Biographical Memoirs: Volume 89 (2007)

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WILLIAM DUWAYNE NEFF October 27, 1912–May 10, 2002 BY JAY M. GOLDBERG AND NELSON Y.-S. KIANG W ILLIAM DUWAYNE (“DEWEY”) NEFF WAS a plainspoken Mid- westerner, who became committed to the study of hear- ing as an undergraduate and maintained that commitment throughout his career. Best known for his use of the abla- tion method to study the functions of the various levels of the auditory pathways, he combined behavioral, electrophysi- ological, and neuroanatomical techniques to define deficits in auditory function more elaborate than simple audiom- etry. Rather than concocting intricate theories, he was con- tent to follow experimental facts wherever they led. Because of his sound judgment and no-nonsense style, he was asked to provide advice to numerous professional and govern- mental agencies. His straightforward, nonpedantic approach to science appealed to graduate students. It may very well be that the large number of graduate students that he trained is one of his most enduring legacies. Many of these gradu- ate students went on to distinguished careers and, follow- ing in his footsteps, trained many successful scientists. Dewey Neff was born on October 27, 1912, in Lomax, Illinois, to Lyman Neff and Emma Jacobson. A sister named Jenona had been born a year earlier. Dewey’s father, like his own father and brothers, was a skilled carpenter, but for 263 

264 BIOGRAPHICAL MEMOIRS much of his adult life he worked as a lumberyard manager. The family spent the first 10 years of Dewey’s life in Lomax, a small town on the Illinois side of the Mississippi River between Burlington, Illinois, and Fort Madison, Iowa. After short stays in Yates City and Brereton, Illinois, and a trip to South Dakota, the family settled in Freeport, Illinois, where the children graduated from the local high school (Jenona in 1929 and Dewey in 1930). In a privately published mem- oir (L. M. Neff, 1969) their father commented on the fact that the children attended schools in many different towns: “I always thought [this] was good for them. They learned to meet strangers and adapt themselves to changing condi- tions.” Dewey entered the University of Illinois in 1930 with the intention of becoming an architect. In those depres- sion years he had to work his way through college, includ- ing spending one year working full time. As a result, it took him six years to complete his undergraduate education. THE GRADUATE YEARS During his undergraduate years, Dewey came under the influence of Elmer A. Culler, a professor of psychology, who persuaded him to continue as a graduate student in experimental psychology. Culler was a well-known physiologi- cal psychologist, whose research combined studies of ani- mal learning and hearing. When Culler moved from Illi- nois to the University of Rochester, he persuaded Dewey to move with him. During his stay at Rochester, Dewey made lasting friendships with fellow graduate students Karl Kryter and J. C. R. Licklider, both of whom went on to distin- guished careers of their own. When Dewey began his graduate work, the dominant question facing hearing researchers was how tonal frequen- 

WILLIAM DUWAYNE NEFF 265 cies were represented in the cochlea and in central audi- tory pathways. Dewey was intrigued by a report from the Johns Hopkins University neurosurgeon, Walter E. Dandy (Dandy, 1934) on the hearing losses resulting from inad- vertent lesions to the auditory nerve when the vestibular nerve was cut to relieve the symptomss of Ménière’s dis- ease. Hearing could be impaired at high frequencies, but never at low frequencies. Dewey realized the potential import of Dandy’s find- ings for the place theory of hearing, in which near-thresh- old tones of any particular frequency were supposed to acti- vate only a restricted and specific set of auditory nerve fibers. Therefore, the question arose whether, in fact, Dandy had ever sectioned the hypothesized low-frequency fibers. The lack of histological controls prevented the question from being resolved in humans. For his doctoral dissertation, Dewey partially cut the auditory nerve in several cats and, compared their hearing preoperatively and posoperatively. The behavioral testing was based on a method developed by Brogden and Culler (Brogden and Culler, 1936). In ad- dition, cochlear electrophysiology was performed at the time of sacrifice and postmortem histology was examined in col- laboration with Prof. E. G. Wever of Princeton University. Dewey’s results in cats were similar to those of Dandy’s in humans. Partial section of the auditory nerve could pro- duce a selective high-frequency hearing loss, but never one confined to low frequencies. The histology provided a ra- tionale for the behavioral findings. In all animals, nerve fiber degeneration started at the cochlear base, where high frequencies are represented, and extended for varying dis- tances toward the apex, the region most sensitive to low frequencies. There was no instance of restricted apical de- generation. From electrophysiology Dewey concluded that cochlear microphonics, which are now recognized to arise 

