Course Organization and Overview



Dorsal Cochlear Nucleus



Oertel, D., and E. D. Young. "What's a cerebellar circuit doing in the auditory system?" Trends Neurosci 27 (2004): 104-110.

Young, E. D., and K. A. Davis. "Circuitry and function of the dorsal cochlear nucleus." In Integrative Functions of the Mammalian Auditory Pathway. Edited by D. Oertel, A. N. Popper, and R. R. Fay. New York, NY: Springer, 2002. ISBN: 9780387989037.

Papers for Discussion

May, B. J. "Role of the dorsal cochlear nucleus in the sound localization behavior of cats." Hear Res 148 (2000): 74-87.

Reiss, L. A., and E. D. Young. "Spectral edge sensitivity in neural circuits of the dorsal cochlear nucleus." J Neurosci 25 (2005): 3680-3691.

Kanold, P. O., and E. D. Young. "Proprioceptive information from the pinna provides somatosensory input to cat dorsal cochlear nucleus." J Neurosci 21 (2001): 7848-7858.

Tzounopoulos, T., Y. Kim, D. Oertel, and L. O. Trussell. "Cell-specific, spike timing-dependent plasticities in the dorsal cochlear nucleus." Nat Neurosci 7 (2004): 719-725.

Bell, C. C. "Memory-based expectations in electrosensory systems." Curr Opin Neurobiol 11 (2001): 481-487.

Levine, R. A. "Somatic (craniocervical) tinnitus and the Dorsal Cochlear Nucleus hypothesis." Am J Otolaryngol 20 (1999): 351-362.

Additional Readings

Manis, P. B. "Membrane properties and discharge characteristics of guinea pig dorsal cochlear nucleus neurons studied in vitro." J Neurosci 10 (1990): 2338-2351.

Nelken, I., and E. D. Young. "Two separate inhibitory mechanisms shape the responses of dorsal cochlear nucleus Type IV units to narrowband and wideband stimuli." J Neurophysiol 71 (1994): 2446-2462.

Hancock, K. E., and H. F. Voigt. "Wideband inhibition of dorsal cochlear nucleus Type IV units in cat: A computational model." Ann Biomed Eng 27 (1999): 73–87.

Brozoski, T. J., C. A. Bauer, and D. M. Caspary. "Elevated fusiform cell activity in the dorsal cochlear nucleus of chinchillas with psychophysical evidence of tinnitus." J Neurosci 22 (2002): 2383-2390.


Descending Systems



Guinan, J. J., Jr. "The physiology of olivocochlear efferents." In The Cochlea. Edited by P. Dallos, A. N. Popper, and R. R. Fay. New York, NY: Springer-Verlag, 1996, pp. 435-502. ISBN: 9780387944494.

Spangler, K. M., and W. B. Warr. "The descending auditory system." In Neurobiology of Hearing: The Central Auditory System. Edited by R. A. Altschuler, R. P. Bobbin, B. M. Clopton, and D. W. Hoffman. New York, NY: Raven Press, 1991, pp. 27-45. ISBN: 9780881678062.

Papers for Discussion

Venecia, R. K. de, M. C. Liberman, J. J. Guinan, Jr., and M. C. Brown. "Medial olivocochlear reflex interneurons are located in the posteroventral cochlear nucleus." J Comp Neurol 487 (2005): 345-360.

Mulders, Wham, I. M. Winter, and D. Robertson. "Dual action of olivocochlear collaterals in the guinea pig cochlear nucleus." Hearing Res 174 (2002): 264-280.

May, B. J., C. A. Prosen, D. Weiss, and D. Vetter. "Behavioral investigation of some possible effects of central olivocochlear pathways in transgenic mice." Hearing Res 171 (2002): 142-157.

Further Reading

Suga, N. "The corticofugal system for hearing: Recent progress." PNAS 97 (2000): 11807-11814.

May, B. J., and S. J. McQuone. "Effects of bilateral olivocochlear lesions on pure-tone intensity discrimination in cats." Auditory Neurosci 1 (1995): 385-400.

