BISC 521/NEUR 542 Spring Semester 2010
Instructors:
Caroline Abdala
Lisa Aziz-Zadeh
Lynne Bernstein
Sarah Bottjer*
Andres Collazo
Laurie Eisenberg
Rick Friedman
Qian-Jie Fu
Louis Goldstein
Federico Kalinec
David Lim
Toby Mintz
Shri Narayanan
Neil Segil
Bob Shannon*
Rachel Walker
Li Zhang
HEI
HNB B20
HEI
HNB 218
HEI
HEI
HEI
HEI
GFS 301G
HEI
HEI
SGM 613
EEB 430
HEI
HEI
MMR 626
213-273-8003
213-821-2970
213-353-7044
213-740-9183
213-353-7075
213-273-8021
213-273-8078
213-273-8036
213-821-4150
213-353-7030
213-353-7021
213-740-2253
213-740-6432
213-273-8082
213-353-7020
323-442-1635 cabdala@hei.org
lazizzad@usc.edu
lbernstein@hei.org
bottjer@usc.edu
acollazo@hei.org
leisenberg@hei.org
rfriedman@hei.org
qfu@hei.org
louisgold@usc.edu
fkalinec@hei.org
dlim@hei.org
tmintz@usc.edu
shri@sipi.usc.edu
nsegil@hei.org
bshannon@hei.org
liizhang@usc.edu
Location: HNB 120 Time: Tuesdays and Thursdays 10-11:50am Units: 4
Course Description:
Grading (letter) : Evaluation will be based on class participation (10%), a midterm (40%) and a final examination (50%). A large pool of possible exam questions will be made available the week before each exam. Approximately 80% of the exam will be composed of these questions.
Course Material : Course materials (syllabus, readings, lecture figures, etc.) will be available on
Blackboard: https://blackboard.usc.edu
. Be sure to check this site frequently. Many important announcements will appear first on Blackboard. Pre-requisite: BISC 421 or NEUR 524.
Students with Disabilities: Students requesting academic accommodations based on a disability are required to register with Disability Services and Programs (DSP) each semester.
A letter of verification for approved accommodations can be obtained from DSP when adequate documentation is filed. Please be sure the letter is delivered to one of the instructors as early in the semester as possible. Disability Services and Programs is located in Student Union 301 and their phone number is (213) 740-0776.
Statement on Academic Integrity: USC seeks to maintain an optimal learning environment.
General principles of academic honesty include the concept of respect for the intellectual property of others, the expectation that individual work will be submitted unless otherwise

allowed by an instructor, and the obligations both to protect one’s own academic work from misuse by others as well as to avoid using another’s work as one’s own. All students are expected to understand and abide by these principles. Scampus, the Student Guidebook, contains the Student Conduct Code in Section 11.00, while the recommended sanctions are located in Appendix A: http://www.usc.edu/dept/publications/SCAMPUS/gov/. Students will be referred to the Office of Student Judicial Affairs and Community Standards for further review, should there be any suspicion of academic dishonesty. The Review process can be found at: http://www.usc.edu/student-affairs/SJACS/ .
SCHEDULE OF LECTURES
Tuesday, Jan 12: From the Ear to Hearing: An Overview of the Auditory System (Shannon,
Bottjer).
A general introduction to hearing and communication science.
Overview of the course (dates of exams, etc)
Thursday, Jan 14: How the Ear Develops: (Collazo, Segil, Lim)
Morphogenesis and cellular development of the peripheral auditory system;
Early embryology and neurogenesis
Barald, K. F. and Kelley, M. W. (2004). From placode to polarization: new tunes in inner ear development.
Development 131, 4119-30.
Fritzsch, B. and Beisel, K. W. (2001). Evolution and development of the vertebrate ear. Brain Res Bull 55,
711-21.
Koundakjian EJ, Appler JL, Goodrich LV. Auditory neurons make stereotyped wiring decisions before maturation of their targets. J Neurosci. 2007 Dec 19;27(51):14078-88.
Raft S, Koundakjian EJ, Quinones H, Jayasena CS, Goodrich LV, Johnson JE, Segil N, Groves AK.Crossregulation of Ngn1 and Math1 coordinates the production of neurons and sensory hair cells during inner ear development. Development.
2007 Dec;134(24):4405-15.
Tuesday, Jan 19: How the Ear Develops: (Collazo, Segil, Lim)
Cochlear morphogenesis and the origin of sensory, non-sensory, and neuronal structures; Cellular differentiation, maturation and regeneration of the organ of Corti.