266 BIOGRAPHICAL MEMOIRS from hair cells, could be spared even in the virtual absence of innervating nerve fibers. World War II delayed full publi- cation of the research until 1947. (Neff, 1947; Wever and Neff, 1947). Dewey’s results left two questions. (1) What would be the consequence of cochlear lesions restricted to the apex? (2) Why did partial sections of the auditory nerve in his experiments never produce selective apical nerve degen- eration? In an attempt to answer the first question, Schu- knecht and Neff (Schuknecht and Neff, 1952) found that cochlear lesions confined to the apex resulted in modest hearing losses limited to low frequencies. A potential an- swer to the second question was suggested by Kiang (Kiang, 1979) at a conference organized by Dewey’s graduate stu- dents and postdoctoral trainees to honor him. Kiang specu- lated that Dewey’s lesions were too medial. As such, they passed through the cochlear nucleus, rather than the audi- tory nerve and were located so as to sever the high-fre- quency innervation of the nucleus, but not the more later- ally situated low-frequency innervation. WARTIME Following completion of his graduate work, Dewey was appointed as a research associate at Swarthmore College, where he collaborated with the famous Gestalt psychologist Wolfgang Köhler. The attack on Pearl Harbor occurred shortly after the start of Dewey’s second year at Swarthmore. Within the month Dewey resigned his position and joined the Underwater Sound Laboratories in New London, Con- necticut, where he worked on the selection and training of submarine crews (Cronbach and Neff, 1949) and auditory discrimination in sonar detection (Neff and Thurlow, 1949). 

WILLIAM DUWAYNE NEFF 267 THE UNIVERSITY OF CHICAGO In 1946 Dewey joined the faculty of the Department of Psychology at the University of Chicago. This was an excit- ing time at the university. A team led by Enrico Fermi had made important contributions to the Manhattan Project, including the first demonstration of a sustained nuclear reaction. Immediately after the war, Edward Teller returned to the University from Los Alamos and Harold Urey, a Nobel laureate in chemistry, came from Columbia University. Paul Weiss, a professor of zoology, was doing pioneering studies in developmental neurobiology, including the first demon- stration of axoplasmic flow. Roger Sperry, having already done his limb re-innervation and optic tectum studies in Weiss’s laboratory, was returning to the university after a postdoctoral fellowship with Karl Lashley at Orange Park. Ralph W. Gerard was a professor of physiology in whose laboratory pioneering work with microelectrodes was tak- ing place. Kenneth S. Cole, having made fundamental ob- servations on action potentials in the giant squid axon, was appointed a professor of biophysics and was perfecting elec- tronic methods needed to control the voltage and current across the nerve membrane. Heinrich Klüver and Stephen Polyak were located on the top floor of Culver Hall, a musty old building. Klüver was continuing his studies of the be- havioral effects of temporal lobe lesions in monkeys—the Klüver-Bucy syndrome—while Polyak, having completed his monumental monograph on the vertebrate retina, was now working on an even more ambitious monograph dealing with the entire visual system. At the same time, a different breed of graduate stu- dent was arriving on campus: war veterans financed by the GI Bill. Compared with graduate students before or since, they were older and more mature, sobered by their war 