Khalfa, S., R. Bougeard, N. Morand, E. Veuillet, J. Isnard, M. Guenot, P. Ryvlin, C. Fischer, and L. Collet. "Evidence of peripheral auditory activity modulation by the auditory cortex in humans." Neurosci 104 (2001): 347-358.

Mulders, Wham, and D. Robertson. "Evidence for direct cortical innervation of medial olivocochlear neurones in rats." Hearing Res 144 (2000): 65-72.


Cell Types and Circuits



Oliver, D. L. "Ascending efferent projections of the superior olivary complex." Microsc Res Tech 51 (2000): 355-63.

Papers for Discussion

Adams, J. C., and E. Mugnaini. "Immunocytochemical evidence for inhibitory and disinhibitory circuits in the superior olive." Hearing Res 49 (1990): 281-298.

Loftus, W. C., D. C. Bishop, R. L. Saint Marie, and D. L. Oliver. "Organization of binaural excitatory and inhibitory inputs to the inferior colliculus from the superior olive." J Comp Neurol 472 (2004): 330-344.

Ryugo, D. K., C-A Haenggeli, and J. R. Doucet. "Multimodal inputs to the granule cell domain of the cochlear nucleus." Exp Brain Res 153 (2003): 477-485.

Further Reading

Peruzzi, D., E. Bartlett, P. H. Smith, and D. L. Oliver. "A monosynaptic GABAergic input from the inferior colliculus to the medial geniculate body in rat." J Neurosci 16 (1997): 3766-3777.

Riquelme, R., E. Saldaña, K. K. Osen, O. P. Ottersen, and M. A. Merchán. "Colocalization of GABA and glycine in the ventral nucleus of the lateral lemniscus in rat: An in situ hybridization and semiquantitative immunocytochemical study." J Comp Neurol 432 (2001): 409-424.


Quantitative Methods



Delgutte, B. "Physiological models for basic auditory percepts." In Auditory Computation. Edited by H. H. Hawkins, T. A. McMullen, A. N. Popper, and R. R. Fay. New York, NY: Springer, 1995, pp. 157-220. ISBN: 9780387978437.

Parker, A. J., and W. T. Newsome. "Sense and the single neuron: Probing the physiology of perception." Annu Rev Neurosci 21 (1998): 227-277.

Buracas, G. T., and T. D. Albright. "Gauging sensory representations in the brain." Trends Neurosci 22 (1999): 303-9.

Papers for Discussion

Fitzpatrick, D. C., R. Batra, T. R. Stanford, and S. Kuwada. "A neuronal population code for sound localization." Nature 388 (1997): 871-4.

Shackleton, T. M., B. C. Skottun, R. H. Arnott, and A. R. Palmer. "Interaural time difference discrimination thresholds for single neurons in the inferior colliculus of guinea pigs." J Neurosci 23 (2003): 716-24.

Takahashi, T. T., A. D. Bala, M. W. Spitzer, D. R. Euston, M. L. Spezio, and C. H. Keller. "The synthesis and use of the owl's auditory space map." Biol Cybern 89 (2003): 378-87.

Note: You can skim the section entitled "Neural computation of auditory space," which lies beyond this theme's focus.

Hancock, K. E., and B. Delgutte. "A physiologically based model of interaural time difference discrimination." J Neurosci 24 (2004): 7110-7.

Chase, S. M., E. D. Young. "Limited segregation of different types of sound localization information among classes of units in the inferior colliculus." J Neurosci 25 (2005): 7575-85.

Furukawa, S., and J. C. Middlebrooks. "Cortical representation of auditory space: information-bearing features of spike patterns." J Neurophysiol 87 (2002): 1749-62.

Further Reading

Buy at MIT Press Rieke, F., D. Warland, de Ruyter Van Steveninck, and W. Bialek. Spikes. MIT Press. 1999. ISBN: 9780262681087. (A remarkably clear introduction to the application of information theory to neuroscience.)

Middlebrooks, J. C, L. Xu, S. Furukawa, and E. A. Macpherson. "Cortical neurons that localize sounds." Neuroscientist 8 (2002): 73-83. (Gives an overview of Middlebrook's work on cortical coding of sound location. Background for the assigned Furukawa and Middlebrooks paper.)

