst we sketch very brieﬂy the recent philosophical and physiological roots. THE MIND The history of such issues as the mind–body problem and epistemology is properly the domain of philosophy, treated extensively in many volumes and well beyond the scope of this chapter and the expertise of these authors. Our focus in this brief section is on the history of the scientiﬁc study of the mind, which really began in the nineteenth century. Perhaps the ﬁrst experimental attacks on the nature of the mind were the observations of Weber as generalized by Gustav Fechner. Ernst Weber, a physiologist, was attempting in 1834 to determine whether the nerves that respond to the state of the muscles also contribute to judgments about weights. He found that the just noticeable difference (jnd) in weight that could be reliably detected by the observer was not some absolute amount but rather a constant ratio of the weight being lifted. The same applied to the pitch of tones and the length of lines. Fechner realized that Weber had discovered a way of measuring the properties of the mind. Indeed, in his Elements of Psychophysics (1860/1966) he felt he had solved the prob- lem of mind and body. He generalized Weber’s observations to state that as the psychological measurement in jnd’s in- creased arithmetically, the intensity of the physical stimulus increased geometrically—the relationship is logarithmic. Fechner, trained as a physicist, developed the classical psy- chophysical methods and the concepts of absolute and differ- ential thresholds. According to Edwin Boring (1942), he had a nervous breakdown and resigned his chair at Leipzig in 1839. During the last 35 years of his life, he devoted himself to panpsychism, the view that mind and matter are one and thus that mind is all. He viewed the psychophysical law as the paradigm for the transformation of the material into the spir- itual. In any event, the methods Fechner developed were of great help to such early experimental psychologists as Wundt 48 Biological Psychology and his student Tichener in their attempts to measure the at- tributes of sensation. Tichener identiﬁed the elements of conscious experience as quality, intensity, extensity, protensity (duration), and at- tensity (clearness) (see Tichener, 1898). But for all their at- tempts at scientiﬁc observation, the basic approach of Wundt and Tichener was introspection, but other observers (e.g., Külpe at Bonn) had different introspections. Boring studied with Tichener and was for many years chair of the psychol- ogy department at Harvard. He attempted to recast Tichener’s views in more modern terms ( The Physical Dimensions of Consciousness, 1933) by emphasizing that the dimensions listed earlier related to discrimination of physical stimuli. His student S. S. Stevens showed that trained observers could re- liably form judgments of sounds in terms of pitch, loudness, “volume,” and “density” (see also Boring, 1950). At Harvard, Stevens later introduced an important new method of psychophysics termed direct magnitude estimation. The subject simply assigned a number to a stimulus, a higher one to a more intense stimulus and a lower number to one that was less intense. Somewhat surprisingly this method gave very reliable results. Using this method, Stevens found that the proper relationship between stimulus intensity and sensation is not logarithmic, as Fechner had argued, but rather a power function: The sensation, that is, sensory magnitude, equaled the stimulus intensity raised to some power, the exponent rang- ing from less than to greater than one. This formulation proved very useful in both psychophysical and physiological studies of sensory processes (see Stevens, 1975). The key point of all this work on psychophysics is that it is not necessary to be concerned at all about subjective experience or introspection. The observer simply pushes a button or states a word or number to describe his or her judgment of the stimu- lus. The more the observer practices, the more reliable the judg- ments become and the more different observers generate the same results. Psychophysics had become purely behavioral. As Hilgard (1987) notes, Fechner was troubled by the ques- tion of where the transformation between stimulus and judg- ment occurs. Fechner distinguished between “inner” and “outer” psychophysics, outer referring to the relation between the mind and external stimuli and inner to the relation between the mind and excitation of the sensory apparatus. Fechner opted for a direct correspondence between excitation and sen- sation, a surprisingly modern view. Indeed, Stevens (1961) ar- gued with evidence that the psychophysical transformation occurs at the receptor–ﬁrst-order neurons, at least for intensity. We take an example from the elegant studies of Mount- castle, Poggio, and Werner (1963). Here they recorded the ac- tion potentials of a neuron in the somatosensory thalamus of a monkey driven by extension of the contralateral knee. The relation between degrees of joint angle (  ) and frequency of neuron discharge ( F)is F  13.9  0.429  24, where 13.9 and 24 are constants determined by conditions. So the power ex- ponent is 0.429, within the general range of exponents for psychophysical judgments of the relation between joint angle and sensation of movement. In other words, the relationship is established by ascending sensory neuron activity before the level of the cerebral cortex, presumably at the receptor– ﬁrst-order neuron. The modern era of psychophysics can perhaps be dated to a seminal paper by John Swets in 1961: Is there a sensory threshold? His answer was no. He and David Green devel- oped the theory and methodology of signal detection theory (Green & Swets, 1966). There is always noise present with signals. When one attempts to detect a signal in noise, the cri- teria used will determine the outcome. This approach has proved immensely useful in ﬁelds ranging from the telephone to psychophysical studies in animals to detection of structural failures in aircraft wings to detection of breast cancer. But where is the mind in decision theory? It has disappeared. The initial hope that psychophysics could measure the mind has been reduced to considerations of observer bias. A similar conclusion led to the downfall of introspection. THE BRAIN Until the nineteenth century, the only method available to study brain function was the lesion, either in unfortunate hu- mans with brain damage or brain lesions done in infrahuman animals. The key intellectual issue throughout the history of the brain sciences was localization. To state the question in simplistic terms: Are psychological traits and functions local- ized to particular regions of the brain or are they widely dis- tributed in the brain? The history of ideas about localization of brain function can be divided roughly into three eras. During the ﬁrst era, which spans from antiquity to about the second century A . D ., debate focused on the location of cognitive function, al- though the discussion revolved around the issue of the soul, that is, what part of the body housed the essence of being and the source of all mental life (for reviews, see Finger, 1994; Gross, 1987; Star, 1989). In an early and particularly prophetic Greek version of localization of function, the soul was thought to be housed in several body parts, including the head, heart, and liver, but the portion of the soul associated with intellect was located in the head (McHenry, 1969). The individual whom many historians have viewed as having the greatest inﬂuence during this era was Galen, an anatomist of Greek origin. Using animals, he performed experiments that The Brain 49 provided evidence that the brain was the center of the ner- vous system and responsible for sensation, motion, and thinking (Finger, 1994; Gross, 1987). In the second era (spanning the second to the eighteenth centuries), the debate focused on whether cognitive functions were localized in the ventricular system of the brain or in the brain matter itself. The inﬂuence of the church during this era cannot be overstated; for example, ethereal spirits (and ideas) were believed to ﬂow through the empty spaces of the brain’s ventricles. Nevertheless, by the ﬁfteenth and sixteenth cen- turies, individuals such as da Vinci and Vesalius were ques- tioning the validity of ventricular localization. Finally, during the seventeenth century, partly as a result of the strongly held views and proliﬁc writings of Thomas Willis, and during the eighteenth century, with the publication of clinical descrip- tions of cognitively impaired patients accompanied by crude descriptions of brain damage (e.g., Baader), the view that in- tellectual function was localized in brain matter and not in the ventricles became solidiﬁed (Clenending, 1942). The nineteenth century to the present makes up the third era, and here debate has focused on how mental activities (or cognitive processes) are organized in the brain. An early idea, which became known as the localizationist view, proposed that speciﬁc mental functions were carried out by speciﬁc parts of the brain. An alternative idea, which became known as the equipotential view, held that large parts of the brain were equally involved in all mental activity and that there was no speciﬁcity of function within a particular brain area (Clark & Jacyna, 1987). Perhaps the most inﬂuential idea about localization of brain function derived from Franz Joseph Gall during the early nineteenth century. Gall had been inﬂuenced somewhat by the earlier ideas of Albrecht von Haller (Clarke & Jacyna, 1987). In the mid-eighteenth century, Haller had developed a doctrine of brain equipotentiality, or a type of action com- mune . He believed that the parts of a distinguishable anatom- ical component of the brain—the white matter, for instance— performed as a whole, each area of white matter having equivalent functional signiﬁcance (Clarke & Jacyna, 1987). Indeed, one might characterize Gall’s ideas as a reaction against the equipotential view of Haller. Gall’s insight was that, despite its similarity in appearance, brain tissue was not equipotential but instead was actually made up of many dis- crete areas that had different and separate functions. Eventu- ally, Gall was able to characterize 27 different regions, or organs, of the brain in a scheme that he called organology. Later, the term phrenology came to be associated with Gall’s work. However, this term was coined by Gall’s colleague, Spurzheim, with whom he had a falling out, and Gall himself never used the term (Zola-Morgan, 1995). Gall’s ideas about the localization of cognitive functions began to tear at the religious and social fabric of the nine- teenth century. In particular, various governmental and reli- gious authorities saw his notion that various mental faculties were represented in different places in the brain as in conﬂict with moral and religious views of the unity of the soul and mind. Gall’s organology, and later versions of phrenology, faced similar critiques from philosophy and science. Clerics and metaphysicians were concerned with the larger theologi- cal implications of the phrenological system. For example, in Flourens’s critique of phrenology in 1846 (dedicated to Decartes), Gall and his followers were declared guilty of un- dermining the unity of the soul, human immortality, free will, and the very existence of God (Harrington, 1991). Rolando, the famous Italian neuroanatomist, recognized the elegance of Gall’s dissection techniques and his tracing of ﬁber tracts from the spinal cord to the cerebrum. However, he found no logical connection between the tracings of the ﬁbers and the distinct organs in the convolutions of the brain proposed to house particular mental faculties. Another scientiﬁc criticism had to do with the question- able way in which Gall had determined the locus and extent of each of the 27 organs. For example, Gall had localized the carnivorous instinct and the tendency to murder (organ 5) above the ear for three reasons: (a) This was the widest part of the skull in carnivores; (b) a prominence was found there in a student who was fond of torturing animals; and © this region was well developed in an apothecary who later be- came an executioner (Barker, 1897). Another scientiﬁc issue critics raised during the nine- teenth century was that Gall never speciﬁed the precise extent or the anatomical borders of any of the organs. This lack of rigor, it was argued, made it impossible to correlate a speciﬁc faculty with the size of an organ or cranial capacity (Sewall, 1839). Related criticisms involved Gall’s seeming failure to acknowledge that there were variations in the thickness of the skull, that is, variations from one individual specimen to an- other and from one locus to another within the same skull (Sewall, 1839). An oft-cited example of a speciﬁc contribution Gall made to our understanding of brain function is the idea that he an- ticipated the discovery by Broca in 1861 of a speciﬁc speech area of the brain (Ackernecht & Vallois, 1956; Bouillaud, 1848). However, we believe that a careful reading of the facts surrounding this discovery tells a somewhat different story. In fact, Broca never mentioned Gall’s name in his 1861 report. Moreover, he referred to Gall’s doctrine in a rather negative way. Nevertheless, Broca’s work stands as a clear example of a modern idea of localization of function built on the foundation and fundamental idea, established by Gall a 50 Biological Psychology half century earlier, that speciﬁc parts of the brain mediate speciﬁc behaviors. Both Gall and Bouillaud seemed to be vindicated in 1861 with the publication of the proceedings from a meeting of the Société d’Anthropologie de Paris. Broca, assisted by Alexandre Ernest Aubertin, Bouillaud’s son-in-law and a strong believer in localization and in Bouillaud’s hypothesis, presented the neuropathological ﬁndings from the brain of his patient, Monsieur Leborgne. [This patient subsequently was referred to by the name “Tan,” the only utterance Broca ever heard Monsieur Leborgne make (Broca, 1861).] Broca’s ﬁnding from his patient Tan has been regarded by some historians as the most important clinical discovery in the history of cortical localization. Moreover, within the decade, what some historians regard as the most important laboratory discovery pertaining to cortical localization was reported when Gustav Fritsch and Eduard Hitzig (1870) dis- covered the cortical motor area in the dog and proved that cortical localization was not restricted to a single function (Finger, 1994). The discoveries of the speech area by Broca and the motor area by Fritsch and Hitzig were seen as vindi- cation for Gall’s ideas and reestablished him as the father of localization. Following the pioneering study by Fritsch and Hitzig on the localization and organization of the motor area of the cerebral cortex, localization of function quickly won the day, at least for sensory and motor systems. In the last three decades of the nineteenth century, the general locations of the visual and auditory areas of the cortex were identiﬁed. The ﬁeld of physiology, in particular neurophysiology—for example, in the work of Sir Charles Sherrington—together with clinical neurology and neuroanatomy, were exciting new ﬁelds at the beginning of the twentieth century. At this time, the only experimental tools for studying brain organization and functions were ablation and electrical stim- ulation. Neuroanatomy was in its descriptive phase; thanks in part to the Golgi method, the monumental work of Ramon y Cajal was completed over a period of several decades begin- ning near the end of the nineteenth century. Neurochemistry was in its descriptive phase, characterizing chemical sub- stances in the brain. The ﬁrst recording of a nerve action potential with a cathode-ray tube was done by Gasser and Erlanger in 1922, but the method was not much used until the 1930s. The human EEG was rediscovered in 1929 by H. Berger, and the method was applied to animal research and human clinical neurology, particularly epilepsy, in the 1930s by, for example, Alexander Forbes, Hallowell Davis, and Donald Lindsley. The pioneering studies of Adrian in England (1940) and of Wade Marshall, Clinton Woolsey, and Philip Bard (1941) at Johns Hopkins were the ﬁrst to record electrical evoked po- tentials from the somatic sensory cortex in response to tactile stimulation. Woolsey and his associates developed the de- tailed methodology for evoked potential mapping of the cerebral cortex. In an extraordinary series of studies, they de- termined the localization and organization of the somatic sensory areas, the visual areas and the auditory areas of the cerebral cortex, in a comparative series of mammals. They initially deﬁned two projection areas (I and II) for each sen- sory ﬁeld; that is, they found two complete functional maps of the receptor surface for each sensory region of the cerebral cortex, for example, two complete representations of the skin surface in the somatic-sensory cortex. In the 1940s and 1950s, the evoked potential method was used to analyze the organization of sensory systems at all levels from the ﬁrst-order neurons to the cerebral cortex. The principle that emerged was strikingly clear and simple—in every sensory system the nervous system maintained recep- totopic maps or projections at all levels from receptors—skin surface, retina, basilar membrane—to cerebral cortex. The receptor maps in the brain were not point-to-point; rather, they reﬂected the functional organization of each system— ﬁngers, lips, and tongue areas were much enlarged in the pri- mate somatic cortex, half the primary visual cortex repre- sented the forea, and so on. The evoked potential method was very well suited to analy- sis of the overall organization of sensory systems in the brain. However, it could reveal nothing about what the individual neurons were doing. This had to await development of the mi- croelectrode (a very small electrode that records the activity of a single cell). Indeed, the microelectrode has been the key to analysis of the ﬁne-grained organization and “feature detec- tor” properties (most neurons respond only to certain aspects, or features, of a stimulus) of sensory neurons. The ﬁrst intra- cellular glass pipette microelectrode was actually invented by G. Ling and R. W. Gerard in 1949; they developed it to record intracellularly from frog muscle. Several investigators had been using small wire electrodes to record from nerve ﬁbers, for example, Robert Galambos at Harvard in 1939 (auditory nerve; see Galambos & Davis, 1943) and Birdsey Renshaw at the University of Oregon Medical School in the 1940s (dorsal and ventral spinal roots). Metal electrodes were generally found to be preferable for extracellular single-unit recording (i.e., recording the spike discharges of a single neuron where the tip of the microelectrode is outside the cell but close enough to record its activity clearly). Metal microelectrodes were improved in the early 1950s; R. W. Davies at Hopkins developed the platinum-iridium glass-coated microelectrode, D. Hubel and T. Wiesel at Harvard developed the tungsten mi- croelectrode, and the search for putative stimulus coding Sensory Processes 51 properties of neurons was on. The pioneering studies were those of Mountcastle