268 BIOGRAPHICAL MEMOIRS experiences. Many were married and already raising fami- lies. They had sublimated their personal agendas to the war effort and now wished to finish graduate school and begin their professional careers. On arriving at the University of Chicago, Dewey taught a summer course in physiological psychology attended by some of these students. Here Dewey could expound his view that a multidisciplinary approach, then practiced in only a few laboratories, was required to make progress in hearing research and, more generally, in brain research. His own doctoral research had exemplified this perspec- tive, combining behavioral testing, electrophysiology, and neuroanatomy. In addition, Dewey was of the opinion that to understand the functions of higher auditory centers, one had to expand the behavioral testing of animals beyond the measurement of hearing thresholds. Here he was inspired by work in the visual system, especially that done by Lashley and Klüver. Dewey and the other psychologists doing animal re- search were assigned laboratory space in a cockroach-in- fested, two-story, prefabricated wooden structure, officially known as the Laboratory of Physiological Psychology, but commonly referred to as the “Prefabs.” Dewey set up sound- proofed experimental chambers to test animal hearing and histological facilities to prepare tissue for microscopic evalu- ation of lesions. He established an electrophysiological labo- ratory in space belonging to the Section of Otolaryngology in Billings Hospital. This was part of a continuing collabo- ration with otolaryngologists in the Surgery Department, including Chief of Otolaryngology John Lindsay, Henry Perlman and Harold Schuknecht (Lindsay et al., 1953; Schuknecht et al., 1951). Dewey’s work with Schuknecht on selective cochlear lesions has already been mentioned and was the start of a lifelong friendship. 

WILLIAM DUWAYNE NEFF 269 As was the case for his mentor, Elmer Culler, much of Dewey’s otolaryngology-related research was partly funded through various otological societies. More importantly, Dewey was able to use contacts he had made during his wartime service to secure funding from the Office of Naval Research (ONR). Nowadays most funding for neuroscience research comes from the National Institutes of Health (NIH) or from the National Science Foundation (NSF). The NSF was founded in 1950, and the Extramural Program of the NIH, although started in 1946, became an important source of funding only in the late 1950s. In the early postwar years the Defense Department became a major source of support for university research, and the ONR was a major source of funding for Dewey’s work. Many of the war veterans who were graduate students in psychology chose to do their doctoral research in Dewey’s laboratory because he exuded a quiet confidence and dis- played a generosity of spirit. He seemed to be a person who knew where he was going and would be happy for others to accompany him. In addition, his multidisciplinary approach to science had an intellectual appeal. With a coherent and, for its time, a well-funded research program, Dewey’s labo- ratory appealed to prospective doctoral candidates as a place where they could successfully learn their craft. As one of his graduate students, Irving Diamond, said at a reunion some 45 years later: “You knew when you came to work in Dewey’s lab that you were joining a winning team.” As the last of the war veterans finished their graduate studies, Dewey was able to recruit younger, more traditional graduate stu- dents. By the time Dewey left the University of Chicago in 1961, he had successfully supervised the doctoral research of over 20 students. There were usually several graduate students working with Dewey at any one time. Concentrating on ablation stud- 

270 BIOGRAPHICAL MEMOIRS ies, Dewey and each student would settle on a thesis topic. It was then the student’s responsibility to do the necessary behavioral testing, usually in cats, but in some cases in rhesus monkeys. Audiometric testing was done in a rotating cage, the device devised by Brogden and Culler. For more sophis- ticated auditory discriminations (e.g., tonal frequency or tonal pattern discriminations) animals were trained in a double-grill box, which required that the animals run be- tween two compartments to avoid shock. Sound localiza- tion was studied in a semicircular arena in which the ani- mal was trained, when released from a restraining box, to go to one of two food wells behind which a buzzer had sounded before release. Once an animal had reached a high level of performance, an ablation of some selected part of the auditory system was done under sterile surgical conditions. Dewey usually did the surgery with the student assisting. After a postoperative recovery, typically a few weeks to a month, the animals were retested and, if necessary, retrained to see if they could once again do the discrimina- tive task. Following postoperative testing, an attempt was made to assess the extent of the lesion with evoked-poten- tial methods, after which the animal was sacrificed and the brain was fixed by transcardiac perfusion as the first step in histological processing. The main topic of interest for Neff and his students was the function of the auditory cortex, although work was also done on lower levels of the auditory pathways. The extent of cortical lesions was based on electrophysiological mapping done by Clinton Woolsey and his colleagues (Woolsey, 1960), among others. A major conclusion of the research was that animals were still able to make simple discriminations after removal of all known auditory cortical areas. This had already been suggested by Harlow Ades and his colleagues, who showed that hearing thresholds (Kryter 

