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Issue 1060 coverThe Neurosciences and Music II: From Perception to Performance Volume 1060 published December 2005
Ann. N.Y. Acad. Sci. 1060: 148–174 (2005). doi: 10.1196/annals.1360.011
Copyright © 2005 by the New York Academy of Sciences
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Articles by TRAMO, M. J.
Articles by BRAIDA, L. D.
Neurophysiology and Neuroanatomy of Pitch Perception: Auditory Cortex

MARK JUDE TRAMOa,b,c, PETER A. CARIANIa,e, CHRISTINE K. KOHa,b, NIKOS MAKRISa,d AND LOUIS D. BRAIDAb

aDepartment of Neurology, Harvard Medical School, and Massachusetts General Hospital; The Institute for Music and Brain Science, Auditory Neuroscience Program, Boston, Massachusetts 02114, USA
bSensory Communication Group, Research Laboratory of Electronics, Massachusetts Institute of Technology, Cambridge, Massachusetts 02139, USA
cEaton-Peabody Laboratory of Auditory Physiology, Massachusetts Eye and Ear Infirmary, Boston, Massachusetts 02114, USA
dCenter for Morphometric Analysis, Martinos Center for Biomedical Imaging, Massachusetts General Hospital, Charlestown, Massachusetts 02129, USA
eDepartment of Physiology, Tufts Medical School, Boston, Massachusetts 02111 USA

Address for correspondence: Mark Jude Tramo, M.D., Ph.D., Director, The Institute for Music and Brain Science, 175 Cambridge Street, Suite 340, Boston, MA 02114. Voice: 617-726-5409. mtramo{at}hms.harvard.edu; http://www.brainmusic.org

We present original results and review literature from the past fifty years that address the role of primate auditory cortex in the following perceptual capacities: (1) the ability to perceive small differences between the pitches of two successive tones; (2) the ability to perceive the sign (i.e., direction) of the pitch difference [higher (+) vs. lower (–)]; and (3) the ability to abstract pitch constancy across changes in stimulus acoustics. Cortical mechanisms mediating pitch perception are discussed with respect to (1) gross and microanatomical distribution; and (2) candidate neural coding schemes. Observations by us and others suggest that (1) frequency-selective neurons in primary auditory cortex (A1) and surrounding fields play a critical role in fine-grained pitch discrimination at the perceptual level; (2) cortical mechanisms that detect pitch differences are neuroanatomically dissociable from those mediating pitch direction discrimination; (3) cortical mechanisms mediating perception of the "missing fundamental frequency (F0)" are neuroanatomically dissociable from those mediating pitch perception when F0 is present; (4) frequency-selective neurons in both right and left A1 contribute to pitch change detection and pitch direction discrimination; (5) frequency-selective neurons in right A1 are necessary for normal pitch direction discrimination; (6) simple codes for pitch that are based on single- and multiunit firing rates of frequency-selective neurons face both a "hyperacuity problem" and a "pitch constancy problem"—that is, frequency discrimination thresholds for pitch change direction and pitch direction discrimination are much smaller than neural tuning curves predict, and firing rate patterns change dramatically under conditions in which pitch percepts remain invariant; (7) cochleotopic organization of frequency-selective neurons bears little if any relevance to perceptual acuity and pitch constancy; and (8) simple temporal codes for pitch capable of accounting for pitches higher than a few hundred hertz have not been found in the auditory cortex. The cortical code for pitch is therefore not likely to be a function of simple rate profiles or synchronous temporal patterns. Studies motivated by interest in the neurophysiology and neuroanatomy of music perception have helped correct longstanding misconceptions about the functional role of auditory cortex in frequency discrimination and pitch perception. Advancing knowledge about the neural coding of pitch is of fundamental importance to the future design of neurobionic therapies for hearing loss.

Key Words: pitch • missing fundamental • psychophysics • periodicity • autocorrelation • auditory cortex • lesion effects • neural coding






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