and associates at Hopkins on the organi- zation of the somatic-sensory system (Mountcastle, Davies, & Berman, 1957), those of Hubel and Wiesel (1959) at Harvard on the visual system (and Maturana and Lettvin’s work at MIT on the optic nerve ﬁbers of frogs, see Maturana, Lettrin, McCulloch, & Pitts, 1960), and those of Rose, Hind, Woolsey, and associates at Wisconsin on the auditory system (see Hind et al., 1960). It was not until many years later that imaging methods were developed to study the organization and functions of the normal human brain (see following text). Heroic studies had been done on human brain functioning much earlier in neuro- surgical procedures (heroic both for the surgeon and the patient, e.g., Penﬁeld & Rasmussen, 1950). However, these patients typically suffered from severe epilepsy. The devel- opment of PET, fMRI, and other modern techniques is largely responsible for the explosion of information in the as- pect of biological psychology termed cognitive neuroscience (see following and the chapter by Leahey in this volume). SENSORY PROCESSES We select two examples of sensory processes, color vision and pitch detection, that illustrate very well the historical develop- ment of the study of sensory systems. They are both extraor- dinary success stories in the ﬁeld of biological psychology. Color Vision Color vision provides an illustrative case history of the de- velopment of a ﬁeld in biological psychology with feet in both physics and physiology. Isaac Newton was perhaps the ﬁrst scientist to appreciate the nature of color. The fact that a prism could break up white light into a rainbow of colors meant that the light was a mixture that could produce spectral colors. But Newton recognized that the light rays themselves had no color; rather, different rays acted on the eye to yield sensations of colors (1704/1931). Oddly, the great German literary ﬁgure Goethe asserted it was impossible to conceive of white light as a mixture of colors (1810/1970). In physics there was an ongoing debate whether light was particle or wave (we know now it is both). Interestingly, Newton favored the particle theory. Thomas Young, an English physicist working a century later, supported the wave theory. Newton had developed the ﬁrst color circle showing that complementary pairs of colors opposite to one another on the circle would mix to yield white light. Young showed that it was possible to match any color by selecting three appropriate colors, red, green, and blue, and suggested there were three such color receptors in the eye. Helmholtz elabo- rated and quantiﬁed Young’s idea into the Young-Helmholtz trichromatic theory. Helmholtz, incidentally, studied with Müller and Du Bois-Reymond. He received his MD in 1842 and published two extraordinary works, the three-volume Treatis on Physiological Optics (1856–1866/1924) and On the Sensations of Tone (1863/1954). He was one of the lead- ing scientists in the nineteenth century and had a profound impact on early developments in psychology, particularly bi- ological psychology. The basic idea in the trichomatic theory is that the three receptors accounted for sensations of red, green, and blue. Yellow was said to derive from stimulation of both red and green receptors, and white was derived from yellow and the blue receptor. But there were problems. The most common form of color blindness is red-green. But if yellow is derived from red and green, how is it that a person with red-green color blindness can see yellow? In the twentieth century, it was found that there are four types of receptors in the human retina: red, green, blue (cones), and light-dark (rods). But what about yellow? Hering (1878) developed an alternative view termed the “opponent-process” theory. He actually studied with Weber and with Fechner and received his MD just two years after Wundt in Heidelberg. Interestingly, Hering disagreed with Fechner about the psychophysical law, arguing that the relationship should be a power function, thus anticipating Stevens. Hering proposed that red-green and blue-yellow acted as opposites, along with white-black. In modern times, Dorothea Jameson and Leo Hurvich (1955) provided an ele- gant mathematical formulation of Herring’s theory that ac- counted very well for the phenomena of color vision. Russell De Valois, now in the psychology department at the University of California, Berkeley, provided the physio- logical evidence to verify the Herring-Jameson-Hurvich theory, using the monkey (see De Valois, 1960). Ganglion neurons in the retina that respond to color show “opponent” processes. One cell might respond to red and be inhibited by green, another will respond to green and be inhibited by red, yet another will respond to blue and be inhibited by yellow, and the last type will respond to yellow and be inhibited by blue. The same is true for neurons in the visual thalamus. De Valois’s work provided an elegant physiological basis for the opponent-process theory of color vision. But Young and Helmholtz were also correct in proposing that there are three color receptors in the retina. It is the neural interactions in the retina that convert actions of the three color receptors into the opponent processes in the ganglion cells. It is remark- able that nineteenth-century scientists, working only with the 52 Biological Psychology facts of human color vision, could deduce the physiological processes in the eye and brain. An interesting chapter in the development of color-vision theory is the work of Christine Ladd-Franklin (Hilgard, 1987). She completed her PhD in mathematics at Johns Hopkins in 1882 but was not awarded the degree because she was a woman. Later she spent a year in Müller’s laboratory in Göttingen, where he gave her private lectures because, as a woman, she was not allowed to attend his regular lectures. She developed a most interesting evolutionary theory of color vision based on the color zones in the retina. The center of the fovia has all colors and the most detailed vision. The next outer zone has red and green sensitivity (as well as blue and yellow), the next outer zone to this has only blue and yel- low sensitivity (and black-white), and the most peripheral regions have only black-white (achromatic) sensitivity. She argued that in evolution, the achromatic sensitivity (rods) developed ﬁrst, followed by evolution of blue and yel- low receptors and ﬁnally red and green receptors. The fact that red-green color blindness is most common is consistent with the idea that it is the most recent to evolve and hence the most “fragile.” Modern molecular biology and genetics actually provide support for Ladd-Franklin’s evolutionary hypothesis. The Old World monkey retina appears to be identical to the human retina: Both macaques and humans have rods and three types of cones. It is now thought that the genes for the cone pigments and rhodopsin evolved from a common ances- tral gene. Analysis of the amino acid sequences in the differ- ent opsins suggest that the ﬁrst color pigment molecule was sensitive to blue. It then gave rise to another pigment that in turn diverged to form red and green pigments. Unlike Old World monkeys, New World monkeys have only two cone pigments, a blue and a longer wavelength pigment thought to be ancestral to the red and green pigments of humans and other Old World primates. The evolution of the red and green pigments must have occurred after the continents separated, about 130 million years ago. The New World monkey retina, with only two color pigments, provides a perfect model for human red-green color blindness. Genetic analysis of the var- ious forms of human color blindness, incidentally, suggests that some humans may someday, millions of years from now, have four cone pigments rather than three and see the world in very different colors than we do now. The modern ﬁeld of vision, encompassing psychophysics, physiology, anatomy, chemistry, and genetics, is one of the great success stories of neuroscience and biological psychol- ogy. We now know that there are more than 30 different visual areas in the cerebral cortex of monkeys and humans, showing degrees of selectivity of response to the various attributes of visual experience, for example, a “color” area, a “movement” area, and so on. We now have a very good un- derstanding of phenomena of visual sensation and perception (see the chapter by Coren in this volume). The ﬁeld con- cerned with vision has become an entirely separate ﬁeld of human endeavor, with its own journals, societies, specialized technologies, and NIH institute. Pitch Detection As we noted, Helmholtz published a most inﬂuential work on hearing in 1863 ( On the Sensation of Tone ). The fundamental issue was how the nervous system codes sound frequency into our sensation of pitch. By this time, much was known about the cochlea, the auditory receptor apparatus. Helmholtz suggested that the basilar membrane in the cochlea func- tioned like a piano, resonating to frequencies according to the length of the ﬁbers. The place on the membrane so activated determined the pitch detected; this view was called the place theory of pitch. The major alternative view was the frequency theory (Rutherford, 1886), in which the basilar membrane was thought to vibrate as a whole due to the frequency of the tone activating it. Boring (1926) presented a comprehen- sive theoretical analysis of these possibilities. One of Boring’s students, E. G. Wever, together with C. W. Bray, recorded from the region of the auditory nerve at the cochlea and found that the recorded electrical signal followed the frequency of the tone up to very high frequencies, many thousands of Hertz (Wever & Bray, 1930). So the frequency theory was vindicated. But there were problems. A single nerve ﬁber cannot ﬁre at much greater than 1,000 Hertz. The attempted answer was the volley theory: Groups of ﬁbers al- ternated in ﬁring to code higher frequencies. Wever and Bray’s discovery is an interesting example of a perfectly good experiment fooled by biology. As it happens, there is a process in the cochlea much like the pizoelectric effect—a tone generates electrical activity at the same fre- quency as the tone, now termed the cochlear microphonic. It is thought to be an epiphenomenon, unrelated to the coding functions of the auditory system. The solution to the question how the cochlea coded tone frequency was provided by Georg von Békésy. Born in Budapest, he received his PhD in physics in 1923 and was a professor at the University of Budapest from 1932 to 1946. In 1947, he accepted a research appointment in the psychology department at Harvard, where he worked until 1964. During his time at Harvard, he was offered a tenured professorship but did not accept it because he disliked formal teaching. Learning and Memory 53 During his years of full-time research at Harvard, he solved the problem of the cochlea, for which he received the Nobel Prize in 1961. In 1964, he accepted a professorship at the University of Hawaii, where he remained until his death. By careful microscopic study of the cochlea, Békésy de- termined the actual movements of the basilar membrane in response to tones (see Békésy, 1947). When William James Hall was built at Harvard to house the psychology depart- ment, a special ﬂoating room was constructed in the base- ment for Békésy’s experiments. The entire room ﬂoated on an air cushion generated by a large air compressor. Further- more, the experimental table ﬂoated within the ﬂoating room on its own compressor. For Békésy’s experiments it was nec- essary to avoid all external building vibrations. (One of the authors, R.F.T., had the opportunity to use this facility when at Harvard.) Békésy discovered that the traveling waves of the basilar membrane induced by a given tone establish a standing wave pattern that maximally displaces a given region for a given tone and different regions for different tones. The pattern of displacement is more complicated than the Helmholtz theory but nonetheless provided a triumph for the place theory. Actually, another kind of physiological evidence provided strong support for the place theory in the 1940s. Woolsey and Walzl (1942) published an extraordinary study in which they electrically stimulated different regions of the auditory nerve ﬁbers in the cochlea (the ﬁbers are laid out along the basilar membrane) in an anesthetized cat and recorded evoked po- tentials in the auditory cortex. The place stimulated on the cochlea determined the region of the auditory cortex acti- vated. An important practical outcome of all this work is the cochlear prosthesis developed for deaf individuals. More recent studies recording the activity of single neu- rons in the auditory cortex have veriﬁed and extended these observations (e.g., Hind et al., 1960). When the ear is stimu- lated with low-intensity pure tones (anesthetized cat), neurons—in particular, narrow dorsal-ventral bands in the primary auditory cortex—respond selectively to tones of dif- ferent frequency. The regions of the cochlea activated by pure tones are represented in an anterior-posterior series of narrow dorsal-ventral bands along the primary auditory cortex, a cochlea-topic representation. Like the visual sciences, the modern ﬁeld of the hearing sciences has become an entirely separate ﬁeld with its own societies, journals, and NIH institute focusing on psy- chophysics and the neurobiology of the auditory system. We know a great deal less about the organization of auditory ﬁelds in the cerebral cortex in primates and humans, inciden- tally, than we do about the visual system. The human auditory areas must be very complex, given our extraordinary species- speciﬁc behavior of speech. LEARNING AND MEMORY Karl Lashley is the most important ﬁgure in the development of physiological psychology and the biology of memory in America. He obtained his PhD at Johns Hopkins University where he studied with John Watson and was heavily inﬂuenced by Watson’s developing notions of behaviorism. While there he also worked with Sheherd Franz at a government hospital in Washington; they published a paper together in 1917 on the ef- fects of cortical lesions on learning and retention in the rat. Lashley then held teaching and research positions at the Uni- versity of Minnesota (1917–1926), the University of Chicago (1929–1935), and at Harvard from 1935 until his death in 1958. During the Harvard years, he spent much of his time at the Yerkes Primate Laboratory in Orange Park, Florida. Lashley devoted many years to an analysis of brain mech- anisms of learning, using the lesion-behavior method, which he developed and elaborated from his work with Franz. Dur- ing this period, Lashley’s theoretical view of learning was heavily inﬂuenced by two congruent ideas—localization of function in neurology and behaviorism in psychology. Lashley describes the origins of his interest in brain sub- strates of memory and Watson’s developing views of behav- iorism in the following letter he wrote to Ernest Hilgard in 1935: In the 1914, I think, Watson called attention of his seminar to the French edition of Bechterev, and that winter the seminar was de- voted to translation and discussion of the book. In the spring I served as a sort of unpaid assistant and we constructed apparatus and planned experiments together. We simply attempted to re- peat Bechterev’s experiments. We worked with withdrawal re- ﬂexes, knee jerk, pupil

bundet

. Watson took the initiative in all this, but he was also trying to photograph the vocal cord, so I did much of the actual experimental work. I devised drainage tubes for the parotid and submaxiallary ducts and planned the salivary work which I published. As we worked with the method, I think our enthusiasm for it was somewhat dampened. Watson tried to es- tablish conditioned auditory reﬂexes in the rat and failed. Our whole program was then disrupted by the move to the lab in Meyer’s clinic. There were no adequate animal quarters there. Watson started work with the infants as the next best material available. I tagged along for awhile, but disliked the babies and found me a rat lab in another building. We accumulated a con- siderable amount of experimental material on the conditioned re- ﬂex which has never been published. Watson saw it as a basis for a systematic psychology and was not greatly concerned with the 54 Biological Psychology nature of the reaction itself. I got interested in the physiology of the reaction and the attempt to trace conditioned reﬂex paths through the nervous system started my program of cerebral work. (Letter of May 14, 1935, K. S. Lashley to E. R. Hilgard, reproduced with the kind permission of E. R. Hilgard) It was in the previous year, 1913, that Watson published his initial salvo in an article entitled “Psychology as the Behav- iorist Views It.” He was elected president of the American Psychological Association in 1914. As we noted earlier, localization of function in the cere- brum was the dominant view of brain organization at the beginning of the twentieth century. In Watson’s behaviorism, the learning of a particular response was held to be the formation of a particular set of connections, a series set. Con- sequently, Lashley argued, it should be possible to localize the place in the cerebral cortex where that learned change in brain organization was stored—the engram. (It was believed at the time that learning occurred in the cerebral cortex.) Thus, behaviorism and localization of function were beauti- fully consistent—they supported the notion of an elaborate and complex switchboard where speciﬁc and localized changes occurred when speciﬁc habits were learned. Lashley set about systematically to ﬁnd these learning locations—the engrams—in a series of studies culminating in his 1929 monograph, Brain Mechanisms of Intelligence . In this study, he used mazes differing in difﬁculty and made lesions of varying sizes in all different regions of the cerebral cortex of the rat. The results of this study profoundly altered Lashley’s view of brain organization and had an extraordi- nary impact on the young ﬁeld of physiological psychology. The locus of the lesions is unimportant; the size is critically important, particularly for the more difﬁcult mazes. These ﬁndings led to Lashley’s two theoretical notions of equipo- tentiality and mass action: that is, all areas of the cerebral cor- tex are equally important (at least in maze learning), and what is critical is the amount of brain tissue removed. Lashley’s interpretations stirred vigorous debate in the ﬁeld. Walter Hunter, an important ﬁgure in physiological- experimental psychology at Brown University who devel- oped the delayed response task in 1913, argued that in fact the rat was using a variety of sensory cues; as more of the sensory regions of the cortex were destroyed, fewer and fewer cues became available. Lashley and his associates countered by showing that removing the eyes has much less effect on maze learning than removing the visual area of the cortex. Others argued that Lashley removed more than the vi- sual cortex. Out of this came a long series of lesion-behavior studies analyzing behavioral “functions” of the cerebral cor- tex. Beginning in the 1940s, several laboratories, including Lashley’s and those of Harry Harlow at the University of Wisconsin and Karl Pribram at Yale, took