WILLIAM DUWAYNE NEFF 271 and Ades, 1943) and intensity differential limens (Raab and Ades, 1946) were unaltered in animals after cortical abla- tions. Work in Dewey’s group showed that frequency dis- criminations could be relearned after very large cortical removals (Butler et al., 1957), but the ability to perform more complex tasks, including tonal pattern discriminations (Diamond and Neff, 1957; Diamond et al., 1962) and sound localization (Neff et al., 1956) were permanently compro- mised. Even the simpler tasks, which could presumably be accomplished by lower auditory centers, had to be relearned after the cortical lesions. The results demonstrated that in the absence of corti- cal receiving areas, the auditory system still has impressive capabilities. That trained discriminations were lost after cor- tical removals might imply that cortical processing has a ubiquitous role when the entire system is intact. The defini- tion of that precise role still remains elusive. In addition, even permanent deficits may be related to the behavioral tasks used to demonstrate discriminative ability, rather than to the sensory discrimination per se. For example, later work showed that animals, after cortical removals, are ca- pable of indicating the location of a sound by unlearned orientation head movements (Thompson and Welker, 1963; Poon, 1979) or simple learned responses (Heffner and Masterton, 1975) even though they are unable to walk to- ward the position of a previously presented sound (Neff et al., 1956; Heffner and Masterton, 1975). It remains unre- solved as to whether this last deficit reflects a defect in sensorimotor integration, short-term memory, or attentive- ness/distractibility. Consistent with his belief in a multidisciplinary approach to the neurosciences, Dewey was instrumental in the found- ing in 1955 of the degree-granting Section of Biopsychol- ogy in the Biological Sciences Division. To be enrolled in 

272 BIOGRAPHICAL MEMOIRS the Biopsychology Section, graduate students had first to be admitted by the Psychology Department, which was in the Social Sciences Division. Once enrolled, the student’s curriculum and research had a large biological sciences com- ponent and the Ph.D. was awarded in the Division of Bio- logical Sciences. The philosophy of the program as stated in the university announcements was as follows: It is expected that the research [person] trained in biopsychology may . . . devote . . . attention to problems falling between the discipline of psychol- ogy and the traditional disciplines, such as physiology, anatomy and zool- ogy. The student is expected, therefore, to become substantially grounded in the physical and biological sciences, in addition to the general field of psychology. Emphasis is placed . . . upon understanding the role of neural and extra-neural mechanisms of the internal environment in the regula- tion of behavior. This nascent neurosciences program served as the focus for the training of his graduate students while Dewey re- mained at Chicago and continued well into the 1990s. BOSTON After spending 15 years in Chicago, Dewey was persuaded to move in 1961 to Bolt, Beranek and Newman (BBN), a high-technology company founded in 1948 by two Massa- chusetts Institute of Technology (MIT) professors in acous- tics, Richard Bolt and Leo Beranek, and a former student of theirs, Robert Newman. Originally an acoustical consult- ing company, BBN obtained many prestigious commissions in architectural acoustics, including the innovative Kresge Auditorium at MIT, the Shed at Tanglewood, and Avery Fisher Hall at Lincoln Center. By the late 1950s the com- pany was looking to diversify. Dewey’s graduate student friends Karl Kryter and J. C. R. Licklider had been recruited to the company. Both had changed their interests from their original training in auditory physiology and psychophysics. Kryter 

WILLIAM DUWAYNE NEFF 273 had become an expert in the effects of noise on human performance (Kryter, 1970) while Licklider grew interested in the relation between humans and computers and is cred- ited with helping to develop computer networking, which eventually led to the Internet (Waldrop, 2001). In recruiting Dewey the company had hoped to ex- pand into the field of auditory physiology. Dewey brought with him two of his recent graduate students, Norman Strominger and Philip Nieder. They set up an experimen- tal suite with a surgery, animal facilities, and testing rooms, and began several promising experiments. Unfortunately, at this time the federal government decided not to support basic research at commercial companies. Since Dewey’s re- search in a corporate environment depended on federal funding, it was clear to him that he had to move on. In 1963 he welcomed the opportunity to join the faculty of Indiana University at Bloomington. The next year he was made director of the newly formed Center for Neural Sci- ences. INDIANA UNIVERSITY In his new position Dewey faced two challenges: (1) to build the Center for Neural Sciences and (2) to reestablish his research program. Dewey attracted several people to the center: Jorgen Fex, an auditory physiologist who pio- neered the study of auditory efferents, came from the Karolinska Institute after stints in Australia and the Na- tional Institutes of Health; Conrad Mueller, a visual scien- tist best known for his work on Limulus eye, came from Columbia University; Willem van Bergeijk, a physicist inter- ested in the evolution of hearing, arrived from Bell Labora- tories; Ilsa Schwartz, an auditory neuroanatomist, came from a postdoctoral fellowship at Albert Einstein Medical School; and Boyd Campbell, a comparative neuroanatomist, moved 

