Learning and Memory 53During 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 MEMORYKarl 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- 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. 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