Borst, A., and F. E. Theunissen. "Information theory and neural coding." Nat Neurosci 2 (1999): 947-57. (A more detailed introduction to information theory than that given by Buracas and Albright.)

Johnson, D. H. "Dialogue concerning neural coding and information theory." August 2003. (This resource may not render correctly in a screen reader.PDF) (Unpublished document.) (Discusses the difficulties of applying Shannon information theory to studies of neural coding.)

Hsu, A., S. M. Woolley, T. E. Fremouw, and F. E. Theunissen. "Modulation power and phase spectrum of natural sounds enhance neural encoding performed by single auditory neurons." J Neurosci 24 (2004): 9201-11. (Application of the information theoretic approach outlined in "Spikes" to the auditory system of the song bird.)


Thalamus and Cortex



Kaas, J. H., and T. A. Hackett. "Subdivisions of auditory cortex and processing streams in primates." PNAS 97 (2000): 11793-11799.

Rauschecker, J. P., and B. Tian. "Mechanisms and streams for processing of "what" and "where" in auditory cortex." Proc Nat Acad Sci 97 (2000): 11800-11806.

Papers for Discussion

Wang, X., T. Lu, R. K. Snider, and L. Liang. "Sustained firing in auditory cortex evoked by preferred stimuli." Nature 435 (2005): 341-6.


Zatorre, R. J., and V. B. Penhune. "Spatial localization after excision of human auditory cortex." J Neurosci 21 (2001): 6321-6328.

Rose, H. J., and R. Metherate. "Auditory thalamocortical transmission is reliable and temporally precise." J Neurophysiol 94 (2005): 2019-30.

Further Reading

Malhotra, S., A. J. Hall, and S. G. Lomber. "Cortical control of sound localization in the cat: unilateral cooling deactivation of 19 cerebral areas." J Neurophysiol 92 (2004): 1625-43. (Complementary approach to the assigned Zatorre and Penhune paper.)



Student Topic Presentations







Ingvar, M., and K. M. Petersson. "Functional maps and brain networks." In Brain Mapping: The Systems. Edited by A. W. Toga, and J. C. Mazziotta. San Diego, CA: Academic Press, 2000, pp. 111-139. ISBN: 9780126925456.

Donchin, E., W. Ritter, and W. C. McCallum. "Cognitive psychophysiology: The endogenous components of the ERP." In Event-Related Potentials in Man. Edited by E. Callaway, P. Tueting, S. H. Koslow. New York, NY: Academic Press, 1979, pp. 349-411. ISBN: 9780121551506.

Papers for Discussion

Furst, M., R. A. Levine, and P. M. McGaffigan. "Click lateralization is related to the beta component of the dichotic brainstem auditory evoked potentials of human subjects." J Acoust Soc Am 78 (1985): 1644-1651.

Zimmer, U., and E. Macalusco. "High binaural coherence determines successful sound localization and increased activity in posterior auditory areas." Neuron 47 (2005): 893-905.

Lewald, J., H. Foltys, and R. Topper. "Role of the posterior parietal cortex in spatial hearing." J Neurosci 22 (2002): RC207.

Kraus, N., T. J. McGee, T. D. Carrell, S. G. Zecker, T. G. Nicol, and D. B. Koch. "Auditory neurophysiologic responses and discrimination deficits in children with learning problems." Science 273 (1996): 971-973.

Beauchamp, M. S., et al. "Unraveling multisensory integration: patchy organization within human STS multisensory cortex." Nat Neurosci 7 (2004): 1190-1192.

Scott, S. K., S. C. Blank, S. Rosen, and R. J. S. Wise. "Identification of a pathway for intelligible speech in the left temporal lobe." Brain 12 (2000): 2400-2406.

Additional Readings

Melcher, J. R. "Cellular generators of the binaural difference potential in cat." Hearing Res 95 (1996): 144-160.

Scott, S. K., and I. S. Johnsrude. "The neuroanatomical and functional organization of speech perception. TINS 26 (2003): 100-107.