274 BIOGRAPHICAL MEMOIRS from Walter Reed Army Medical Center. Through no fault of Dewey’s, the center did not flourish. Van Bergeijk met an untimely death. Fex left to become director of the Labo- ratory of Neuro-otology at the National Institutes of Health. Schwartz left to take an offer at the University of California, Los Angeles. Campbell left to resume clinical training. Of the people Dewey recruited, only Mueller remained at Indi- ana University. Dewey did resume his own research. With his first Indi- ana graduate student, J. I. (“Pete”) Casseday, he studied the role of acoustic commissures in sound localization. Build- ing on the graduate research of Colston Nauman Moore, done at Chicago, they were able to show that sound local- ization was unperturbed by section of the corpus callosum or the commissure of the inferior colliculus, but was im- paired by section of the trapezoid body, the main fiber bundle allowing for the integration of binaural inputs in the superior olivary complex (Moore et al., 1974; Casseday and Neff, 1975). Two other graduate students worked with Dewey on studies of relatively simple auditory behaviors. Paul Poon confirmed that animals could indicate the loca- tion of a sound by unlearned orientation head movements (Poon, 1979), while C.-K. (“Joseph”) Chan defined some of the brain stem pathways involved in various acoustic re- flexes (Chan, 1983). Dewey summarized his own studies on auditory cortex in a brief review (Neff, 1977) and wrote a comprehensive survey of behavioral studies in the auditory system (Neff et al., 1975). NATIONAL SERVICE AND AWARDS In addition to his wartime service Dewey served for a year (1953-1954) in London as a scientific liaison officer for the Office of Naval Research. He was a long-standing 

WILLIAM DUWAYNE NEFF 275 member of the Committee on Hearing and Bioacoustics of the National Research Council, as well as a member of the Executive Council of the Acoustical Society of America and of the Psychonomics Society. He served on the Editorial Boards of the Journal of Neurophysiology and the Annual Review of Psychology. Along with Wolf Keidel, he edited volume V (“Auditory System”) of the Handbook of Sensory Physiology. He was the sole editor of a series, Contributions to Sensory Physiology, which published reviews on sensory processing; eight volumes were published between its found- ing in 1965 and 1983. In recognition of his contributions, Dewey was elected to the National Academy of Sciences in 1964 and the American Academy of Arts and Sciences. He received the Annual Award of the Beltone Institute for Hear- ing Research and the Award of Merit of the Association for Research in Otolaryngology. IN SUMMARY During Dewey’s professional career, much of experi- mental psychology moved from emphasizing speculative doctrines, as exemplified by various learning theories, to obtaining more empirical descriptions of the brain, behav- ior, and the relation between the two. He and his students never established a school of thought wedded to specific dogmas but concentrated on obtaining rigorous experimental results with proper controls and making limited generaliza- tions. A list of his graduate students at Chicago and Indi- ana and the titles of their Ph.D. dissertations are included at the end of this memoir, and it will be evident that they moved in many directions, contributing in diverse ways to the neurosciences. On balance, one of Dewey’s most lasting contributions may be his students. He created an environment where young people could find themselves and develop their individual 