Goodyear, R., Kros, C. and Richardson, G. (2006). The Development of Hair Cells in the Inner Ear. In
Vertebrate Hair Cells , pp. 20-94.
Kelley MW.; Regulation of cell fate in the sensory epithelia of the inner ear. Nat Rev Neurosci. 2006
Nov;7(11):837-49. Review.
Eddison M, Le Roux I, Lewis J. Notch signaling in the development of the inner ear: lessons from
Drosophila.
Proc Natl Acad Sci U S A. 2000 Oct 24;97(22):11692-9.
White PM, Doetzlhofer A, Lee YS, Groves AK, Segil N. Mammalian cochlear supporting cells can divide and trans-differentiate into hair cells. Nature. 2006 Jun 22;441(7096):984-7.
Stone JS, Cotanche DA Hair cell regeneration in the avian auditory epithelium. Int J Dev Biol. 2007;51(6-
7):633-47. Review.
Stone JS, Oesterle EC, Rubel EW. Recent insights into regeneration of auditory and vestibular hair cells.
Curr Opin Neurol. 1998 Feb;11(1):17-24. Review.
Thursday, Jan 21: How the Ear Develops: (Collazo, Segil, Lim)
Inner ear homeostasis: The stria vascularis and inner ear fluid and ion regulation related to hearing and deafness; Causes of deafness and their current and future treatment.
Chen J, Nathans J . “Estrogen-related receptor beta/NR3B2 controls epithelial cell fate and endolymph production by the stria vascularis.”
Dev Cell.
2007 Sep;13(3):325-37.
Nickel R, Forge A. Gap junctions and connexins in the inner ear: their roles in homeostasis and deafness.
Curr Opin Otolaryngol Head Neck Surg . 2008 Oct;16(5):452-7. Review.
Tuesday, Jan 26: How We Hear: Turning on the System —The Functional Development of the
Peripheral Auditory System (Abdala, Kalinec, Collazo)
The physics of sound and the evolution of the ear.

L Robles & M Ruggero (2001) “Mechanics of the Mammalian Cochlea”. Physiol. Rev.
81(3):1305-1352
J Ashmore (2008) “Cochlear Outer Hair Cell Motility”. Physiol Rev 88: 173-210.
GA Manley (2000) “Cochlear Mechanisms from a Phylogenetic Viewpoint”. PNAS (USA) 97(22):11736-
11743
AJ Hudspeth (2008) “Making an Effort to Listen: Mechanical Amplification in the Ear”.
Neuron 59:530-545.
GI Frolenkov et al.
(2004) “Genetic Insights into the Morphogenesis of Inner Ear Hair Cells”.
Nature Rev
Genetics 5:489-498
Thursday, Jan 28: How We Hear: Turning on the System —The Functional Development of the
Peripheral Auditory System (Abdala, Kalinec, Collazo)
Biomechanics and electrophysiology of hearing and the cochlea.
The cochlear amplifier —active mechanisms and otoacoustic emissions.
Hudspeth, A. J. (2008). Making an effort to listen: mechanical amplification in the ear. Neuron 59, 530-45.
Manley, G. A. (2001). Evidence for an active process and a cochlear amplifier in nonmammals. J
Neurophysiol 86, 541-9.
Riley, B. B. and Phillips, B. T. (2003). Ringing in the new ear: resolution of cell interactions in otic development. Dev Biol 261, 289-312.
Rubel, E. W. and Fritzsch, B. (2002). Auditory system development: Primary auditory neurons and their targets. Annu Rev Neurosci 25, 51-101.
James O. Pickles An Introduction to the Physiology of Hearing , Second Edition (1988) Academic Press,
Chapter 3: The Cochlea and Chapter 5: Mechanisms of Transduction and Excitation in the
Cochlea.
Tuesday, Feb 2: How We Hear: Turning on the System —The Functional Development of the
Peripheral Auditory System (Abdala, Kalinec, Collazo)
Getting the signal to the brain: cochlear innervation. Type 1 and Type 2 afferent neurons, lateral and crossed efferent neurons. Synaptic patterns on inner and outer hair cells.
Dallos, Popper and Fay (Eds), 1996, The Cochlea , Springer-Verlag, Chapter 8; Physiology of Olivocochlear
Efferents
Thursday, Feb 4: How We Hear: The Auditory Pathways of the Brain (Bottjer, Zhang)
Auditory Pathways I: Functional circuits for brainstem and midbrain
—processing and integration; auditory localization - physiology of the auditory nerve (rate-intensity functions, phase locking, high and low spontaneous rate fibers), transduction
Neuroscience , 4th Edition. D. Purves et al. (eds.), Chapter 13, "The Auditory System" (pp. 313-342).