up the search for the more complex functions of association cortex using mon- keys and humans. Perhaps the most important single discovery in this ﬁeld came from Brenda Milner’s studies with patient H. M. who, following bilateral temporal lobectomy (removing the hip- pocampus and other structures), lives forever in the present. Work on higher brain functions in monkeys and humans is one of the key roots of modern cognitive neuroscience, to be treated later. Since Milner’s work with H. M., the hippocam- pus has been of particular interest in biological psychology. Another facet of hippocampal study in the context of the biological psychology of memory is long-term potentiation (LTP), discovered by Bliss and Lomo (1973). Brief tetanic stimulation of monosynaptic inputs to the hippocampus causes a profound increase in synaptic excitability that can persist for hours or days. Many view it as a leading candidate for a mechanism of memory storage, although direct evi- dence is still lacking. Yet another impetus to study of the hippocampus in the re- markable discovery of “place cells” by John O’Keefe (1979). When recording from single neurons in the hippocampus of the behaving rat, a give neuron may respond only when the animal is in a particular place in the environment (i.e., in a box or maze), reliably and repeatedly. There is great interest now in the possibility that LTP may be the mechanism form- ing place cells. A number of laboratories are making use of genetically altered mice to test this possibility. Lashley’s inﬂuence is felt strongly through the many emi- nent physiological psychologists who worked or had contact with him. We select two examples here—Austin Riesen and Donald O. Hebb. We discuss Roger W. Sperry’s work next in the context of cognitive neuroscience. The basic problem of the development of perception fascinated Lashley and his students. How is it that we come to perceive the world as we do? Do we learn from experience or is it told to us by the brain? Riesen did pioneering studies in which he raised mon- keys for periods of time in the dark and then tested their vi- sual perception. They were clearly deﬁcient. This important work served as one of the stimuli for Hebb to develop a new theory of brain organization and function, which he outlined in The Organization of Behavior (1949). This book had an immediate and profound impact on the ﬁeld. Hebb effectively challenged many traditional notions of brain organization and attempted to pull together several dis- cordant themes—mass action and equipotentiality, effects of dark rearing on perception, the preorganization of sensory cortex, the lack of serious intellectual effects of removal of an entire hemisphere of the brain in a human child—into a Learning and Memory 55 coherent theory. Important inﬂuences of Gestalt notions can be seen in Hebb’s theory. He is a connectionist but in a mod- ern sense: Connections must underlie brain organization but there is no need for them to be in series. One concept in Hebb’s book has come to loom large (too large perhaps) in modern cognitive-computational neuro- science—the Hebb synapse: When an axon of Cell A is near enough to excite a cell B and re- peatedly or persistently takes part in ﬁring it, some growth process or metabolic change takes place in one or both cells such that A’s efﬁciency, as one of the cells ﬁring B, is increased. (1949, p. 62) Lashley’s pessimistic conclusions in his 1929 monograph put a real but temporary damper on the ﬁeld concerned with brain substrates of memory. But other major traditions were developing. Perhaps the most important of these was the in- ﬂuence of Pavlov. His writings were not readily available to Western scientists, particularly Americans, until the publica- tion of the English translation of his monumental work Con- ditioned Reﬂexes in 1927. It is probably fair to say this is the most important single book ever published in the ﬁeld of be- havioral neuroscience. Pavlov developed a vast and coherent body of empirical results characterizing the phenomena of conditioned responses, what he termed “psychic reﬂexes.” He argued that the mind could be fully understood by analy- sis of the higher order learned reﬂexes and their brain sub- strates. As an example of his inﬂuence, Clark Hull, in his Principles of Behavior (1943), wrote as though he were a student of Pavlov. W. Horsley Gantt, an American physician, worked with Pavlov for several years and then established a Pavlovian laboratory at Johns Hopkins. He trained several young psy- chologists, including Roger Loucks and Wulf Brogden, who became very inﬂuential in the ﬁeld. Perhaps the most impor- tant modern behavioral analyses of Pavlovian conditioning are the works of Robert Rescorla and Allan Wagner (1972). Although Pavlov worked with salivary secretion, most studies of classical conditioning in the West tended to utilize skeletal muscle response, à la Bechterev. Particularly pro- ductive have been Pavlovian conditioning of discrete skeletal reﬂexes (e.g., the eyeblink response), characterized behaviorally by Isadore Gormezano and Allan Wagner and analyzed neuronally by Richard Thompson and his many stu- dents, showing localization of the basic memory trace to the cerebellum (Thompson, 1986). Masao Ito and associates in Tokyo had discovered the phenomenon of long-term depres- sion (LTD) in the cerebellar cortex (see Ito, 1984). Repeated conjunctive stimulation of the two major inputs to the cerebellum, mossy-parallel ﬁbers and climbing ﬁbers, yields a long-lasting decrease in the excitability of parallel ﬁbers— Purkinje neuron synapses. Ito developed considerable evi- dence that this cerebellar process underlies plasticity of the vestibular-ocular reﬂex. Thompson and associates developed evidence, particularly using genetically altered mice, that cerebellar cortical LTD is one of the mechanisms underly- ing classical conditioning of eyeblink and other discrete responses. Fear conditioning was characterized behaviorally by Neal Miller and analyzed neuronally by several groups, particu- larly Michael Davis (1992), Joseph LeDoux (2000), and Michael Fanselow (1994), and their many students. They showed that at least for classical conditioning of fear, the es- sential structure is the amygdala, which may contain the basic memory trace for this form of learning (but see just below). The process of LTP may serve to code the amygdalar fear memory. Duncan’s discovery in 1949 of the effects of electrocon- vulsive shock on retention of simple habits in the rat began the modern ﬁeld of memory consolidation. Hebb and Gerard were quick to point out the implication of two memory processes, one transient and fragile and the other more per- manent and impervious. James McGaugh and his associates (1989) have done the classic work on the psychobiology of memory consolidation. He and his colleagues demonstrated memory facilitation with drugs and showed that these effects were direct and not due to possible reinforcement effects of the drugs (and similarly for ECS impairment). The amygdala is critical for instrumental learning of fear. McGaugh and his associates demonstrated that for both pas- sive and active avoidance learning (animals must either not respond, or respond quickly, to avoid shock) amygdala le- sions made immediately after training abolished the learned fear. Surprisingly, if these same lesions were made a week after training, learned fear was not abolished, consistent with a process of consolidation (see McGaugh, 2000). The appar- ent difference in the role of the amygdala in classical and in- strumental learning of fear is a major area of research today. Chemical approaches to learning and memory are recent. The possibility that protein molecules and RNA might serve to code memory was suggested some years ago by pioneers such as Gerard and Halstead. The RNA hypothesis was taken up by Hyden and associates in Sweden and by several groups in America. An unfortunate by-product of this approach was the “transfer of memory” by RNA. These experiments, done by investigators who shall remain nameless, in the end could not be replicated. At the same time, several very productive lines of investi- gation of neurochemical and neuroanatomical substrates of 56 Biological Psychology learning were developing. In 1953, Krech and Rosenzweig began a collaborative study of relationships between brain chemistry and behavior. Krech didclassic earlywork inanimal learning (under his earlier name, Kreshevsky) and was a col- league of and collaborator with Tolman. Mark Rosenzweig re- ceived his PhD in physiological psychology at Harvard in 1949 and joined the psychology department at the University of California, Berkeley, in 1951. Soon after they began their joint work in 1953 they were joined by E. L. Bennett and later by M. C. Diamond. Their initial studies concerned brain levels of AChE in relation to the hypothesis behavior and included analysis of strain differences (see Krech, Rosenzweig, & Bennett, 1960). More recently, they discovered the striking differences in the brains of rats raised in “rich” versus “poor” environments.William Greenough (1984), at the University of Illinois, replicated and extended this work to demonstrate dra- matic morphological changes in the structures of synapses and neurons as a result of experience. The use of model biological systems has been an impor- tant tradition in the study of neural mechanisms of learning. This approach has been particularly successful in the analysis of habituation, itself a very simple form or model of learning. Sherrington did important work on ﬂexion reﬂex “fatigue” in the spinal animal at the turn of the century. In 1936, Prosser and Hunter completed a pioneering study comparing habitu- ation of startle response in intact rats and habituation of hindlimb ﬂexion reﬂex in spinal rats. They established, for habituation, the basic approach of Sherrington, namely that spinal reﬂexes can serve as models of neural-behavioral processes in intact animals. Sharpless and Jasper (1956) es- tablished habituation as an important process in EEG activity. Modern Russian inﬂuences have been important in this ﬁeld—the key studies of Evgeny Sokolov (1963), ﬁrst on habituation of the orienting response in humans and more re- cently on mechanisms of habituation of responses in the sim- pliﬁed nervous system of the snail. The deﬁning properties of habituation were clearly estab- lished by Thompson and Spencer in 1966, and the analysis of mechanisms began. Several laboratories using different preparations— Aplysia withdrawal reﬂex; Kandel and his many associates (see Kandel, 1976); vertebrate spinal re- ﬂexes; Thompson, Spencer, Farel; crayﬁsh tail ﬂip escape; Krasne (1969), Kennedy—all arrived at the same underlying synaptic mechanism—a decrease in the probability of trans- mitter release from presynaptic terminals of the habituating pathway. Habituation is thus a very satisfying ﬁeld; agree- ment ranges from deﬁning behavioral properties to synaptic mechanisms. In a sense, the problem has been solved. Habituation also provides a most successful example of the use of the model biological systems approach to analysis of neural mechanisms of behavioral plasticity (see Groves & Thompson, 1970). Special mention must be made of the elegant and detailed studies by Eric Kandel and his many associates on long- lasting neuronal plasticity in the Aplysia gill-withdrawal circuit (Kandel, 1976; Hawkins, Kandel, & Siegelbaum, 1993). This simpliﬁed model system (together with work on the hippocampus) made it possible to elucidate putative processes that result in long-lasting synaptic plasticity, for example, biochemical models of memory formation and stor- age. Eric was awarded the Nobel Prize for Physiology and Medicine in 2000 in part for this work. MOTIVATION AND EMOTION Physiological and neural mechanisms of motivation and emotion have been a particular province of biological psy- chology and physiology in the twentieth century. In more re- cent years, the ﬁelds of motivation and emotion have tended to go separate ways (see Brown, 1961, 1979). However mo- tivation and emotion have common historical origins. In the seventeenth and eighteenth centuries, instinct doctrine served as the explanation for why organisms were driven to behave (at least infrahuman organisms without souls). Darwin’s emphasis on the role of adaptive behavior in evolutionary survival resulted in the extension of instinct doctrine to human behavior. Major sources of impetus for this were Freud’s and McDougall’s notions of instinctive human moti- vation. Watson rebelled violently against the notion of in- stinct and rejected it out of hand, together with all biological mechanisms of motivation. As Lashley (1938) put it, he “threw out the baby with the bath.” Emotion The dominant theory of emotion in the ﬁrst two decades of the century was that of James and Lange—“We feel afraid because we run” (see James, 1884). Actually, James focused more on the subjective experience of emotion, and Lange, a Danish anatomist, focused on the physiological phenomena, believing that subjective experience is not a proper topic for science. But between them they developed a comprehensive theory of emotion. The basic idea is that we ﬁrst perceive an emotionally arousing situation or stimulus (“a bear in the woods” is a favorite example), which leads to bodily (physi- ological) changes and activities, which result in the experi- enced emotion. This general view was challenged by the American physi- ologist Walter B. Cannon in the 1920s and 1930s. He actually Motivation and Emotion 57 agreed with James and Lange that the initial event had to be perception of an emotion-arousing situation but argued that the development of autonomic (sympathetic) responses— release of epinephrine and other bodily changes—occurred concomitantly with the subjective feelings (see Cannon, 1927). However, his primary interest was in the physiology, particularly the peripheral physiology. Cannon’s view was championed by the distinguished Johns Hopkins physiologist Philip Bard, who stressed the key role of the brain, particu- larly the thalamus and hypothalamus, in both emotional behavior and experience (see Bard, 1934). Cannon, inciden- tally, also contributed the notion of homeostasis, which he developed from Bernard’s concept of the milieu interieur. A key issue in these theories was the role of sympathetic arousal or activation in the experience of emotion. This issue was tested in a classic study by Stanley Schachter and Jerome Singer at Columbia University in 1962. They injected human subjects with either effective doses of epinephrine or a placebo. The epinephrine activated the sympathetic signs of emotions (pounding heart, dry mouth, etc.). Both groups of subjects were told they were receiving a shot of a new vita- min. Stooges acted out euphoria or anger in front of the sub- jects. The subjects were either informed of what the injection might do, for example, the autonomic side effects, or not in- formed. Results were dramatic. Uninformed epinephrine subjects reported emotional experiences like those of stooges but informed epinephrine subjects did not report any emotion at all. Emotion is more than sympathetic arousal—cognitive factors are also important. Experimental work on brain substrates of emotion may be said to have begun with the studies of Karplus and Kreidl in 1910 on the effects of stimulating the hypothalamus. In 1928, Bard showed that the hypothalamus was responsible for “sham rage.” In the 1930s, S. W. Ranson and his associates at Northwestern, particularly H. W. Magoun, published a clas- sic series of papers in the hypothalamus and its role in emo- tional behavior (Ranson & Magoun, 1939). In the same period, W. R. Hess (1957) and his collaborators in Switzer- land were studying the effects of stimulating the hypothala- mus in freely moving cats. A most important paper by H. Klüver and P. Bucy reported on “psychic blindness and other symptoms following bilateral temporal lobectomy in rhesus monkeys” in 1937. This came to be known as the Klüver- Bucy syndrome. The animals exhibited marked changes in motivation and aggressive behavior. Pribram (Bucy’s ﬁrst resident in neurosurgery) developed the surgical methods necessary to analyze the Klüver-Bucy syndrome.This analysis led to his discovery of the functions of theinferotemporal cortex invision and tothe exploration of the suggestions of J. W. Papez (1937) and P. D. MacLean (1949) thatthe structures ofthe limbic system(the “Papez” circuit)are concerned with motivation and emotion. However, modern neuroanatomy deconstructed the Papez circuit. The emphasis is now on the hypothalamus-pituitary axis, on descending neural systems, and on the amygdala. Motivation Today most workers in the ﬁeld prefer the term motivated behaviors to emphasize the speciﬁc features of behaviors re- lating to hunger, thirst, sex, temperature, and so forth. Karl Lashley was again a prime mover. His 1938 paper, “Experi- mental Analysis of Instinctive Behavior,” was the key. He ar- gued that motivated behavior varies and is not simply a chain of instinctive or reﬂex acts, is not dependent on any one stim- ulus, and involves central state. His conclusions, that “physi- ologically, all drives are no more than expression of the activity of speciﬁc mechanisms” and that hormones “activate some central mechanism which maintains excitability and ac- tivity,” have a very modern ring. Several key ﬁgures in the modern development of the psychobiology of motivation are Clifford Morgan, Eliot Stellar, Kurt Richter, Frank Beach, Neal Miller, Philip Teitelbaum, and James Olds. Morgan went to graduate school at Rochester, where his professors included E. A. K. Culler and K. U. Smith and his fellow graduate students in- cluded D. Neff, J. C. R. Licklider, and P. Fitts. He then be- came an instructor at Harvard, where he ﬁrst worked in Lashley’s laboratory in 1939. He later moved to Johns Hop- kins, where he remained until 1959. As a graduate student and later at Harvard, Morgan came to doubt Cannon’s then current notion that hunger was the result of stomach con- tractions. Morgan did a series of studies showing this could not be a complete or even satisfactory account of hunger and feeding behavior. Eliot Stellar and Robert McCleary, then undergraduates at Harvard, worked with Morgan. They focused on hoarding behavior and completed a classic analysis of the internal and environmental factors control- ling the behavior. Lashley’s general notion of a central mechanism that maintains activity was developed by Beach in an important series of papers in the 1940s and by Morgan in the ﬁrst edi- tion of his important text, Physiological Psychology (1943), into a central excitatory mechanism and ultimately a central theory of drive. This view was given a solid physiological basis by Donald B. Lindsley from the work he and H. W. Magoun, G. Moruzzi, and associates were doing on the as- cending reticular activating system. Lindsley sketched his ac- tivation theory of emotion in his important chapter in the Stevens Handbook (1951). Hebb (1955) and Stellar (1954) 58 Biological Psychology pulled all these threads together into a general central theory of motivation. Eliot Stellar worked with Clifford Morgan as an under- graduate at Harvard. After obtaining his doctorate in 1947 at Brown