276 BIOGRAPHICAL MEMOIRS strengths. He was patient, encouraging, and available to members of his laboratory including students, technical staff, and visitors. The comforting atmosphere universally felt by the people who worked with him was directly related to his social skill in providing education under the guise of friendly chats. He was able to attract some of the best students avail- able and they always benefited from his relaxed yet rigor- ous instructional style. These personal qualities also made him a highly regarded, informed advisor to his peers and to a variety of scientific organizations. Dewey had a daughter, Carol, and a son, Peter, by his first wife, Ernestine, a professor of English, and a total of five grandchildren. His second wife, Palmer, contributed to auditory research by studying the effects of muscular dys- trophy on the middle-ear reflexes. His professional family was large and his influence remains strong on all who were fortunate enough to know him. DOCTORAL DISSERTATIONS OF STUDENTS TRAINED BY W. D. NEFF UNIVERSITY OF CHICAGO Arnott, G. P. Impairment following ablation of the primary and secondary areas of the auditory cortex, 1953. Butler, R. A. The role of the auditory cortex in frequency discrimi- nation, 1951. Diamond, I. T. The function of the auditory cortex: The effect of ablation of the auditory cortex on the discrimination of tempo- ral frequency patterns, 1953. Gerken, G. M. The evoked electrocortical response to acoustic stimu- lation and its relation to behavioral conditioning, 1959. Goldberg, J. M. Frequency discrimination after central nervous sys- tem lesions, 1960. Hahn, J. F. Effects of partial ablations of visual areas I and II on visual form discrimination in the cat, 1952. Hind, J. E. An electrophysiological study of tonotopic organization in the auditory cortex of the cat, 1952. 

WILLIAM DUWAYNE NEFF 277 Jerison, H. J. Effect of auditory cortex ablation on tone pattern discrimination in the rhesus monkey, 1954. Kennedy, T. T. K. An electrophysiological study of the auditory projection areas of the cortex in monkey (Macaca mulatta), 1955. Kiang, N. Y.-S. An electrophysiological study of cat auditory cortex, 1955. Lindquist, S. E. Stimulation deafness: A study of temporary and permanent hearing losses resulting from exposure to noise and to blast impulses, 1949. Nauman, C. Sound localization: The role of the commissural path- ways of the auditory system of the cat, 1958. Nieder, P. C. Electrophysiological recording and stimulation stud- ies of the auditory system of unanesthetized cats, 1960. Odoi, H. An investigation into the functions of the visual cortical areas in the cat, 1951. Oesterreich, R. E. The role of higher neural auditory centers in the discrimination of differential sound intensities, 1960. Rosner, B. S. Higher auditory nervous centers, 1950. Stoughton, G. S. S. The role of the cortex in retention of a condi- tioned response to auditory signals, 1954. Strominger, N. L. Localization of sound after central nervous sys- tem lesions, 1961. Sutton, S. Threshold changes for evoked cortical potentials follow- ing cochlear lesions, 1955. Wegener, J. G. The role of special cerebral and cerebellar systems in sensory discrimination, 1954. INDIANA UNIVERSITY Casseday, J. H. Auditory localization: The role of the brainstem auditory pathways of the cat, 1970. Chan, C.-K. Neural centers and pathways involved in startle, orient- ing, and middle-ear reflexes to acoustic stimuli, 1983. 

278 BIOGRAPHICAL MEMOIRS Poon, P. W. F. Cortical centers and midbrain pathways involved in sound localization in space, 1979. REFERENCES Brogden, W. J., and E. A. Culler. 1936. Device for the motor condi- tioning of small animals. Science 83:269-270. Butler, R. A., I. T. Diamond, and W. D. Neff. 1957. Role of auditory cortex in discrimination of changes in frequency. J. Neurophysiol. 20:108-120. Casseday, J. H., and W. D. Neff. 1975. Auditory localization: role of auditory pathways in the brain stem of the cat. J. Neurophysiol. 38:842-858. Chan, C.-K. 1983. Neural centers and pathways involved in startle, orienting, and middle-ear reflexes to acoustic stimuli. Doctoral dissertation: Center for Neural Sciences, Indiana University, Bloomington, Ind. Cronbach, L. J., and W. D. Neff. 1949. Selection and training. In A Survey Report on Human Factors in Undersea Warfare, pp. 491- 513. Washington, D.C.: Committee on Undersea Warfare. Na- tional Research Council. Dandy, W. E. 1934. Effects on hearing after subtotal section of the cochlear branch of the auditory nerve. Bull. Johns Hopkins Hosp. 55:240-243. Diamond, I. T., and W. D. Neff. 1957. Ablation of temporal cortex and discrimination of auditory patterns. J. Neurophysiol. 20:300- 315. Diamond, I. T., J. M. Goldberg, and W. D. Neff. 1962. Tonal dis- crimination after ablation of auditory cortex. J. Neurophysiol. 25:223-235. Heffner, H., and B. Masterton. 1975. Contribution of auditory cor- tex to sound localization in the monkey (Macaca mulatta). J. Neurophysiol. 38:1340-1358. Kiang, N. Y.-S. 1979. A reexamination of “the effects of partial sec- tion of the auditory nerve.” In Festschrift for W. D. Neff, eds. J. M. Goldberg and R. A. Butler, pp 46-71. Chicago, Ill.: Privately published. Kryter, K. D. 1970. The Effects of Noise on Man. New York: Aca- demic Press. 