Sinauer Associates, Sunderland MA, 2008.
Tuesday, Feb 9: How We Hear: The Auditory Pathways of the Brain (Bottjer, Zhang)
Cochlear nucleus (SCC, choppers, octopus cells, bushy cells, AVCN, PVCN, DCN), olive complex (medial and lateral, laminaris in birds), lateral lemniscus, inferior colliculus.
Neuroscience , 4th Edition. D. Purves et al. (Eds.). Chapter 13, "The Auditory System" (pp. 313-342),
Sinauer Associates, Sunderland MA, 2008.
Thursday, Feb 11: How We Hear: The Auditory Pathways of the Brain (Bottjer, Zhang)
Auditory Pathways II: Functional circuits for auditory thalamus and cortex; representing complex sounds; role of experience in cortical development.
Auditory Pathways III: Cortex.
Chapter 13, "The Auditory System" (pp. 313-342) Neuroscience , 4th Edition. D. Purves et al., editors.
Sinauer Associates, Sunderland MA, 2008.
Tuesday, Feb 16: Genetics of Hearing and Communications: The Awesome Power of Modern
Genetics (Friedman).
Identifying genes: From Mendelian genetics to whole genome studies;
The genetics of susceptibility: Natural variation, and why some of us lose our hearing;
The genetics of language.
Ballenger's Otorhinolaryngology Head and Neck Surgery, 17th edition , James B. Snow Jr. and P. Ashley
Wackym (Eds.), Chapter 26 - Hereditary Hearing Loss (Dr. Friedman is an author on this chapter)

Leibovici M, Safieddine S, Petit C. Mouse models for human hereditary deafness. Curr Top Dev Biol.
2008;84:385-429. Review.
Steel KP, Kros CJ. A genetic approach to understanding auditory function.
Nat Genet.
2001 Feb;27(2):143-
9. Review.
Vernes SC, Newbury DF, Abrahams BS, Winchester L, Nicod J, Groszer M, Alarcón M, Oliver PL, Davies
KE, Geschwind DH, Monaco AP, Fisher SE. A functional genetic link between distinct developmental language disorders.
N Engl J Med.
2008 Nov 27;359(22):2337-45.
White SA, Fisher SE, Geschwind DH, Scharff C, Holy TE. Singing mice, songbirds, and more: models for
FOXP2 function and dysfunction in human speech and language. J Neurosci. 2006 Oct
11;26(41):10376-9. Review.
Friedman et al. GRM7 variants confer susceptibility to age-related hearing impairment. Hum Mol Genet;
18(4), 785
Genome-wide association studies for common diseases and complex traits. Hirschorn and Daly, Nature
Genetics Reviews
Thursday, Feb 18: How We Hear: Normal Perception and its Disorders (Shannon, Eisenberg,
Fu, Abdala)
Auditory Psychophysics —loudness, pitch, temporal processing, frequency selectivity, binaural hearing.
Moore BCJ. Hearing (Handbook of Perception and Cognition, Second Edition) Academic Press.
Tuesday, Feb 23: How We Hear: Normal Perception and its Disorders (Shannon, Eisenberg,
Fu, Abdala)
Normal and Abnormal Perceptual Development
—development of frequency tuning, developmental psychophysics.
Dallos, P. (1992). The Active Cochlea. J. of Neuroscience, 12, 4575
Thursday, Feb 25: How We Hear: Normal Perception and its Disorders (Shannon, Eisenberg,
Fu, Abdala)
Audiology and Hearing Impairment —Clinical assessment of hearing, electrophysiology, otoacoustic emissions, conductive vs. cochlear hearing loss, flat vs. sloping audiograms, loss of sharpness in auditory filters, auditory neuropathy.
Dirks, D.D., Ahlstrom, J.B., & Morgan, D.E. (2000). Auditory sensitivity: Air and bone conduction. In Canalis,
R.F., & Lambert, P.R. (Eds.), The Ear: Comprehensive Otology , Chapter 10. Philadelphia:
Lippincott Williams & Wilkins.
Dubno, J.R., & Dirks, D.D. (2000). Measures of auditory function using speech stimuli. In Canalis, R.F., &
Lambert, P.R. (Eds.), The Ear: Comprehensive Otology , Chapter 11. Philadelphia: Lippincott
Williams & Wilkins.