University, he spent several years at Johns Hopkins and joined the psychology department at the University of Pennsylvania in 1954. Stellar did extensive work on brain mechanism of motivation. He coauthored the revision of Morgan’s text in 1950 and published his inﬂuential central theory of drive in 1954. Philip Teitelbaum (1955) did the classic work on charac- terization of, and recovery from, the lateral hypothalamic “aphagia” syndrome. He discovered the striking parallel with the ontogenetic development of feeding behavior. In addi- tion, he discovered more general aspects of the syndrome, for example, “sensory neglect.” Frank Beach received his doctorate from the University of Chicago under Lashley in 1940 and then joined the American Museum of Natural History in New York. He moved to Yale in 1946, and then to the University of California, Berkeley, in 1958. From the beginning, he focused on brain mechanisms of sexual behavior (see Beach, 1951). As the study of sexual behavior developed, hormonal factors came to the fore and the modern ﬁeld of hormones and behavior developed. Beach played a critical role in the development of this ﬁeld, as did the biologist W. C. Young of the University of Kansas. They and their students shaped the ﬁeld as it exists today. Even within the ﬁeld of hormones and behavior, several ﬁelds have developed. Sexual behavior has become a ﬁeld unto itself. Another important ﬁeld is the general area of stress. The endocrinologist Hans Selye was an important in- tellectual inﬂuence. Kurt Richter, a pioneering ﬁgure in this ﬁeld, took his BS at Harvard in 1917 and his doctorate at Johns Hopkins in 1921 and was a dominant inﬂuence at Hopkins. His early work was on motivation and feeding (see Richter, 1927). His pioneering “cafeteria studies” in rats are still a model (if given a wide choice of foods, they select a relatively balanced diet). Richter then focused on the adrenal gland, its role in diet and in stress. He also did pioneering work on circadian rhythms in mammals. The modern ﬁeld of stress focuses on hormonal-behavioral interactions, particu- larly adrenal hormones, as in the work of Seymore Levine (1971). Neal Miller represents a uniquely important tradition in biological psychology. From the beginning of his career, Miller was interested in physiological mechanisms of both motivation and learning. He took his doctorate at Yale in 1935 and stayed on at Yale for many years, with a year out in 1936 at the Vienna Psychoanalytic Institute. Throughout his career he has exempliﬁed superb experimentation and an unusual ability to synthesize. He was a pioneer in early stud- ies of punishing and rewarding brain stimulation and their roles in learning and in the study of conditioned fear (see Miller, 1948, 1961). In later years, his work focused on mechanisms of instrumental conditioning of autonomic responses—biofeedback techniques—and brain mechanisms of learning. The impact of his work is much wider than bio- logical psychology, inﬂuencing learning theory, psychiatry, and clinical medicine as well. James Olds, whose untimely death in 1976 cut short an ex- traordinary career, made the most important discovery yet in the ﬁeld of motivation—rewarding electrical self-stimulation of the brain. He got his doctorate at Harvard and worked with Richard Solomon. Solomon, although primarily a behavioral student of learning, had considerable impact on biological psychology through his theoretical-experimental analysis of hypothetical central factors in learning. As a graduate student Olds read and was much inﬂuenced by Hebb’s Organization of Behavior and obtained a postdoctoral fellowship with Hebb at McGill in 1953. He began work there with Peter Milner. In his own words: Just before we began our own work (using Hess’s technique for probing the brain), H. R. Delgado, W. W. Roberts, and N. E. Miller at Yale University had undertaken a similar study. They had located an area in the lower part of the mid-line system where stimulation caused the animal to avoid the behavior that provoked the electrical stimulus. We wished to investigate posi- tive as well as negative effects (that is, to learn whether stimula- tion of some areas might be sought rather than avoided by the animal). We were not at ﬁrst concerned to hit very speciﬁc points in the brain, and, in fact, in our early tests the electrodes did not al- ways go to the particular areas in the mid-line system at which they were aimed. Our lack of aim turned out to be a fortunate happening for us. In one animal the electrode missed its target and landed not in the mid-brain reticular system but in a nerve pathway from the rhinecephalon. This led to an unexpected discovery. In the test experiment we were using, the animal was placed in a large box with corners labeled A, B, C, and D. Whenever the animal went to corner A, its brain was given a mild electric shock by the experimenter. When the test was performed on the animal with the electrode in the rhinencephalic nerve, it kept returning to corner A. After several such returns on the ﬁrst day, it ﬁnally went to a different place and fell asleep. The next day, however, it seemed even more interested in corner A. At this point we assumed that the stimulus must provoke curiosity; we did not yet think of it as a reward. Further exper- imentation on the same animal soon indicated, to our surprise, that its response to the stimulus was more than curiosity. On the second day, after the animal had acquired the habit of returning Cognitive Neuroscience 59 to corner A to be stimulated, we began trying to draw it away to corner B, giving it an electric shock whenever it took a step in that direction. Within a matter of ﬁve minutes the animal was in corner B. After this the animal could be directed to almost any spot in the box at the will of the experimenter. Every step in the right direction was paid with a small shock; on arrival at the appointed place the animal received a longer series of shocks. After conﬁrming this powerful effect of stimulation of brain areas by experiments with a series of animals, we set out to map the places in the brain where such an effect could be obtained. We wanted to measure the strength of the effect in each place. Here Skinner’s technique provided the means. By putting the an- imal in the “do-it-yourself” situation (i.e., pressing a lever to stimulate its own brain) we could translate the animal’s strength of “desire” into response frequency, which can be seen and measured. The ﬁrst animal in the Skinner box ended all doubts in our minds that electric stimulation applied to some parts of the brain could indeed provide a reward for behavior. The test displayed the phenomenon in bold relief where anyone who wanted to look could see it. Left to itself in the apparatus, the animal (after about two to ﬁve minutes of learning) stimulated its own brain regu- larly about once very ﬁve seconds, taking a stimulus of a second or so every time. (1956, pp. 107–108) We think now that this brain reward circuit Olds discov- ered underlies addictive behaviors. It includes the medial forebrain bundle (MRB) containing the ascending dopamine (and other neurotransmitters) projection system to the nu- cleus accumbens and prefrontal cortex. Activation of this sys- tem appears to be a common element in what keeps drug users taking drugs. This activity is not unique to any one drug; all addictive drugs affect this circuit. Another direction of research in motivation and emotion relating to brain stimulation concerns elicited behaviors, par- ticularly from stimulation in the region of the hypothalamus. This work is in some ways a continuation of the early work by Hess. Thus, Hess described directed attack, from hypo- thalamic stimulation in cats, as opposed to the “sham” rage of decerebrate animals and certain other brain stimulation stud- ies (“sham” because the animal exhibited peripheral signs of rage without integrated behavior) (see Hess, 1957). John Flynn, in a most important series of studies, was able to elicit two quite different forms of attack behavior in cats—one a quiet predation that resembled normal hunting and the other a rage attack (Flynn, Vonegas, Foote, & Edwards, 1970). Elliot Valenstein analyzed a variety of elicited consumatory-like behaviors—eating, drinking, gnawing, and so forth—from hypothalamic stimulation and their possible relations to the rewarding properties of such stimulation (Valenstein, Cox, & Kakolweski, 1970). Current focus in the study of motivated behaviors is on de- tailed physiological processes, particularly involving mecha- nisms of gene expression of various peptide hormones in the hypothalamus and their actions on the pituitary gland, and on descending neural systems from the hypothalamus that act on lower brain systems to generate motivated behaviors (see e.g., Swanson, 1991). But we still do not understand the neural circuitries underlying the fact that seeing the bear in the woods makes us afraid. COGNITIVE NEUROSCIENCE The term cognitive neuroscience is very recent, dating per- haps from the 1980s. The Journal of Cognitive Neuroscience was ﬁrst published in 1989. Indeed, Posner and Shulman’s comprehensive chapter on the history of cognitive sci- ence (1979) does not even mention cognitive neuroscience (human imaging techniques were not yet much in use then). The cognitive revolution in psychology is treated in the chap- ter by Leahey in this volume. Here we note brieﬂy the bio- logical roots of cognitive neuroscience (see Gazzaniga, 1995). Karl Lashley was again a key ﬁgure. One of the most im- portant aspects of cognitive neuroscience dates from the early days at the Orange Park laboratory, where young scien- tists like Chow and Pribram began studies of the roles of the association areas of the monkey cerebral cortex in learning, memory, and cognition. The 1950s was an especially rich time of discovery re- garding how cognitive function was organized in the brain. Pribram, Mortimer Mishkin, and Hal Rosvold at NIMH, using lesion studies in monkeys, discovered that the temporal lobe was critical for aspects of visual perception and mem- ory. Work with neurologic patients also played a critical role in uncovering the neural substrates of cognition. One partic- ular discovery became a landmark in the history of memory research. “In 1954 Scoville described a grave loss of recent memory which he had observed as a sequel to bilateral medial temporal resection in one psychotic patient and one patient with intractable seizures. In both case s… removals extended posteriorly along the medial surface of the temporal lobe s…and probably destroyed the anterior two-thirds of the hippocampus and hippocampal gyrus bilaterally, as well as the uncus and amygdala. The unexpected and persistent memory deﬁcit which resulted seemed to us to merit further investigation.” That passage comes from the ﬁrst paragraph of Scoville and Milner’s 1957 report, “Loss of Recent Memory after Bilateral Hippocampal Lesions.” This publication became a 60 Biological Psychology landmark in the history of memory research for two reasons. First, the severe memory impairment (or amnesia) could be linked directly to the brain tissue that had been removed, sug- gesting that the medial aspect of the temporal lobe was an important region for a particular aspect of cognition, that is, memory function. Second, comprehensive testing of one of the patients (H. M.) indicated that memory impairment could occur on a background of otherwise normal cognition. This observation showed that memory is an isolatable function, separable from perception and other cognitive and intellec- tual functions. The ﬁndings from patient H. M. (Scoville & Milner, 1957) identiﬁed a region of the brain important for human memory, that is, the medial portion of the temporal lobe. The damage was originally reported to have included the amygdala, the periamygdaloid cortex (referred to as the uncus in Scoville & Milner, 1957), the hippocampal region (referred to as the hippocampus), and the perirhinal, entorhinal, and parahip- pocampal cortices (referred to as the hippocampal gyrus). Recently, magnetic resonance imaging of patient H. M. has shown that his medial temporal lobe damage does not extend as far posteriorly as originally believed and that damage to the parahippocampal cortex is minimal (the lesion extends caudally from the temporal pole approximately 5 cm, instead of 8 cm, as originally reported; Corkin, Amaral, Gonzalez, Johnson, & Hyman, 1997). While these observations identiﬁed the medial temporal lobe as important for memory, the medial temporal lobe is a large region including many different structures. To deter- mine which structures are important required that studies be undertaken in which the effects of damage to medial tempo- ral lobe structures could be evaluated systematically. Accord- ingly, soon after the ﬁndings from H. M. were reported, efforts were made to develop an animal model of medial tem- poral lobe amnesia. During the next 20 years, however, ﬁnd- ings from experimental animals with intended hippocampal lesions or larger lesions of the medial temporal lobe were inconsistent and difﬁcult to interpret. In 1978, Mishkin introduced a method for testing memory in monkeys that captured an important feature of tests sensi- tive to human memory impairment (Mishkin, 1978). This method allowed for the testing of memory for single events at some delay after the event occurred. The task itself is known as the trial-unique delayed-nonmatching-to-sample task, and it measures object recognition memory. In Mishkin’s study, three monkeys sustained large medial temporal lobe lesions that were intended to reproduce the damage in patient H. M. The operated monkeys and three unoperated monkeys were given the delayed-nonmatching-to-sample task in order to as- sess their ability to remember, after delays ranging from eight seconds to two minutes, which one of two objects they had re- cently seen. The monkeys with medial temporal lobe lesions were severely impaired on the nonmatching task, consistent with the severe impairment observed in patient H. M. on delay tasks. Thus, lesions that included the hippocampal region, the amygdala, as well as adjacent perirhinal, entorhinal, and parahippocampal cortices caused severe memory impairment. This work, together with work carried out in the succeeding few years, established a model of human amnesia in nonhu- man primates (Mishkin, Spiegler, & Saunders, 1982; Squire & Zola-Morgan, 1983). Although other tasks have been useful for measuring memory in monkeys (object discrimination learning, the visual paired-comparison task; see below), much of the information about the effects of damage to medial tem- poral lobe structures has come, until recently, from the delayed-nonmatching-to-sample task. Once the animal model was established, systematic and cu- mulative work eventually identiﬁed the structures in the me- dial temporal lobe that are important for memory. The important structures are the hippocampal region and the ad- jacent perirhinal, entorhinal, and parahippocampal cortices (for reviews, see Mishkin & Murray, 1994; Zola-Morgan & Squire, 1993). The amygdala proved not to be a component of this memory system, although it can exert a modulatory action on the kind of memory that depends on the medial tem- poral lobe system (Cahill & McGaugh, 1998). The medial temporal lobe is necessary for establishing one kind of memory, what is termed long-term declarative or ex- plicit memory. Declarative memory refers to the capacity for conscious recollection of facts and events (Squire, 1992). It is specialized for rapid, even one-trial learning, and for forming conjunctions between arbitrarily different stimuli. It is typically assessed in humans by tests of recall, recognition, or cued recall, and it is typically assessed in monkeys by tests of recognition (e.g., the delayed-nonmatching-to-sample task). The medial temporal lobe memory system appears to perform a critical function beginning at the time of learn- ing in order that representations can be established in long- term memory in an enduring and usable form (see also Eichenbaum, Otto, & Cohen, 1994). Another important discovery that paralleled in time the work on the medial temporal lobe system involved the un- derstanding that there is more than one kind of memory. Speciﬁcally, work with amnesic patients and with experi- mental animals who sustained lesions to speciﬁc brain regions showed that other kinds of abilities (including skills, habit learning, simple forms of conditioning, and the phe- nomenon of priming, which are collectively referred to as nondeclarative memory) lie outside the province of the me- dial temporal lobe memory system. Nondeclarative forms of Cognitive Neuroscience 61 memory are intact in amnesic patients and intact in monkeys with medial temporal lobe lesions. For example, classical delay conditioning of skeletal musculature depends on the cerebellum (Thompson & Krupa, 1994), conditioning of emotional responses depends on the amygdala (Davis, 1992; LeDoux, 2000), and habit learning (win-stay, lose-shift re- sponding) depends on the neostriatum (Packard, Hirsh, & White, 1989; Salmon & Butters, 1995). Nondeclarative memory thus refers to a variety of ways in which experience can lead to altered dispositions, preferences, and judgments without providing any conscious memory content. Further work with monkeys has demonstrated that the severity of memory impairment depends on the locus and extent of damage within the medial temporal lobe memory system. Damage limited to the hippocampal region causes signiﬁcant memory impairment, but damage to the adjacent cortex increases the severity of memory impairment. It is im- portant to note that the discovery that larger medial temporal lobe lesions produce more severe amnesia than smaller le- sions is compatible with the idea that structures within the medial temporal lobe might make qualitatively different con- tributions to memory function. This is because anatomical projections carrying information from different parts of the neocortex enter the medial temporal lobe memory system at different points (Suzuki & Amaral, 1994). Another important brain area for memory is the dien- cephalon. However, the critical regions in the diencephalon that when damaged produce amnesia have not at the time of writing been identiﬁed with certainty. The important struc- tures appear to include the mediodorsal thalamic nucleus, the anterior nucleus, the internal medullary lamina, the mammillo-thalamic tract, and the mammillary nuclei. Be- cause diencephalic amnesia resembles medial temporal lobe amnesia in many ways, these two regions together probably form an anatomically linked, functional system. These ﬁndings in monkeys are fully consistent with the ﬁndings from human amnesia. Damage limited to the hip- pocampal region is associated with moderately severe amne- sia (Rempel-Clower, Zola, & Squire, 1996; Zola-Morgan, Squire, Rempel, Clower, & Amarel, 1992), and more exten- sive damage that includes the hippocampal region as well as adjacent cortical regions is associated with more severe memory impairment (Corkin, 1984; Mishkin, 1978; Rempel- Clower et al., 1996; Scoville & Milner, 1957). The same principle, that more extensive damage produces more severe impairment, has also been established for the