WILLIAM DUWAYNE NEFF 279 Kryter, K. D., and H. W. Ades. 1943. Studies on the function of the higher acoustic nerve centers in the cat. Am. J. Psychol. 56:501- 536. Lindsay, J. R., W. D. Neff, H. F. Schuknecht, and H. B. Perlman. 1953. Obliteration of the ductus endolymphaticus and their ac- companying venous channels. Acta Otolaryngol. 43:176-189. Moore, C. N., J. H. Casseday, and W. D. Neff. 1974. Sound localiza- tion: Role of the commissural pathways of the auditory system of the cat. Brain Res. 82:13-26. Neff, L. M. 1969. A Neff history and autobiography: Privately pub- lished. Neff, W. D. 1947. The effects of partial section of the auditory nerve. J. Comp. Physiol. Psychol. 40:203-215. Neff, W. D. 1977. The brain and hearing: Auditory discrimination affected by brain lesions. Ann. Otol. Rhinol. Laryngol. 86:500- 506. Neff, W. D., and W. R. Thurlow. 1949. Auditory discrimination in sonar operation. In A Survey Report on Human Factors in Un- dersea Warfare, pp. 219-230. Washington, D.C.: Committee on Undersea Warfare. National Research Council. Neff, W. D., I. T. Diamond, and J. H. Casseday. 1975. Behavioral studies of auditory discrimination: Central nervous system. In Handbook of Sensory Physiology, vol. V/2, eds. W. D. Keidel and W. D. Neff, pp. 307-400. Berlin: Springer-Verlag. Neff. W. D., J. F. Fisher, I. T. Diamond, and M. Yela. 1956. Role of auditory cortex in discrimination requiring localization of sound in space. J. Neurophysiol. 19:500-512. Poon, P. W. F. 1979. Cortical centers and midbrain pathways in- volved in sound localization in space. Doctoral dissertation: Cen- ter for Neural Sciences, Indiana University, Bloomington, Ind. Raab, D. H., and H. W. Ades. 1946. Cortical and midbrain media- tion of a conditioned discrimination of acoustic intensities. Am. J. Psychol. 59:59-83. Schuknecht, H. F., and W. D. Neff. 1952. Hearing losses after apical lesions in the cochlea. Acta Oto-laryngol. 42:263-274. Schuknecht, H. F., W. D. Neff, and H. B. Perlman. 1951. An experi- mental study of auditory damage following blows to the head. Ann. Otol. Rhinol. Laryngol. 60:273-289. Thompson, R. F., and W. I. Welker. 1963. Role of auditory cortex in 

280 BIOGRAPHICAL MEMOIRS reflex head orientation by cats to auditory stimuli. J. Comp. Physiol. Psychol. 56:996-1002. Waldrop, M. M. 2001. The Dream Machine. New York: Viking/Pen- guin Putnam. Wever, E. G., and W. D. Neff. 1947. A further study of the effects of partial section of the auditory nerve. J. Comp. Physiol. Psychol. 40:217-226. Woolsey, C. N. 1960. Organization of cortical auditory system: A review and a synthesis. In Neural Mechanisms of the Auditory and Vestibular Systems, eds. G. L. Rasmussen and W. F. Windle, pp. 165-180. Springfield, Ill.: Thomas. 