Pediatric Otology and Neurotology (1998) Eds Anil Lalwani and Kenneth Grundfast. Lippencott – Raven.
Chapter 9. Physiologic Assessment of Hearing.
Tuesday, Mar 2: How We Hear: Normal Perception and its Disorders (Shannon, Eisenberg, Fu,
Abdala)
Cochlear Implants - electric activation of neurons, safety concerns of electric stimulation of neurons, cochlear implant history and development.
Loizou, P.C. (1998). Mimicking the human ear, IEEE Signal Processing Magazine , 15(5) 101-130.
Shannon, R.V. (1993). Psychophysics of electrical stimulation, in Cochlear Implants: Audiological
Foundations , R.S. Tyler (Ed.), Singular Pub. Grp., San Diego, pp. 357-388.
Shannon, R.V. (1992). A model of safe levels for electrical stimulation, IEEE Transactions on Biomedical
Engineering , 39(4), 424-426.
Thursday, Mar 4: How We Hear: Normal Perception and its Disorders (Shannon, Eisenberg, Fu,
Abdala)
Cochlear Implant psychophysics and speech perception, speech processing for implants, central pattern processing in implants
Shannon, R.V., Zeng, F-G., Kamath, V., Wygonski, J., and Ekelid, M (1995). Speech recognition with primarily temporal cues. Science, 270, 303-304.

Fu, Q.-J. and Shannon, R.V.
(1999).
Recognition of spectrally degraded and frequency-shifted vowels in acoustic and electric hearing, Journal of the Acoustical Society of America , 105(3), 1889-1900.
Tuesday, Mar 9: How We Hear: Normal Perception and its Disorders (Shannon, Eisenberg, Fu,
Abdala)
Auditory Brainstem Implants and Auditory midbrain implants, implant psychophysics
(loudness, pitch, temporal processing, spatial selectivity, etc).
Shannon, R.V. and Otto, S.R. (1990). Psychophysical measures from electrical stimulation of the human cochlear nucleus, Hearing Res.
, 47, 159-168.
Colletti V and Shannon RV (2005). Open Set Speech Perception with Auditory Brainstem Implant? The
Laryngoscope 115:1974-1978.
Otto SR, Shannon RV, Brackmann DE, Hitselberger WE, McCreery D, Moore J, Wilkinson E (2008).
Audiological Outcomes with the Penetrating Electrode Auditory Brainstem Implant, Otology and
Neurotology , 29:1147-1154.
Thursday, Mar 11: Midterm Exam
Tuesday, Mar 16: Spring Break; Thursday, Mar 18: Spring Break
Tuesday, Mar 23: How We Use Hearing: Speech and Language (Eisenberg, Walker, Byrd,
Goldstein, Mintz)
Speech Production -- Articulation and linguistic contrast, vocal tract anatomy, source filter theory, parameters of linguistic contrast, co-articulation and parallel transmission,
Instrumental techniques for articulation research.
Goldstein, L. & Rubin, P. (2007). Speech - Dances of the Vocal Tract. Odyssey Magazine , v.16:no.1, pp. 14-
15. http://www.haskins.yale.edu/Reprints/HL1452.pdf
"Phonetics" chapter from Discovering Speech, Words, and Mind (Byrd & Mintz); to be provided
Thursday, Mar 25: How We Use Hearing: Speech and Language (Eisenberg, Walker, Byrd,
Goldstein, Mintz)
Speech Perception and Sound Patterns in Language -- Typology of sound patterns in language, perception of consonants and vowels, syllable structures and speech perception, (mis-)perception of speech and theory of language typology.
Bergman, Anouschka, Kathleen Currie Hall, & Sharon Miriam Ross (eds.). 2007. Implicational laws. In
Language Files 10 , pp. 122-126. Columbus: The Ohio State University Press.
Wright, Richard. 2004 A review of perceptual cues and cue robustness. In Bruce Hayes, Robert Kirchner, and Donca Steriade, eds., Phonetically Based Phonology . Cambridge: Cambridge University
Press.
Tuesday, Mar 30: How We Use Hearing: Speech and Language (Eisenberg, Walker, Byrd,
Goldstein, Mintz)
Development of Speech Perception -- Infants' innate capacities for categorizing speech sounds, the tuning of perceptual categories in the first year of life and proposed mechanisms, the fragility of perceptual contrasts in early word learning, Infants' segmentation of continuous speech in to words.