hippocampus proper in the case of the rat (E. Moser, Moser, & Andersen, 1993; M. Moser, Moser, & Forrest, 1995). The dorsal hippocampus of the rat is essential for spatial learning in the water maze, and progressively larger lesions of this region produce a correspondingly larger impairment. Thus, in all three species it has turned out that the brain is organized such that memory is a distinct and separate cognitive func- tion, which can be studied in isolation from perception and other intellectual abilities. Information is still accumulating about how memory is organized, what structures and connec- tions are involved, and what functions they support. The dis- ciplines of both psychology and neuroscience continue to contribute to this enterprise. Roger Sperry was another key player in the origins of cog- nitive neuroscience. He received his doctorate in zoology at the University of Chicago and then joined Lashley for a year at Harvard and moved with Lashley to the Yerkes Primate Laboratory at Orange Park, where he stayed for some years. Sperry did his pioneering studies on the selective growth of brain connections during this time (see Sperry, 1951). Lashley was fascinated by the mind–brain issue—the brain substrates of consciousness (although he never wrote about it)—and often discussed this problem with his younger col- leagues at Orange Park (Sperry, personal communication). In more recent years, Sperry and his associates at the California Institute of Technology tackled the issue with a series of com- missurotomy patients—the human “split-brain” studies. This work proved to be extraordinary, perhaps the most important advance in the study of consciousness since the word itself was developed many thousands of years ago (Sperry, 1968). Another key origin of the modern ﬁeld of cognitive neuro- science is the study of humans with brain damage, as in Milner’s work on H. M. noted earlier. Other inﬂuential scien- tists in the development of this ﬁeld were Hans-Lukas Teuber and Brenda Milner. Karl Pribram also played a critical role. Teuber received his early training at the University of Basel, obtained his doctorate at Harvard, and studied with Karl Lashley. He became chairman of the psychology department at MIT in 1961. In the 1940s, he published an important se- ries of papers in collaboration with Bender and others on per- ceptual deﬁcits following penetrating gunshot wounds of the brain. Later he also investigated the effects of frontal lesions on complex performance in humans. Brenda Milner received her undergraduate training at Cambridge; then after the war she came to Canada and stud- ied for her PhD at McGill University under Hebb’s supervi- sion. Hebb arranged for her to work with Wilder Penﬁeld’s neurosurgical patients at the Montreal Neurological Institute. Her work on temporal lobe removal in humans, including H. M., really began modern study of the memorial functions of the hippocampus (see earlier). She also collaborated on studies with Roger Sperry and with Karl Pribram. Another very important inﬂuence in modern cognitive neuroscience comes from the Soviet scientist Alexander 62 Biological Psychology Luria, who died in 1977. Luria approached detection and evaluation of damage to higher regions of the human brain both as a clinician with extraordinary expertise in neurology and as a scientist interested in higher functions of the nervous system (e.g., his book Language and Cognition, 1981). Yet another origin of cognitive neuroscience is recording the activity of the human brain, initially using the EEG. Donald Lindsley was a pioneer in this work. Lindsley did his graduate work at Iowa and worked with L. E. Travis, himself an important ﬁgure in psychophysiological record- ing. Lindsley then took a three-year postdoctoral at Har- vard Medical School (1933–1935). The neurophysiologist Alexander Forbes was at Harvard doing pioneering studies on brain-evoked potentials and EEG in animals. The ﬁrst human EEG recording laboratory was set up at Harvard, and Lindsley and other pioneering ﬁgures such as Hallowell Davis did the ﬁrst EEG recording in America (Lindsley, 1936). More recently, the method of averaging evoked potentials recorded from the human scalp made it possible to detect brain signals relevant to behavioral phenomena that could not be detected with individual trial recording. Donald Lindsley was a pioneer in this ﬁeld as well, doing early studies on evoked potential correlates of attention. E. Roy John and oth- ers developed complex, comprehensive methods of quantita- tive analysis of EEG and evoked potential recordings. But the techniques that have revolutionized the study of normal human brain organization and functions are of course the methods of imaging. The ﬁrst such method was X-ray- computed tomography, developed in the early 1970s. The major innovation beyond simple X rays was complex mathe- matical and computer techniques to reconstruct the images. Somewhat later, positron emission tomography (PET) was developed. It is actually based on a long used method in animal neuroanatomy—autoradiography. In this technique, a radioactive substance that binds to a particular type of mole- cule or brain region is infused and brain sections are prepared and exposed to X-ray ﬁlm. For humans PET involves inject- ing radioactive substances, for example, radiolabeled oxygen ( 15 O), in water. Increased neuronal activity in particular re- gions of the brain causes a rapid increase in blood ﬂow to the regions, as shown years earlier in work by Seymore Kety and others. Consequently, the radioactive water in the blood be- comes more concentrated in active brain areas and is de- tectable by radioactivity detectors. The most widely used method at present is magnetic reso- nance imaging (MRI). This is based on the fact that changes in blood ﬂow cause changes in the blood’s magnetic proper- ties, which can be detected as changes in a strong imposed magnetic ﬁeld. This method was ﬁrst used in 1990 (Ogawa, Lee, Kay, & Tank). The current procedure is termed functional MRI (fMRI), involving very fast acquisition of images. A landmark publication in human brain imaging is the elegant book by two pioneers in the ﬁeld, Michael Posner and Marcus Raichle, Images of Mind (1994). The fMRI pro- cedures have several advantages, such as the fact that they are noninvasive—no radioactive substance is injected—and provide better spatial resolution than does PET imaging. Functional magnetic resonance imaging exploits variations in magnetic susceptibility that arise from molecular binding of oxygen to hemoglobin, which can be used to detect blood ﬂow changes associated with neuronal activity. At the present time, these neuronal activity-related signals can be derived from areas of the brain with a spatial resolution of 1 to 2 mm. Moreover, the temporal resolution of this functional imaging technique is compatible with the time course needed to carry out most perceptual and cognitive operations. An important and promising strategy for the use of fMRI is its use in con- junction with other kinds of neurobiological techniques, in- cluding neurophysiology and anatomical and behavioral analyses. Thus, fMRI provides an extraordinary new window through which one can probe the neural machinery of cogni- tion (Albright, 2000). CONCLUSION Physiological psychology, the ﬁeld concerned with biologi- cal substrates of behavior and experience (mind), has to be the most important discipline in psychology and the life sci- ences. The two great questions in science are the nature of the universe and the nature of the mind. Over the past century, the ﬁeld of physiological psychology has spun off a number of areas that are now separate ﬁelds in their own right: vision, audition, psychophysiology, behavioral genetics, behavioral neuroscience, and cognitive neuroscience. It seems that in this sense physiological psychology is destined to self- destruct. But to participate in the process is surely among the most exciting intellectual endeavors of our time. REFERENCES Ackernecht, E. H., & Vallois, H. V. (1956). Franz Joseph Gall, in- ventor of phrenology and his collection (C. St. Leon. Trans.). Madison, WI: Medical School, Department of History. Adrian, E. D. (1940). Double representation of the feet in the sen- sory cortex of the cat. Journal of Physiology, 98, 16. Albright, T. D. (2000). Functional magnetic resonance imaging of the brain in nonhuman primates: A prospectus for research on aging. In P. C. Stern & L. L. Cartensen (Eds.), The aging mind. Washington, DC: National Academy Press. 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DEWSBURY 67 EARLY HISTORY 68 FORERUNNERS OF COMPARATIVE PSYCHOLOGY 68 COMPARATIVE PSYCHOLOGY BEFORE WORLD WAR I 69 BETWEEN THE WORLD WARS 71 Leaders of the Reconstruction 71 New Blood for Comparative Psychology 71 The State of Comparative Psychology between the Wars 73 COMPARATIVE PSYCHOLOGY SINCE WORLD WAR II 74 Personnel 74 Funding 75 Research Centers 75 Journals 76 Academic Societies 76 Soul-Searching 76 THREE IMPORTANT POSTWAR INFLUENCES 76 European Ethology 77 Sociobiology, Behavioral Ecology, and Evolutionary Psychology 77 Comparative Cognition 78 CONCLUSION: PERSISTENT ISSUES 79 REFERENCES 81 Comparative psychology has been a part of American psy- chology since its emergence as a separate discipline. As early as 1875, William James wrote to Harvard University presi- dent Charles W. Eliot “that a real science of man is now being built up out of the theory of evolution and the facts of ar- chaeology, the nervous system and the senses” (James, 1875/1935, p. 11). G. Stanley Hall (1901), founder of the American Psychological Association (APA), regarded the study of the evolution of the human soul as “the newest and perhaps richest ﬁeld for psychology” (pp. 731–732). Future Yale University president James Rowland Angell (1905) wrote that “if the evolutionary doctrine is correct, there seems to be no reason why we should not discover the fore- runners of our human minds in a study of the consciousness of animals” (p. 458). Although the ﬁeld has changed greatly over more than a century, some of the problems addressed during this earlier era remain relevant today (Boakes, 1984; Dewsbury, 1984). There is no universally accepted deﬁnition of comparative psychology, although there is general agreement concerning which research is included, excluded, or falls near its bound- aries. Comparative psychology may be regarded as that part of the ﬁeld of animal psychology, the psychology of nonhu- man animals, not included within either physiological psy- chology or process-oriented learning studies. Such research generally is conducted on either species or behavioral pat- terns not generally utilized in those ﬁelds. Comparative psy- chology ﬁts within the broad ﬁeld of animal behavior studies, which includes research by scientists from many disciplines. Much research within comparative psychology includes no overt comparisons among species. The goals are to develop a complete understanding of general principles governing mind and behavior including its origins (evolutionary, ge- netic, and developmental), control (internal and external), and consequences (for the individual, the surrounding envi- ronment, and for subsequent evolution). Comparison is but one method of reaching such understanding. Comparative psychologists take seriously the effects of behavior on differ- ential reproduction and, ultimately, evolutionary change. In an article on the contributions of comparative psychology to child study, a favorite approach of Hall’s, Linus Kline (1904) used the term zoological psychology as a label for the ﬁeld; this may be a more accurate descriptive title than compara- tive psychology because it highlights the connection of comparative psychology with zoology—especially so-called whole-animal biology. In this chapter, I trace the history of comparative psychol- ogy from early cave paintings to the present. This entails ﬁrst a consideration of the British forerunners of comparative psy- chology and the emergence of the ﬁeld prior to World War I. 68 Comparative Psychology This was followed by a postwar period of decline, as younger comparative psychologists were unable to sustain careers, and then by a resurgence of activity between the world wars. The ﬁeld has remained active since World War II and has been strongly inﬂuenced by developments in European ethology, sociobiology, and cognitive science. EARLY HISTORY Humans have a long history of interest in animal behavior. Perhaps the ﬁrst evidence of this is from the cave paintings depicting animals in southern Europe dating from the Upper Paleolithic period, 35,000 to 10,000 years before the present. Domestication of animals began about 11,500 years ago in the Middle East and Asia (Singer, 1981). Among the ancient Greeks, Herodotus (c. 425 B . C .) described habits and behav- ior of animals and made observations on animal physiology. Interest in animals was brought to a new level by Aristotle (384–322 B . C .). He relied on observation and inductive rea- soning, not just speculation, to develop a natural history of many species. Aristotle believed in the continuity of species, though he believed species to be ﬁxed rather than evolving. He also proposed the notion of a Scala naturae, a single di- mension along which all species could be ordered. Although this idea, transformed from dealing with the characteristics of the animals’ souls to their level of intelligence, is still popu- lar today, it is widely regarded as fallacious. Evolution is branching, and species do not lie along a single continuum. During the long period from the ancient Greeks to the mid-nineteenth century, interest in animal behavior was strong in three areas. Such individuals as Frederick II of Hohenstaufen (1194–1250), John Ray (1627–1705), and Charles George Leroy (1723–1757), studied animal behavior in nature and developed the area of natural history. A second area was applied animal behavior, where domestication and selective breeding of livestock, dogs, and other species con- tinued and was perfected. Falconers developed remarkable skills in the control of behavior (Mountjoy, 1980). Finally, the relation between human and nonhuman animals became an area of interest to philosophers. The seventeenth-century French philosopher René Descartes is credited with popularizing the view that there is an absolute gulf between humans and all other species. According to Descartes, humans are the only ones to possess the immate- rial rational soul that enables abstract reasoning and self- awareness; animals are automata that can carry on simple mental functions but cannot think or have language. Darwin’s work would discredit this dichotomy. An interesting di- chotomy developed between the British and continental philosophers regarding the developmental origins of ideas. British philosophers such as John Locke and David Hume be- lieved that all knowledge originated in experience. For Locke, themind was a tabula rasa, orblankslate. Continental philoso- phers,such as ImmanuelKant, proposed theexistence of anac- tive mind with a priori properties, suchas categories, that acted on experience to produce knowledge. This geographic differ- ence can be seen in contrasting the British and continental ap- proaches to the ﬁeld of ethology in the twentieth century. FORERUNNERS OF COMPARATIVE PSYCHOLOGY The intellectual grounding for a comparative psychology was provided in the nineteenth century with the development of the theory of evolution. The notion that evolution had oc- curred did not originate with Charles Darwin but rather de- veloped with the work of such individuals as Erasmus Darwin (his grandfather), Jean-Baptiste Pierre Antoinne de Monet de Lamarck, and Robert Chambers. Darwin provided a viable mechanism, the theory of natural selection, and es- tablished that no mystical forces affected the direction of evolutionary change. Change is the result of differential re- production under prevailing circumstances. What was critical for comparative psychology was the solidiﬁcation of the idea that human and nonhuman animal behavior is continuous and thus both can be studied and compared with similar methods. This need not imply that there are no important differences between humans and nonhuman animals (henceforth called animals ), but only that there are similarities and that any dif- ferences will best be revealed through careful comparisons. Although his Origin of Species (1859) and Descent of Man (1871) are Darwin’s best-known works, The Expression of the Emotions in Man and Animals (1872) was especially im- portant for comparative psychology because it showed how a comparative study of behavior might be conducted. Among Darwin’s many contributions to comparative psychology, we should remember that in the 1871 work Darwin laid out im- portant principles of sexual selection, the manner in which individual males and females ﬁnd mates and achieve repro- ductive success. Sexual selection has been an important topic in the ﬁeld of comparative psychology in recent years. Darwin’s protégé was George John Romanes, an excellent scientist, who worked with jellyﬁsh, starﬁsh, and sea urchins (Romanes, 1885). He was also committed to demonstrating Darwin’s principle of continuity in instinct and mind in hu- mans and animals. In Animal Intelligence (1882), Romanes, like most of his contemporaries, relied heavily on anecdotes, reports of single instances of behavior provided by various Comparative Psychology before World War I 69 associates. Although he tried to be careful in selecting these, some of them are rather far-fetched and have led to a viliﬁca- tion of Romanes and his methods. His reputation was fur- ther tarnished because, in his efforts to establish continuity, he tended to anthropomorphize (i.e., attribute human proper- ties to animals). Romanes’s many contributions are often neglected. A more conservative approach to animal behavior was taken by another Englishman, C. Lloyd Morgan, in his book An Introduction to Comparative Psychology (1894). Al- though this was a multifaceted work, Morgan is best remem- bered for one sentence, which has come to be known as Lloyd Morgan’s Canon: In no case may we interpret an action as the outcome of the ex- ercise of a higher psychical faculty, if it can be interpreted as the outcome of the exercise of one which stands lower in the psy- chological scale. (p. 53) Morgan clearly believed in a hierarchy of psychological processes, with some processes being higher, or more com- plex, than others. He suggested that we can only invoke the higher processes when behavior cannot be explained in terms of lower, or simpler, psychological processes. This principle is often confused with a related dictum, the law of parsimony (Dewsbury, 1984; Newbury, 1954). The terms “law of parsi- mony” and “Occam’s razor” can be used interchangeably for most purposes. These terms refer to the assumptions made in providing an explanation rather than to the complexity of the psychological processes that are invoked. Thus, other things being equal, we should strive for explanations that do not multiply explanatory principles and that are simple explana- tions in that sense. Morgan (1894), by contrast, noted that “the simplicity of an explanation is no necessary criterion of its truth” (p. 54). It would be possible to construct