Hall, D. A. "Auditory pathways: are 'what' and 'where' appropriate?" Curr Biol 13 (2003): 406-408.


Speech Motor Control

Guenther / Perkell


Barlow, S. M., G. R. Farley, and R. D. Andreatta. "Neural Systems in Speech Physiology." Chapter 2 in Handbook of Clinical Speech Physiology. San Diego, CA: Singular Publishing Group, 1999, pp. 101-165. ISBN: 9781565932678.

Guenther, F. H., S. S. Ghosh, and J. A. Tourville. "Neural modeling and imaging of the cortical interactions underlying syllable production." Brain and Language. 2005. (E-print ahead of publication.)

Papers for Discussion

Houde, J. F., and M. I. Jordan. "Sensorimotor adaptation in speech production." Science 279 (1998): 1213-1217.

Gomi, H., M. Monda, T. Ito, and E. Z. Murano. "Compensatory articulation during bilabial fricative production by regulating muscle stiffness." J Phonetics 30: 261-279.

Averbeck, B. B., M. V. Chafee, D. A. Crowe, and A. P. Georgopoulos. "Parallel processing of serial movements in prefrontal cortex." Proc Natl Acad Sci U.S.A. 99 (2002): 13172-7.

Further Reading

Houde, J. F., and M. I. Jordan. "Sensorimotor adaptation of speech I: Compensation and adaptation." J Speech Langu Hear Res 45 (2002): 295-310. (A more detailed version of the assigned Houde and Jordan (1998) paper.)

Amaral, D. G. "The functional organization of perception and movement." Chapter 19 in Principles of Neural Science. Edited by E. R. Kandel, J. H. Schwartz, and T. M. Jessell. New York, NY: McGraw-Hill, 2000, pp. 337-348. ISBN: 9780838577011.

Rizzolatti, G., G. Luppino, and M. Matelli. "The organization of the cortical motor system: New concepts." Electoencephalography and Clinical Neurophysiology 106 (1998): 283-296.




Refer to readings for Ses #15


Cortical Language Processing



Caplan, D., J. L. Gould. "Language and communication." In Fundamental Neuroscience. Edited by L. R. Squire, F. E. Bloom, S. K. McConnell, J. L. Roberts, N. C. Spitzer, and M. J. Zigmond. 2nd ed. Boston, MA: Academic, 2002, pp. 1333-1352. ISBN: 9780126603033.

Friederici, A. D. "Towards a neural basis of auditory sentence processing." Trends Cog Sci 6 (2002): 1364-1366.

Papers for Discussion

Dronkers, N. F., D. P. Wilkins, R. D. Van Valin, B. B. Redfern, and J. J. Jaeger. "Lesion analysis of the brain areas involved in sentence comprehension." Cognition 92 (2005): 145-177.

Caplan, D., E. Chen, and G. Waters. "Syntactic processing is accomplished only in Broca's area." 2005. (To be submitted to Science.)

Rizzolatti, G., and M. A. Arbib. "Language within our grasp." Trends Neurosci 21 (1998): 188-94.

Additional Reading

Rizzolatti, G., and L. Craighero. "The mirror-neuron system." Ann Rev Neurosci 27 (2004): 169-192. (Additional information about mirror neurons.)

Hickok, D., and D. Poeppel. "Towards a functional neuroanatomy of speech perception." Trends Cog Sci 4 (2000): 131-138. (Review of brain areas involved in speech perception with emphasis on low-level processing.)


Special Lecture

Guest Lecturer: Donata Oertel, Department of Physiology, University of Wisconsin, Madison



Student-initiated Topics



Lai, C. S., S. E. Fisher, J. A. Hurst, F. Vargha-Khadem, and A. P. Monaco. "A forkhead-domain gene is mutated in a severe speech and language disorder." Nature 413 (2001): 519-23.

MacDermot, K. D., E. Bonora, N. Sykes, A. M. Coupe, C. S. Lai, S. C. Vernes, F. Vargha-Khadem, F. McKenzie, R. L. Smith, A. P. Monaco, and S. E. Fisher. "Identification of FOXP2 truncation as a novel cause of developmental speech and language deficits." Am J Hum Genet 76 (2005): 1074-80.