WILLIAM DUWAYNE NEFF 281 SELECTED BIBLIOGRAPHY 1947 The effects of partial section of the auditory nerve. J. Comp. Physiol. Psychol. 40:203-215. With E. G. Wever. A further study of the effects of partial section of the auditory nerve. J. Comp. Physiol. Psychol. 40:217-226. 1949 With L. J. Cronbach. Selection and training. In A Survey Report on Human Factors in Undersea Warfare, pp. 491-513. Washington, D.C.: Committee on Undersea Warfare. National Research Coun- cil. With W. R. Thurlow. Auditory discrimination in sonar operation. In A Survey Report on Human Factors in Undersea Warfare, pp. 219-230. Washington, D.C.: Committee on Undersea Warfare. National Research Council. 1951 With H. F. Schuknecht and H. B. Perlman. An experimental study of auditory damage following blows to the head. Ann. Otol. Rhinol. Laryngol. 60:273-289. 1952 With H. F. Schuknecht. Hearing losses after apical lesions in the cochlea. Acta Oto-laryngol. 42:263-274. 1954 With S. E. Lindquist and H. F. Schuknecht. Simulation deafness: A study of hearing losses resulting from exposure to noise or to blast impulses. J. Comp. Physiol. Psychol. 47:406-411. 1956 With I. T. Diamond, J. F. Fisher, and M. Yela. Role of auditory cortex in discrimination requiring localization of sound in space. J. Neurophysiol.19: 500-512. 

282 BIOGRAPHICAL MEMOIRS 1957 With R. A. Butler and I. T. Diamond. Role of auditory cortex in discrimination of changes in frequency. J. Neurophysiol. 20:108- 120. With I. T. Diamond. Ablation of temporal cortex and discrimina- tion of auditory patterns. J. Neurophysiol. 20:300-315. 1958 With I. T. Diamond and K. L. Chow. Degeneration of caudal medial geniculate body following cortical lesion ventral to auditory area II in the cat. J. Comp. Neurol. 109:349-362. 1961 With J. M. Goldberg. Frequency discrimination after bilateral abla- tion of cortical auditory areas. J. Neurophysiol. 24:119-128. With J. M. Goldberg. Frequency discrimination after bilateral sec- tion of the brachium of the inferior colliculus. J. Comp. Neurol. 116:265-289. With P. C. Nieder. Auditory information from subcortical electrical stimulation in cats. Science 133:1010-1011. 1962 With I. T. Diamond and J. M. Goldberg. Tonal discrimination after ablation of auditory cortex. J. Neurophysiol. 25:223-235. 1963 With G. M. Gerken. Experimental procedures affecting evoked re- sponses recorded from auditory cortex. Electroenceph. Clin. Neurophysiol. 15:947-957. 1965 With D. P. Scharlock and N. L. Strominger. Discrimination of tone duration after bilateral ablation of cortical auditory areas. J. Neurophysiol. 28:673-681. 1971 With R. E. Oesterreich and N. L. Strominger. Neural structures mediating differential sound intensity discrimination in the cat. Brain Res. 27:251-270. 

WILLIAM DUWAYNE NEFF 283 1974 With C. N. Moore and J. H. Casseday. Sound localization: The role of the commissural pathways of the auditory system of the cat. Brain Res. 82:13-26. 1975 With J. H. Casseday. Auditory localization: Role of auditory path- ways in brain stem of the cat. J. Neurophysiol. 38:842-858. With T. R. Dolan, H. W. Ades, and G. Bredberg. Inner ear damage and hearing loss after exposure to tones of high intensity. Acta Otolaryngol. 80:343-352. 1976 With J. L. Cranford, S. J. Ladner, and C. B. Campbell. Efferent projections of the insular and temporal neocortex of the cat. Brain Res. 117:195-210. 1977 With J. H. Casseday. Effects of unilateral ablation of auditory cortex on monaural cat’s ability to localize sound. J. Neurophysiol. 40:44- 52. 1980 With N. L. Strominger and R. E. Oesterreich. Sequential auditory and visual discriminations after temporal lobe ablation in mon- keys. Physiol. Behav. 24:1149-1156.