an inter- pretation based on lower psychological processes but that introduces numerous additional assumptions and is thus con- sistent with Morgan’s Canon but inconsistent with the law of parsimony or one that is parsimonious but in violation of the canon. The canon implies, for example, that we should be very careful in attributing consciousness to animals. By no means did Morgan wish to suggest that animals lack con- sciousness; rather, he meant that we could invoke such a process only when necessary to explain observations that could not be explained with psychologically lower complex processes. Other investigations in the growing ﬁeld of animal behavior studies were conducted by such Britishers as Douglas A. Spalding, Sir John Lubbock, and L. T. Hobhouse and Americans such as Lewis Henry Morgan, T. Wesley Mills, George W. Peckham, and Elizabeth Peckham. Espe- cially notable was the work of Charles H. Turner on the com- parative psychology of crayﬁsh, ants, spiders, bees, and other invertebrates. Turner was an African American scientist of the time who published signiﬁcant research in major journals (see Cadwallader, 1984). COMPARATIVE PSYCHOLOGY BEFORE WORLD WAR I Building on these foundations, comparative psychology emerged as a signiﬁcant, visible discipline during the late nineteenth and early twentieth centuries in the universities of the United States (see Dewsbury, 1992). Hall had been called to the presidency of Clark University and brought with him Edmund C. Sanford, who ran the laboratory. They taught courses and attracted students to comparative psychology. The laboratory course included work on microscopic ani- mals, ants, ﬁsh, chicks, white rats, and kittens. Graduate stu- dent Linus Kline (1899), who did some of the teaching, suggested that “a careful study of the instincts, dominant traits and habits of an animal as expressed in its free life—in brief its natural history should precede as far as possible any experimental study” (p. 399). The best known of the early Clark studies were those on maze learning published by Willard S. Small (1901). Kline mentioned to Sanford that he had observed runways built by feral rats under the porch of his father’s cabin in Virginia, and Sanford suggested the use of a Hampton Court maze as an analog of the learning re- quired of rats in nature (Miles, 1930). Small and Kline con- structed the mazes and other devices in which to study the learning process in rats. Thus, the early studies were designed to mimic situations the subjects faced under natural condi- tions. The Clark program was not limited to such studies. Under the inﬂuence of Hall, there was a strong developmen- tal focus, as in Small’s (1899) study of the development of behavior in rats and in Conradi’s (1905) study of the devel- opment of song in English sparrows. James P. Porter (1906) analyzed the naturally occurring behavioral patterns of two genera of spiders. Robert M. Yerkes, under the inﬂuence of William James and Hugo Münsterberg, was a mainstay of comparative psychology during this period at Harvard. He studied the be- havior of a wide variety of invertebrates such as crayﬁsh (Yerkes & Huggins, 1903) and published one of the early classics of the ﬁeld, The Dancing Mouse (Yerkes, 1907), a comprehensive study of a mutant mouse strain. Yerkes and his students also studied a variety of behavioral patterns and species, including sensory function, such as Cole’s (1910) 70 Comparative Psychology study of the reactions of frogs to four chlorides; genetics and development, such as Yerkes and Bloomﬁeld’s (1910) study of the reactions of kittens to mice; and learning, such as Coburn and Yerkes’s (1915) study of crows. Edward Bradford Titchener dominated psychology at Cornell University. Although he is often portrayed as having opposed comparative psychology, he conducted a number of studies in the ﬁeld early in his career (Dewsbury, 1997). A prize student at Cornell was his ﬁrst PhD, Margaret Floy Washburn, who later became the second woman elected to the presidency of the APA. Her most notable contribution to comparative psychology was her book The Animal Mind (1908), that went through four editions and was the standard textbook in comparative psychology into the 1930s. Re- search at Cornell included a study of vision in ﬁsh (M. F. Washburn & Bentley, 1906) and one on learning in parame- cia (Day & Bentley, 1911). Even Edwin G. Boring (1912), future historian of psychology, published a study of phototro- pisms in ﬂatworms. The pride of the program at the University of Chicago, directed by Angell, was John Broadus Watson. Although Watson became famous later in his career for his writings on behaviorism, he did work in comparative psychology dur- ing his younger years. His dissertation, Animal Education (Watson, 1903), was an early study in developmental psy- chobiology, as Watson tried to correlate the development of learning in rats with the development of the nervous system. Watson also studied imitation in monkeys and spent several summers studying noddy and sooty terns on the Dry Tortugas Islands off Florida (e.g., Watson & Lashley, 1915). This study anticipated some later research in ethology. Many psycholo- gists who know only his writings on behaviorism are surprised by his earlier thinking on instinctive behavior (Watson, 1912). Most of the other students in animal psy- chology at Chicago worked on rats, although Clarence S. Yoakum (1909) studied learning in squirrels. Edward L. Thorndike had a brief, but extremely inﬂuen- tial, career in comparative psychology. After conducting some research with William James at Harvard, Thorndike moved to Columbia University, where he completed his PhD under James McKeen Cattell in 1898. After a year at Western Reserve University, he returned to Columbia, where he spent the remainder of his career, most of it as an educational psy- chologist. His dissertation, Animal Intelligence (Thorndike, 1898), was a classic study of cats learning to escape from puz- zle boxes; Thorndike (1911) later expanded this work with the addition of several previously published articles. He be- lieved that cats used simple trial and error to learn to operate manipulanda to escape from the compartments in which they had been enclosed; they kept emitting different behavioral patterns until one was successful. Further, he believed that virtually all learning in all species followed the same laws of trial-and-error and reward (the law of effect). This provided little impetus for comparative analysis. Thorndike’s major contribution was the development of precise methods for careful study of learning in the laboratory. In the tradition of C. L. Morgan, Thorndike generally sought to explain behav- ior in terms of relatively simple processes and eschewed no- tions of insight in creative problem solving. T. Wesley Mills took a very different approach, closer to that of Romanes than to that of Morgan. This led to a bitter exchange of mutually critical articles. Mills emphasized the importance of testing under natural conditions, writing of Thorndike’s puzzle box experiments that one might “as well enclose a living man in a cofﬁn, lower him, against his will, into the earth, and attempt to deduce normal psychology from his conduct” (Mills, 1899, p. 266). Thorndike (1899) defended his research as the only way “to give us an explanatory psychology and not fragments of natural history” (p. 415). Karl S. Lashley, best known as a physiological psycholo- gist, also had a lifelong interest in comparative psychology. He was inﬂuenced by Watson at Johns Hopkins and spent one summer working with him on the tern project. Lashley inﬂu- enced comparative psychology not only through his research and integrative writings but also through his students. Harry M. Johnson was another Hopkins-trained comparative psy- chologist, as exempliﬁed in his study of visual pattern dis- crimination in dogs, monkeys, and chicks (Johnson, 1914). Other comparative psychologists in graduate school dur- ing this period included John F. Shepard at the University of Michigan, who did many studies of learning in ants and rats (see Raphelson, 1980), and William T. Shepherd at George Washington University, who worked on a variety of species (e.g., Shepherd, 1915). Perhaps the most inﬂuential foreign-born comparative psychologist was Wolfgang Köhler, who completed a doctor- ate at the University of Berlin in 1909. Much of his career was devoted to the development and promotion of Gestalt psychology. His major work in comparative psychology was conducted on the island of Tenerife in the Canary Islands dur- ing World War I. Köhler’s best-known studies were of prob- lem solving with chimpanzees. These studies used such tasks as the stacking of boxes to reach a banana suspended above the animals’ enclosure and stick problems in which the chim- panzees had to manipulate sticks of one sort or another to reach a banana that was placed outside of the enclosure where it could be reached with a stick but not without it (Köhler, 1925). Little original theory was created during this period. The guiding theoretical framework came from the theory of Between the World Wars 71 evolution. Attention was devoted to building an empirical foundation for the ﬁeld. The range of species studied was ex- tensive. Although the study of learning came to be especially prominent, there was much research on sensory function, de- velopment, and social behavior as well. Although only a few comparative psychologists studied animal behavior in the ﬁeld, many were aware of the place of their study subjects in nature and used that awareness in understanding behavior. Although the foundations for a stable ﬁeld of comparative psychology appeared to have been laid, it was not to be—at least not yet. A number of problems arose. The major difﬁ- culty lay in the place of comparative psychology, as a study of behavior in animals, in psychology, a discipline most viewed as the study of mind and behavior in humans. Despite its intellectual and historical connections with the rest of psy- chology, comparative psychology was perceived as a periph- eral ﬁeld. Pressures were brought to bear on those trained in comparative psychology to switch and move to other re- search areas, especially applied ﬁelds. At Harvard, for exam- ple, Münsterberg (1911) wrote of Yerkes to President Abbott Lawrence Lowell that “anyone interested in those animal studies alone is in no way a real psychologist, and really no longer belongs in the philosophy department.” The situation was complicated because psychologists doing laboratory studies of animals required special facilities that were both expensive and viewed by some as undesirable because of odor and atmosphere. Some had philosophical objections to animal research. It became clear to many that the path to pro- motion was to leave comparative psychology for applied ﬁelds (Dewsbury, 1992). As a result, most comparative psy- chologists educated during these years followed such paths and left the ﬁeld. The American entry into World War I and the loss of personnel to military endeavors exacerbated the situation. BETWEEN THE WORLD WARS The period running from the late 1910s and through the 1920s was a nadir for the ﬁeld. With the old foundation for the ﬁeld gone, a new one had to be constructed. Leaders of the Reconstruction Few psychologists were in the universities to engage in re- construction. Harvey Carr and Walter Hunter, products of the Chicago program, remained active, as did Karl Lashley, who was inﬂuential in the careers of many aspiring compara- tive psychologists. After the war, Yerkes spent several years in Washington before Angell, then the president of Yale University, brought him to New Haven in 1924. Yerkes and Lashley would be pivotal in the redevelopment of compara- tive psychology that would help to establish it as a ﬁeld that has been strong ever since. Several other individuals who would lead the reformulation of comparative psychology were educated in other programs scattered about the country. Although Yerkes functioned as an administrator in Washington until 1924, he never lost sight of his plan for a re- search station where nonhuman primates might be studied (Yerkes, 1916). In 1915, he took a half-year sabbatical to conduct research on primates in California. In 1923 he pur- chased two animals, Chim and Panzee, for study, primarily at his summer home in New Hampshire. The following sum- mer, he studied primates in the colony of a private collector, Madame Rosalia Abreu in Havana, Cuba. All the while, he was publishing material on primate research (e.g., Yerkes, 1925) and lobbying various private foundations for funds for a primate facility. Finally, in 1925 the Rockefeller Founda- tion appropriated funds to support a primate facility in New Haven and, in 1929, for a feasibility study for a remote pri- mate station. Later that year, $500,000 was granted and Yerkes established the Anthropoid Station of Yale University (later renamed the Yale Laboratories of Primate Biology when it was incorporated in 1935 and the Yerkes Laborato- ries of Primate Biology upon its founder’s retirement). The facility would remain in Orange Park until 1965 and was a focal point of research on the great apes. Lashley moved to the University of Minnesota in 1917 and, with an interlude of work in Washington, D.C., remained there until he moved to Chicago, ﬁrst to the Behavior Research Fund of the Institute for Juvenile Research in 1926 and then to the University of Chicago in 1929. He moved to Harvard University in 1935, and in 1942, he became the sec- ond director of the Yerkes Laboratories of Primate Biology in Orange Park, Florida, from which he retired in 1955. New Blood for Comparative Psychology A cluster of comparative psychologists of lasting impact completed graduate training during the 1920s and 1930s. Per- haps the ﬁrst of the new generation of comparative psycholo- gists was Calvin P. Stone, who completed his PhD under Lashley at Minnesota in 1921. Stone went on to a long career at Stanford University, where he was noted for his studies of sexual behavior and the development of behavior, for his ed- itorial work, and for mentoring numerous students. Zing-Yang Kuo, a native of Swatow, Kwangton, China, completed a doctorate with Edward C. Tolman at the Univer- sity of California at Berkeley in 1923. The primary issue with which Kuo grappled during his career was the nature of 72 Comparative Psychology development and the relative roles of nature and nurture. At various stages of his career he concluded that there was either little evidence of genetic effects or that genetic and environ- mental inﬂuences were so intimately entwined that it was im- possible to separate them. Although he was able to publish some articles throughout his career, his difﬁculty in ﬁnding employment in the United States and his involvement in ad- ministrative and political turmoil in China greatly limited his inﬂuence. Carl J. Warden completed a PhD at the University of Chicago in 1922; he spent much of his career at Columbia University. Among his contributions to the ﬁeld were his writings on the history of comparative psychology, text- books, and research. The latter often entailed use of the Columbia Obstruction Box, in which a rat had to cross a shock grid in order to reach an incentive (e.g., Warden, 1931). The greater the intensity of the shock the animal was willing to endure, the greater was the animal’s drive believed to be. Henry W. Nissen completed a PhD with Warden at Columbia in 1929. He spent much of career working on pri- mate behavior under the inﬂuence of Yerkes, ﬁrst at Yale University and later at the Yerkes Laboratories of Primate Biology. He was the director of the latter facility from 1955 to 1958. Nissen is said to have known more about chim- panzees and their behavior than anyone else of his time but was a self-effacing psychologist whose inﬂuence was limited by his reticence. Nevertheless, his career was prominent enough to earn him election to the prestigious National Academy of Sciences of the United States. A remarkable cluster of students worked with Lashley at the University of Chicago during the early 1930s (Dewsbury, in press-a). Norman R. F. Maier completed a PhD with Shepard at Michigan in 1928 and, after a year on the faculty at Long Island University, went to work with Lashley during 1929–1931. He then spent most of his career back at Michigan. In comparative psychology, Maier is best known for his studies of problem solving in which he suggested that rats do not learn to solve complex processes via the simple associative processes suggested by Thorndike but rather use a process of reasoning (e.g., Maier, 1937). This was part of a fairly substantial interest in cognitive approaches to behavior during the 1930s (Dewsbury, 2000). He was also interested in the abnormal behavior, including ﬁxations and seizures, that sometimes occurred in his testing situations. Theodore C. Schneirla also completed his doctorate with Shepard at Michigan in 1928. Shepard interested Schneirla in studies of the behavior of ants, which became the focus of Schneirla’s career. In 1927, he moved to New York Uni- versity, combining his duties there with a position at the American Museum of Natural History during much of his career. He went to work with Lashley in Chicago during 1930–1931. Schneirla was a primary exemplar of the role of ﬁeld research in comparative psychology, as he made many trips to study the complex adaptive patterns of various species of ants at many sites. He also conducted notable lab- oratory research on learning in ants. Schneirla also engaged in theory construction. He advocated a concept of integrative levels, occupied by different species. With this concept, he called for caution in generalizing across widely diverse taxa. He also was a strong advocate of the epigenetic approach to development and opposed the notion that some behavioral patterns are innate. He believed that tendencies to approach toward and withdraw from stimuli of varying intensities played an important role in development (see Aronson, Tobach, Rosenblatt, & Lehrman, 1969). Frank A. Beach completed an MA degree at the Kansas State Teachers College in Emporia before going to Chicago to complete his doctorate. He worked with Lashley during 1933–1934, taught high school for a year, and then returned to Chicago for further graduate work. Lashley was gone by then, but Beach followed him to Harvard in 1936. He com- pleted the ﬁnal requirements for the Chicago PhD in 1940. Beach spent his career at the American Museum of Natural History, Yale University, and the University of California, Berkeley. He is probably best known today for a series of incisive articles he wrote about the state of comparative psy- chology and the conceptual foundations thereof. The best- known example is his “The Snark Was a Boojum” (Beach, 1950). Beach argued that throughout the ﬁrst half of the twentieth century, comparative psychologists had become interested in a more narrow range of behavioral patterns and progressively fewer species, primarily white rats. He suggested that this was not a healthy development. In “The Descent of Instinct” (1955), he criticized simplistic concep- tions of the concept of instinct. His research program was a broadly based attack directed primarily at the determinants of