Papers for Discussion

Liegeois, F., T. Baldeweg, A. Connelly, D. G. Gadian, M. Mishkin, and F. Vargha-Khadem. "Language fMRI abnormalities associated with FOXP2 gene mutation." Nat Neurosci 6 (2003): 1230-7.

Shu, W., J. Y. Cho, Y. Jiang, M. Zhang, D. Weisz, G. A. Elder, J. Schmeidler, R. De Gasperi, M. A. Sosa, D. Rabidou, A. C. Santucci, D. Perl, E. Morrisey, and J. D. Buxbaum. "Altered ultrasonic vocalization in mice with a disruption in the Foxp2 gene." Proc Natl Acad Sci U.S.A. 102 (2005): 9643-8.

Teramitsu, I., L. C. Kudo, S. E. London, D. H. Geschwind, and S. A. White. "Parallel FoxP1 and FoxP2 expression in songbird and human brain predicts functional interaction." J Neurosci 24 (2004): 3152-63.


Student-initiated Topics



Sumby, W. H., and I. Pollack. "Visual contribution to speech intelligibility in noise." J Acoust Soc Am 26 (1954): 212-215.

McGurk, H., and J. MacDonald. "Hearing lips and seeing voices." Nature 264 (1976): 746-748.

Calvert, G. A., E. T. Bullmore, M. J. Brammer, R. Campbell, S. C. R. Williams, P. K. McGuire, P. W. R. Woodruff, S. D. Iversen, and A. S. David. "Activation of auditory cortex during silent lipreading." Science 276 (1997): 593-596.

Papers for Discussion

Bushara, K. O., T. Hanakawa, I. Immisch, K. Toma, K. Kansaku, and M. Hallet. "Neural correlates of cross-modal binding." Nature Neurosci 6 (2003): 190-195.

Calvert, G. A., and R. Campbell. "Reading speech from still and moving faces: The neural substrates of visible speech." J Cogn Neurosci 15 (2003): 57-70.

Komura, Y., R. Tamura, T. Uwano, H. Nishijo, and T. Ono. "Auditory thalamus integrates visual inputs into behavioral gains." Nature Neurosci 8 (2005): 1203-1209.


Student-initiated Topics



Pasternak, T., and M. W. Greenlee. "Working memory in primate sensory systems." Nature Reviews Neurosci 6 (2005): 97-107.

Papers for Discussion

Gottlieb, Y., E. Vaadia, and M. Abeles. "Single unit activity in the auditory cortex of a monkey performing a short-term memory task." Exp Brain Res 74 (1989): 139-148.

Colombo, M., M. R. D'Amato, H. R. Rodman, and C. G. Gross. "Auditory association cortex lesions impair auditory short-term memory in monkeys." Science 247 (1990): 336-338.

Arnott, S. R., C. L. Grady, S. J. Hevenor, S. Graham, and C. Alain. "The functional organization of auditory working memory as revealed by fMRI." J Cog Neurosci 17 (2005): 819-831.

Further Reading

Sakurai, Y. "Cells in the rat auditory cortex have sensory-delay correlates during the performance of an auditory working memory task." Behav Neurosci 104 (1990): 856-868. (This study compares single-unit activity in different auditory brain areas while rats perform a short-term auditory memory task.)

Colombo, M., H. R. Rodman, and C. G. Gross. "The effects of superior temporal cortex lesions on the processing and retention of auditory information in monkeys (cebus apella)." J Neurosci 16 (1996): 4501-4517. (A detailed follow up to the Colombo et al. (1990) study featuring a comparison between the effects of unilateral and bilateral lesions and considerable discussion.)

Gaab, N., C. Gaser, T. Zaehle, L. Jancke, and G. Schlaug. "Functional anatomy of pitch memory-an fMRI study with sparse temporal sampling. " NeuroImage 19 (2003): 1417-1426. (Another fMRI study of short-term auditory memory with several interesting methodological features.)