reproductive behavior. He was interested in the neural bases, endocrine correlates, evolution, development, and situational determinants of reproductive and social behavioral patterns. Isadore Krechevsky, later David Krech, studied ﬁrst at New York University but completed his doctorate with Edward C. Tolman at the University of California at Berke- ley. He then moved to the University of Chicago, initially with Lashley, where he remained from 1933 to 1937. A polit- ical activist, he had to change afﬁliations with some fre- quency because of difﬁculties with administrators, but he spent the last part of his career, beginning in 1947, at Berke- ley. Krechevsky (1932) showed that as rats learn mazes, they appear to form “hypotheses,” systematic runs of choices gov- erned by different rules, each of which is tried as a solution Between the World Wars 73 is sought. As with Maier’s work, this was part of the 1930s effort in comparative cognition. The ﬁfth important comparatively oriented student to work with Lashley at Chicago was Donald Olding Hebb. Hebb moved to Chicago in 1934 and accompanied his men- tor to Harvard after one year. He received a Harvard PhD in 1936. He then ﬁlled positions in Canada and, in 1942, re- joined Lashley in Orange Park. In 1947, Hebb joined the fac- ulty at McGill, from which he retired in 1974. Like Lashley, Hebb is best remembered for his contributions to physiologi- cal psychology. His The Organization of Behavior (1949) was important in the reinvigoration of physiological psychology after World War II and introduced the so-called Hebb synapse to psychology. Like Beach and Schneirla, Hebb worked to- ward reinterpretation of behavioral patterns that appeared to be innate (Hebb, 1953). His comparative interests are also ap- parent in his efforts to get studies of animal social behavior more recognition in the ﬁeld of social psychology (Hebb & Thompson, 1954). Two important comparative psychologists completed PhDs under Stone at Stanford. Harry F. Harlow completed the PhD in 1930 and spent the rest of his career at the University of Wisconsin. He spent much of his career study- ing learning in rhesus monkeys, where he developed an error factor theory, according to which the primary process during learning often involved the manner in which errors were eliminated. Harlow is best known, however, for his work on behavioral development. He found the social and reproduc- tive behavior of rhesus monkeys reared in the absence of their parents and siblings to be greatly distorted. Deﬁcits in learning were found to be much less severe. With many im- portant students educated in his program and with his editor- ial and administrative work, Harlow was a very inﬂuential comparative psychologist. The other Stanford-Stone graduate was C. Ray Carpenter, who completed his studies in 1932 with work on endocrine inﬂuences on pigeons. He is best known, however, as the “father” of primate ﬁeld research. With the help of Yerkes, Carpenter began a series of ﬁeld studies in locations such as Panama, Southeast Asia, and India (e.g., Carpenter, 1934). He established a colony of rhesus monkeys on the island of Cayo Santiago, off Puerto Rico. This was the ﬁrst sophisti- cated work on primates in their native habitats. This ﬁeld has exploded in recent years with the work of such scientists as Jane Goodall and George Schaller. Many people are sur- prised to learn of the role of a psychologist in establishing the subdiscipline of primate ﬁeld research. Carpenter spent much of his career at the Pennsylvania State University, where he also devoted much effort to documenting studies of primates and other species on ﬁlm. Other comparative psychologists completing graduate work during this era included Curt P. Richter (Johns Hopkins, 1921), Carl Murchison (Johns Hopkins, 1923), Leonard Carmichael (Harvard, 1924), Lucien H. Warner (Columbia, 1926), Otto L. Tinklepaugh (Berkeley, 1927), Winthrop N. Kellogg (Columbia, 1929), and Meredith P. Crawford (Columbia, 1935). The State of Comparative Psychology between the Wars The comparative psychologists educated during the 1920s and 1930s placed comparative psychology on a ﬁrm footing. Unlike the pre–World War I cadre, this group was successful in securing research support and in educating a next genera- tion of comparative psychologists who would carry on the tradition. Nevertheless, much was not well. This group of comparative psychologists, which appears to coalesce as a coherent unit when viewed in retrospect, did not appear so when viewed in its time. There were a number of reasons for this. Disciplines and subdisciplines become recognizable and inﬂuential with the development of a set of institutional land- marks including departments, textbooks, courses, research facilities, organizations, meetings, and journals. During this period, comparative psychology was well established in many departments, and courses were a staple in many places. In other respects, however, it lacked elements that foster cohesion. Textbooks The 1930s saw the greatest burst of publication for textbooks in the history of the ﬁeld. Margaret Floy Washburn’s The An- imal Mind had been dominant since 1908. Her fourth edition appeared in 1936. The most inﬂuential book of the era was Maier and Schneirla’s Principles of Animal Psychology (1935). The textbook provided a comprehensive overview of the ﬁeld, beginning with 11 chapters organized according to animal taxa. Material concerning receptor equipment, sensi- tivity, conduction, and the action system is provided for each group. The second part of the book is concerned with natively determined behavior, sensory function, and neural mecha- nisms in mammals. Part III addresses learning and mental processes. The most comprehensive of the works was the three- volume Comparative Psychology: A Comprehensive Treatise (1935, 1936, 1940) by Warden, T. N. Jenkins, and Warner. The ﬁrst volume deals with principles and methods; the second volume with plants and invertebrates; and the third volume with vertebrates. 74 Comparative Psychology A tradition of edited textbooks in the ﬁeld began with F. A. Moss’s (1934) Comparative Psychology . The 15 chapters included information on maturation, motivation, sensory function, learning, individual differences, animal social psy- chology, and a set of related topics. This work was followed by similar volumes from various editors at regular intervals in 1942, 1951, 1960, and 1973. An interesting approach was taken by Normal L. Munn (1933) with his An Introduction to Animal Psychology . The book is concerned solely with the behavior of laboratory rats and provides a comprehensive review of many characteristics of rats. Other Characteristics In some respects, the textbooks were the only bright spot in the institutionalization of comparative psychology during the period between the wars. One problem was that of deﬁnition, a difﬁculty that still affects the ﬁeld today. There was clearly a cadre of comparative psychologists of the sort included by my deﬁnition. The term comparative psychology, however, was used in a variety of ways. Often, it referred to all animal psychology. Important work was being done in the ﬁelds of animal learning and cognition during this period. Much of the work, however, was done within a more process-oriented framework than most work in comparative psychology as de- ﬁned here. Similarly, numerous physiological studies were conducted. The true comparative tradition was obscured, in part, because the ﬁeld lacked a clearly differentiating name, clear deﬁnition, and less permeable boundaries. A landmark was the beginning of the publication of the Journal of Comparative Psychology in 1922. The ﬁeld had had other journals, including the Journal of Comparative Neurology and Psychology (1904–1910), the Journal of Ani- mal Behavior (1911–1917), and Psychobiology (1917–1920). With the Journal of Comparative Psychology, however, the ﬁeld ﬁnally appeared to have a named journal to provide unity for the discipline. However, it was not to be. The journal be- came one of animal psychology and the primary vehicle for the publication of research in all ﬁelds of animal psychology, thus muddying the deﬁnitional problem even further. Indeed, during the 1920s, the Journal of Comparative Psychology in- cluded a signiﬁcant number of studies of human behavior (Dewsbury, 1998). A complete perspective on comparative psychology re- quires consideration of its ﬂaws as well as its accomplish- ments. Some of the writings of the time appear to be racist, at least by contemporary standards. The early volumes of the Journal of Comparative Psychology included numerous arti- cles on race differences regarding performance on intelligence tests, emotional traits, and physical development. In addition, Watson (1919) wrote that “psychologists persistently main- tain that cleanliness is instinctive, in spite of the ﬁlth of the negro, of the savage, and of the child” (p. 260), and Yerkes (1925) wrote that “certainly these three types of ape [chim- panzees, orangutans, and gorillas] do not differ more obvi- ously than do such subdivisions of mankind as the American Indian, the Caucasian, and the Negro” (p. 56). There are many aspects of the history of comparative psychology that are wor- thy of pride; a balanced view must include aspects lacking in such worth. Perhaps underlying the looseness of organization of com- parative psychology was a lack of identity among the leaders in the ﬁeld. Although all would probably have accepted the title of comparative psychologist, there was no sense of unity or effort to differentiate their work from that of other animal psychologists who often were included as “comparative psy- chologists.” There was no unifying theory of the sort devel- oped by the followers of B. F. Skinner. There was no agenda of the sort later promulgated by the European ethologists. Most comparative psychologists of the era were independent- minded individuals concerned with doing their research, re- porting it at existing meetings, and publishing it in mainline journals. There were no efforts to form new organizations or otherwise band together to deﬁne the developing tradition with any precision. As a result, the individual researchers gained respect and prestige for their efforts but they lacked real inﬂuence as a group. The subdiscipline that seems so clear in retrospect was not developed as an entity. COMPARATIVE PSYCHOLOGY SINCE WORLD WAR II The story of comparative psychology since World War II is one of developments within the ﬁeld and response to inﬂu- ences from outside. The war caused some interruption in the efforts that could be devoted to comparative psychology. The improved funding environment and the growth of universi- ties after the war, however, fueled rapid growth. Personnel Most critical was the availability of personnel. With stable positions, most of the prewar generation of comparative psy- chologists were able to develop active laboratories and pro- duce a continued output of research. As universities grew and fellowships became available, this generation, in turn, produced a new generation of comparative psychologists. In 40 years at the University of Wisconsin, Harlow alone Comparative Psychology since World War II 75 supervised 35 PhDs, including such names as Abraham Maslow, Donald R. Meyer, John M. Warren, Gerald E. McClearn, Allen M. Schrier, Leonard A. Rosenblum, and Stephen J. Suomi (Suomi & Leroy, 1982). During his career, Beach supervised 41 predoctoral and postdoctoral students (McGill, Dewsbury, & Sachs, 1978). Schneirla left a legacy of inﬂuential students including Daniel S. Lehrman, Jay S. Rosenblatt, and Ethel Tobach. Similar programs were devel- oped elsewhere. Then, of course, these students found jobs, built laboratories, and began educating yet another genera- tion. Comparative psychology still had a problem in that many who published animal research early in their careers left to become prominent in other ﬁelds of psychology. Ex- amples include Maslow, William Bevan, Jerome S. Bruner, William K. Estes, Eugene Galanter, Eleanor J. Gibson, Jerome Kagan, Quinn McNemar, M. Brewster Smith, and Dael L. Wolﬂe. Comparative psychology was always a small part of the big picture of American psychology. Nevertheless, there was a solid cadre of comparative psychologists carrying on the tradition. Funding Critical to the growth of comparative psychology was the availability of funding. Prior to World War II, most fund- ing for research came either from local sources or from private foundations, with prospective recipients making the rounds seeking research support. An exception was the Rockefeller Foundation–funded Committee for Research in Problems of Sex (Aberle & Corner, 1953). The explosive growth of support for scientiﬁc research not only increased the funding available but changed the pattern to one that in- volved the submission of research proposals that were subse- quently subject to peer review. I have analyzed funding patterns for comparative psychol- ogy for 1948–1963 at both the National Institute of Mental Health (NIMH) and the National Science Foundation (NSF) (Dewsbury, in press-b). According to my analysis, the NIMH awarded a total of 117 grants in comparative psychology for approximately $5.6 million during this period. The mean grant was for 2.5 years with an annual budget of under $20,000. The NSF program in psychobiology, not begun until 1952, awarded 72 grants in comparative psychology for a total of over $1.4 million with a mean size much smaller than those from the NIMH. The top-10 grant getters at the NIMH were Harlow, Lehrman, John Paul Scott, Richter, Eckhard Hess, Nissen, Beach, William Mason, M. E. Bitterman, and Schneirla. Half of those—Beach, Harlow, Lehrman, Nissen, and Richter—were elected to the National Academy of Sciences of the United States. Nissen, Schneirla, and Richter also were among the top-10 grant getters in comparative psy- chology at the NSF. The leading research topic in the NIMH grants was behavioral development. The NSF grants were less concentrated, with a greater emphasis on sensation and perception and general studies of behavior. This input of funding helped to create a great surge of research in compar- ative psychology, still small relative to the rest of psychology but substantial relative to that which had come before. Research Centers Although most comparative psychologists were scattered about the country in various universities, this funding en- abled the development of several centers for research. The Yerkes Laboratories of Primate Biology in Orange Park, Florida, were pivotal. Yerkes remained as director from its founding in 1930 until 1941 (Yerkes, 1943). He was suc- ceeded in turn by Lashley and Nissen. Arthur J. Riopelle and Geoffroy Bourne were the ﬁnal two directors in Orange Park. When the federal government established a program of Regional Primate Centers, Emory University, which then owned the Laboratories, moved them to their home campus in Atlanta. In addition to its directors, many other scientists such as Roger Sperry, Kenneth Spence, Austin Riesen, Paul Schiller, Hebb, Mason, and many others worked in Orange Park. From 1930 to 1965, the total budget was over $2.5 mil- lion. During the early years, the funding came almost exclu- sively from university and private foundation sources. This was reversed, and during the last ﬁve years for which data are available, over two-thirds of the funds came from the federal government. With greatly increased funding, the facility has thrived in Atlanta, albeit with a more biomedical emphasis. Harlow established and directed a primate laboratory at the University of Wisconsin. With the founding of a Regional Primate Research Center in Madison in 1964, Harlow as- sumed its directorship as well. Behavior programs also thrived in regional primate research centers in New England, Louisiana, Oregon, Washington state, and Davis, California. The behavior program at the Jackson Laboratory in Bar Harbor, Maine, was founded by John Paul Scott, who was ed- ucated as a geneticist but functioned in departments of psy- chology during much of his career. Joined by John L. Fuller, Walter C. Stanley, John A. King, and others, the program re- ceived substantial grant support and became a center for re- search on inbred strains of house mice and ﬁve breeds of dogs. It was also the site of two important conferences that helped to coalesce the ﬁeld of animal behavior studies. Another focal point developed in the New York City area. In 1937, Beach moved to the American Museum of Natural History, where he founded the Department of Animal 76 Comparative Psychology Behavior in 1942. Schneirla and Lester R. Aronson joined him in the department; Schneirla succeeded Beach as curator when Beach left for Yale University in 1946. Students such as Tobach, Rosenblatt, and Howard Topoff graduated and re- mained in the New York area. After graduating, one promi- nent student in the program, Daniel S. Lehrman, moved to the Newark, New Jersey, campus of Rutgers University, where he founded the Institute of Animal Behavior in 1959. This, too, became an important center for education and research. The focus of this whole group was on an epigenetic approach to development, and this New York epigeneticist group pro- duced numerous students and programs in the ﬁeld. Journals Beginning in 1947, the Journal of Comparative Psychology adopted a name more descriptive of its coverage: the Journal of Comparative and Physiological Psychology . That title re- mained until 1983, when it was split into three journals even more descriptive of the three prominent parts of animal psy- chology at the time: Behavioral Neuroscience, the Journal of Experimental Psychology: Animal Behavior Processes, and the reformulated Journal of Comparative Psychology . The scope of the growth of animal psychology can be seen in an analysis of the articles appearing in the Journal of Com- parative and Physiological Psychology in 1963 as compared with 1949 (Dewsbury, in press-b). The number of articles published increased by a factor of nearly 3.5 from 60 to 208. There were few footnote credits to federal funding sources in 1949; by 1963, just 14 years later, it had risen eightfold. Comparative psychologists published in other American journals as well. Some comparative psychologists found American journals uncongenial and published in the grow- ing stable of European journals, including Animal Behav- iour, Behaviour, and the Zeitschrift für Tierpsychologie (now Ethology ). Academic Societies No one academic society has emerged as the primary home for comparative psychologists. The APA remains the leading organization of psychologists, but its Division of Behavioral Neuroscience and Comparative Psychology (Division 6) is but a small part of the APA. Some psychologists have given allegiance to other psychological organizations such as the Psychonomic Society or the American Psychological Society. The Animal Behavior Society in NorthAmerican, the Associ- ation for the Study of Animal Behavior in Great Britain, and the International Ethological Congress have become the lead- ing organizations in the ﬁeld of animal behavior studies. Many comparative psychologists participate in these. The International Society for Comparative psychology, founded in 1983, has great possibilities that have not yet been realized. Soul-Searching Comparative psychologists have often assessed the state of their discipline and often criticized the directions taken. Three major articles stand out. In his famous “The Snark Was a Boojum,” mentioned earlier, Beach (1950) argued that comparative psychology had begun as the study of a wide range of topics in a wide range of species but had degenerated into the study of learning in rats. He stressed the need for a resurrection of the breadth that had earlier characterized the ﬁeld. Although Beach’s analysis was ﬂawed (Dewsbury, 1998), it was quite inﬂuential. In the second major critique, Hodos and Campbell (1969) criticized comparative psychologists’ perspective on evolu- tionary history. They argued that comparative psychologists still utilized the concept of a Scala naturae, derived from Aristotle, that implies that all species can be placed along a single great chain of being. They pointed to the branching na- ture of evolutionary history and to the need for a more realis- tic selection of species in comparative analyses. In “Reﬂections on the Fall of Comparative Psychology: Is There a Lesson for Us All?” Lockard (1971) detailed 10 myths that he thought plagued the ﬁeld. He incorporated the problems discussed by both Beach (1950) and Hodos and Campbell (1969) and added that comparative psychologists had devoted too little effort to the study of individual differ- ences, species differences, genetics and evolution, and ﬁeld research. Lockard advocated a more realistic biological ap- proach for comparative psychology. All three were effective critiques that provoked much dis- cussion and appear to have helped to stimulate change. How- ever, all three appear to have been overstatements of the problems. This may have been caused, in part, by the lack of a clear differentiation of true comparative psychology from other important, but different, parts of animal psychology. A survey published in 1980 revealed that comparative psychol- ogists remained divided with regard to both the deﬁnition and status of the ﬁeld (Demarest, 1980).

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THREE IMPORTANT POSTWAR INFLUENCES There were three major inﬂuences on comparative psychol- ogy after World War II. The ﬁrst was from the important ﬁeld developed by European zoologists that became know as ethology. The second stemmed from the elaboration of Three Important Postwar Inﬂuences 77 Darwinian principles that led to sociobiology, behavioral ecology, and evolutionary psychology. Finally, the so-called cognitive revolution had important effects on the ﬁeld. European Ethology Three major inﬂuences have affected and reshaped compara- tive psychology since World War II. The ﬁrst was the full- blown arrival of European ethology. Ethology is as difﬁcult to deﬁne as comparative psychology. It is a school of animal behavior studies that was developed in Europe by Konrad Lorenz, Niko Tinbergen, and their associates (Thorpe, 1979). According to a common, though greatly exaggerated, way of differentiating ethology from comparative psychology, the former developed in European zoology with birds, ﬁsh, and insects as subjects in observational studies and ﬁeld experi- ments designed to understand instinct and evolution. Com- parative psychology is said to have developed within North American comparative psychology with mammals, espe- cially laboratory rats, as subjects in laboratory research emphasizing experimental control and statistical analysis de- signed to understand learning and development. Although these differences in emphasis are instructive, the extent to which they are exaggerations of comparative psychology should be apparent to the reader. On the other side, Lorenz himself conducted little ﬁeld research. Although European ethologists and American compara- tive psychologists had numerous interactions prior to World War II, it was only after the war that contact became exten- sive. The two disciplines that had each developed as “the ob- jective study of behavior” along different lines came into direct conﬂict. Ethologists criticized psychologists for em- phasizing laboratory research that may produce results irrele- vant to the natural habitat; psychologists bemoaned the lack of experimental control in ﬁeldwork of the ethologists. The most critical issues, however, centered about behav- ioral development and the nature of instinctive behavior. The battle over instinct had gone on in the United States for most of the century, and during the 1950s, few psychologists ac- cepted the concept; most emphasized the complex interaction of genes and environment in epigenesis. Based on their ﬁeld observations, ethologists, by contrast, were quite comfortable with discussions of instinctive behavior (e.g., Lorenz, 1950). Lehrman (1953) wrote a scathing rebuttal to Lorenz’s ap- proach, with particular criticism directed at the instinct con- cept. On the one hand, the critique shocked ethologists and polarized the ﬁelds; on the other hand, however, it placed the issues out in the open where they could be debated and re- solved. At a series of conferences, many of the differences were softened and mutual understanding increased. One anecdote is revealing. Lehrman was discussing the issues in a rather stiff and formal disagreement with two ethologists, Gerard Baerends and Jan van Iersel, in a hotel room in Montreal. At one point, van Iersel’s attention was distracted by a birdsong coming from the garden. Lehrman, a lifelong bird-watcher, replied that it was a hermit thrush. Van Iersel wanted to observe it, so the three went off on a birding expe- dition. When they returned to the debate, their shared feeling for animals lessened the hostility and fostered give-and-take discussion (Beer, 1975). Although some psychologists, such as Schneirla, saw little value in ethology, others, like Lehrman and Beach, saw much. Beach and Carpenter facilitated interaction by agree- ing to serve on the founding editorial board of the ethological journal Behaviour . Many students were attracted to the natu- ralism of the ethological analyses and traveled to Europe to work in ethological laboratories (Dewsbury, 1995). The two ﬁelds changed in each other’s directions. Tinbergen moved from the Netherlands to Oxford University and developed a program with English-speaking ethologists. In general, they moved more toward the American position than did the German-speaking ethologists. The result of this activity was a more uniﬁed approach to animal behavior studies, with more psychologists even more concerned with naturally oc- curring behavior and principles that would be valid in nature and more ethologists concerned with control and experimen- tal procedures. More and more psychologists participated in the biannual International Ethological Conferences, and more Europeans came to the United States for either short or ex- tended periods of time. In the process, both ﬁelds beneﬁted. Sociobiology, Behavioral Ecology, and Evolutionary Psychology The second major postwar inﬂuence came from the ﬁeld that has become known as sociobiology. Many scientists had been studying behavior from an evolutionary perspective for many years. However, some of the implications of Darwinian theory had not been fully thought through; this led to some question- able interpretations. During the 1960s and early 1970s, theo- rists such as William D. Hamilton, John Maynard Smith, and Robert Trivers reinterpreted Darwinian principles.Although a number of principles were reevaluated, two were critical. One principle concerns the level at which natural selection acts. Much had been written about how behavior evolves for the good of the species or group. Thus, for example, reproduction had been thought to have evolved for the perpetuation of the species. On reinterpretation, it was concluded that natural selection works primarily at the level of the individual or gene, not the group or species. This had many important 78 Comparative Psychology implications and permitted reinterpretation of many phenom- ena. If a nest of eggs is left unguarded in a colony of gulls, for example, other members of the colony will eat the eggs. This makes little sense if all were there to perpetuate the species but is easy to understand if competition was at the level of the in- dividual, with each selected to get its genes into the next gen- eration. It had been believed that, at high densities, some species temporarily curtail reproduction in order not to over- exploit the available environmental resources. This became reinterpreted as a temporary “strategy” that could beneﬁt the lifetime reproductive success of an individual by preserving energy during hard times. The extreme form of this approach was Richard Dawkins’s (1976) selﬁsh gene theory. The second new principle concerned inclusive ﬁtness. The term ﬁtness refers to the relative contribution of different in- dividuals to future gene pools. It was noted that an individual shares more genes with close relatives than with those to which one is unrelated. There are thus at least two ways in which one’s genes can be transmitted to the next generation. One is through reproduction (direct ﬁtness); the other is through facilitating the reproduction of close relatives (indi- rect ﬁtness or kin selection). Inclusive ﬁtness is, essentially, the sum of the two. Thus, what might appear to be altruistic behavior, which might lower one’s ﬁtness, might be adaptive in the long run. One might lower one’s direct ﬁtness but gain even more indirect ﬁtness in the process. The bottom line became lifetime inclusive ﬁtness. These two principles, and some others associated with them, led many students of animal behavior, both inside and outside of psychology, to reorient their research programs. Many students became interested in the study of the evolu- tionary causes of behavior (ultimate causation) as opposed to the immediate and developmental causes (proximate causa- tion). There were many studies of the role of kinship in be- havior. A large number of species were found to modulate their behavior depending on the degree of kinship shared with others with which they interacted (e.g., Holmes & Sherman, 1982; R. H. Porter, 1988). Sexual selection, a topic emphasized by Darwin in 1871, became a major focus of research, with psychologists emphasizing studies of mate choice (e.g., Beauchamp, Yamazaki, & Boyse, 1985) and male–male competition for mates (e.g., LeBoeuf, 1974). Oth- ers sharpened evolutionary interpretations of phenomena of research on learning (e.g., Hollis, 1990; Kamil & Clements, 1990; Timberlake, 1990). Evolutionary perspectives had been a part of comparative psychology since its founding; now studies became more reﬁned and were addressed more speciﬁcally to evolutionary principles. This orientation had been somewhat diffuse until E. O. Wilson (1975) organized and named the ﬁeld in his Sociobi- ology: The New Synthesis . Although Wilson predicted the demise of comparative psychology, the ﬁeld was, in fact, strengthened by the new perspective. The ﬁrst 26 chapters of Wilson’s book provided a synthesis of much work related to the ultimate causation of animal behavior. In the ﬁnal chap- ter, however, Wilson applied these principles to human be- havior. This approach proved highly controversial. Although some careful research on human behavior stemmed from this approach (e.g., Daly & Wilson, 1978), other writings rested on less solid ground. A split developed between the contro- versial studies of human behavior and those less controver- sial studies of animals. The term behavioral ecology was coined, in part, so that students of animal behavior could dis- tance themselves from some of the more speculative studies of humans. More recently, sociobiology has been reborn as evolu- tionary psychology . An emphasis in the older sociobiology had been upon the ways in which existing patterns of human behavior enhanced ﬁtness. In the newer perspective, the focus shifted to adaptiveness at the time that mecha- nisms of behavior evolved in the ancestors of humans, and it was recognized that many behavior patterns and tenden- cies, such as our attraction to sweet foods, might not be as adaptive under present conditions (e.g., Cosmides & Tooby, 1987). Comparative Cognition The third major inﬂuence on comparative psychology since World War II was from the “cognitive revolution.” According to the received view, the hegemony of behaviorism precluded cognitive approaches to behavior prior to the 1960s and a major revolution occurred thereafter. In fact, however, re- search with a cognitive orientation has long been a part of comparative psychology (e.g., Dewsbury, 2000; Wasserman, 1993). Although a case can be made that there was no true revolution (Leahey, 1992), it is clear that in recent years cog- nitive perspectives have achieved a prominence in compara- tive psychology that had not been apparent previously. This increased emphasis on cognitive perspectives was a part of the broad upsurge of interest in cognitive processes throughout psychology and related disciplines. Develop- ments in information processing, computers, and mathemati- cal logic fostered a reconstruction of psychology as a science of information as well as behavior. Some psychologists moved to overcome what they perceived as the counterpro- ductive constraints imposed by more behavioristic theories. Interest was directed in the manner in which animals represent their worlds and the consequences of such repre- sentation for behavior. Among the topics receiving renewed interest were atten- tion, memory, timing, concept formation, counting, social Conclusion: Persistent Issues 79 cognition, and language (Roitblat, 1987; Shettleworth, 1998; Vauclair, 1996). The most publicized research in this ﬁeld has been that of language learning. Although there had been earlier attempts to teach language to apes, such as that of Cathy and Keith Hayes at the Yerkes Laboratories (Hayes, 1951), three major approaches developed during the 1960s and 1970s. Gardner and Gardner (1969) taught American sign language to chimpanzee Washoe; Rumbaugh (e.g., Rumbaugh & Gill, 1976) studied acquisition of a computer- based language in chimpanzee Lana; and Premack (1971) used a system of sentence formation using pieces of plastic with chimpanzee Sarah. This groundbreaking research was both heavily criticized and staunchly defended. This pro- duced a fallow period, due largely to an absence of funding, during which little language research was conducted. This period was followed by a reformulation and rebirth of studies of animal language, the most remarkable of which were of bonobo Kanzi, who learned a symbol-based language with- out overt training and became efﬁcient in interpreting human speech (e.g., Savage-Rumbaugh et al., 1993). Other studies of language acquisition were conducted with an African grey parrot (Pepperberg, 1999), dolphins (Herman, 1987), and sea lions (Schusterman & Krieger, 1984). Many of the studies in animal cognition were derived from, and closely related to, traditional research in animal learning. Other scientists made an effort to more completely revolutionize the ﬁeld of animal cognition using language suggestive of conscious processes in animals. Leading the latter effort was American ethologist Donald R. Grifﬁn (e.g., Grifﬁn, 1976b); the ﬁeld became known as cognitive ethol- ogy. Advocates of this approach contended that, with ad- vances in methodology, there are now available methods that can provide windows to the minds of animals. According to Grifﬁn (1976a), “the hypothesis that some animals are indeed aware of what they do, and of internal images that affect their behavior, simpliﬁes our view of the universe by removing the need to maintain an unparsimonious assumption that our species is qualitatively unique in this important attribute” (p. 534). Critics disagreed, contending that no methods were yet available that enable scientists to observe the internal processes of animal minds. Among the focal areas of research in cognitive ethology have been studies of self-recognition in mirrors (e.g., Gallup, 1985). According to Gallup, humans, chimpanzees, orang- utans, and some gorillas are the only species to show evi- dence of self-recognition when presented with mirrors. The key evidence comes from “dot tests,” in which a dot is painted onto the forehead of an anesthetized animal to see if the ani- mal selectively touches the dot when awakened and presented with a mirror. Gallup believes that such behavior suggests awareness, self-awareness, and mind in chimpanzees. Critics disagree (Heyes, 1994). In some ways, the ﬁeld had returned to questions addressed a century ago. Psychologists disagree as to whether the results achieved during that century now permit a return to these questions in a more sophisticated manner or whether it is regressing to an earlier state. CONCLUSION: PERSISTENT ISSUES Looking back over a little more than a century of compara- tive psychology, a number of characteristics are apparent. The conceptual foundations of comparative psychology have changed greatly. Some of these changes have been generated from within the ﬁeld; many more have been stimulated from related ﬁelds. Comparative psychologists have excelled in doing research and greatly expanded the body of data avail- able. It is this mass of information, sometimes well organized and sometimes rather scattered, that has enabled a century of change. Comparative psychologists have been empiricists working to expand the observational and experimental foun- dation of the ﬁeld; there are few postmodern comparative psychologists. In spite of the changes that have occurred, however, throughout the century there have been some per- sistent issues that have characterized the ﬁeld. Surely the ﬁrst issue lies in the very deﬁnition of compar- ative psychology. Although comparative psychologists have written much about this problem, few have been truly both- ered by it. Even lacking a clear deﬁnition, most comparative psychologists have ignored the fuzzy boundaries of the ﬁeld and concentrated on the business of studying animals and building general principles of behavior. Underlying the research effort have been issues of method- ology. The ﬁeld has been characterized by eclecticism, as most researchers have used whatever methods have appeared appropriate for the problem under study. Although some com- parative psychologists, such as Carpenter, Schneirla, Yerkes, Mason, and others, have conducted ﬁeld research, most have preferred the controlled conditions of the laboratory. Not all comparative psychologists make overt comparisons among species; the goal is not one of comparison for the sake of com- parison but rather the development of principles of generality. Although some, such as Kline, Schneirla, and James V. McConnell, have studied invertebrates, most have concen- trated on vertebrates, especially mammals. Some, such as Beach, Lehrman, and Carpenter, have concentrated on natu- rally occurring behavioral patterns; others have turned their attention to the study of learning and motivation in a compar- ative context. Although experimentation is the preferred method, many observational and correlational studies have been important. The anecdotes that characterized early re- search disappeared as laboratory methodology became more