Saffran, J. R., Werker, J. F., & Werner, L. A. (2006). The infant's auditory world: Hearing, speech, and the beginnings of language. In D. Kuhn and R. Siegler (Eds.), Handbook of Child Psychology Sixth
Edition, Vol. 2: Cognition, Perception, and Language . New Jersey: Wiley. pp. 72-91.
Thursday, Apr 1: How We Use Hearing: Speech and Language (Eisenberg, Walker, Byrd,
Goldstein, Mintz)
Development of Speech Production -- Emergence of phonological structure, selforganization models, sensorimotor abilities of young infants, onset and structure of babbling, attunement of vocal actions to the ambient language, phonetic and phonological structure of early words.
Kuhl, P. and Meltzoff (1996). Infant vocalizations in response to speech: Vocal imitation and developmental change. Journal of the Acoustical Society of America. 100, 2425-2438.

Davis, B. and MacNeilage (1995). The articulatory basis of babbling. Journal of Speech and Hearing
Research, 38, 1199-1211.
Studdert-Kennedy, M. and Goldstein, L. (2003) Launching language: Gestural origin of discrete infinity.
Christiansen, M. & Kirby, S. (eds.) Language evolution: The States of the Art. (pp. 235-254).
Oxford: Oxford University Press.
Tuesday, Apr 6: How We Use Hearing: Speech and Language (Eisenberg, Walker, Byrd,
Goldstein, Mintz)
Development of Speech Perception and Production in Children with Hearing Loss --
Phonetic contrast and phoneme perception, word and sentence recognition, primitive and complex vocalizations, phonemic-syllabic vocal representations.
Johnson, K..C., Eisenberg, L.S., & Martinez, A.S. (in preparation). Moderate to profound sensorineural hearing loss in children” In Gravel, J., Seewald, R, & Tharpe, A.M., Comprehensive Handbook of
Pediatric Audiology.
San Diego: Plural Publishing.
Eisenberg, L.S. (2007). Current state of knowledge: Speech recognition and production in children with hearing impairment. Ear and Hearing , 28, 766-772.
Tuesday, Apr 13: How We Use Hearing: Auditory Cognitive Neuroscience (Bernstein, Aziz-
Zadeh, Lu, Narayanan, Fu)
The evolution of auditory perception.
Kohler E, Keysers C, Umiltà MA, Fogassi L, Gallese V, Rizzolatti G. Hearing sounds, understanding actions: action representation in mirror neurons. Science . 2002 Aug 2;297(5582):846-8.
Gazzola V, Aziz-Zadeh L, Keysers Empathy and the somatotopic auditory mirror system in humans C.Curr
Biol.
2006 Sep 19;16(18):1824-9.
Thursday, Apr 15: How We Use Hearing: Auditory Cognitive Neuroscience (Bernstein, Aziz-
Zadeh, Lu, Narayanan, Fu)
Signal processing of speech —time representation of speech (voiced and unvoiced, F0), spectral representation of speech (formants, glides).
Rabiner and Schafer: Digital Processing of Speech Signals , Prentice Hall, 1978.
Deller, Hansen, and Proakis: Discrete-time Processing of Speech Signals , 2000.
T. Quatieri: Discrete-time Speech Signal Processing , Prentice Hall, 2001.
Tuesday, Apr 20: How We Use Hearing: Auditory Cognitive Neuroscience (Bernstein, Aziz-
Zadeh, Lu, Narayanan, Fu)
Extraction of features from speech —automatic speech recognition, noise reduction processing for speech.
Fundamentals of Speech Recognition , Rabiner and Juang, Prentice Hall, 1993
Spoken Language Processing: A guide to theory, algorithm and system development , X. Huang, A. Acero,
H-W. Hon, Prentice Hall 2001
Thursday, Apr 22: How We Use Hearing: Auditory Cognitive Neuroscience (Bernstein, Aziz-
Zadeh, Lu, Narayanan, Fu)
Speech recognition and language processing.
Hickok, G. and D. Poeppel .
The cortical organization of speech processing. Nature Reviews: Neuroscience ,
2007. 8 (5): p. 393-402.
Scott, S.K. and I.S. Johnsrude .
The neuroanatomical and functional organization of speech perception.
Trends in Neurosciences , 2003. 26 (2): p. 100-107.
Tuesday, Apr 27: How We Use Hearing: Auditory Cognitive Neuroscience (Bernstein, Aziz-
Zadeh, Lu, Narayanan, Fu)
Multi-sensory integration —eyes, ears and skin.
Ghazanfar, A.A. and C.E. Schroeder .
Is neocortex essentially multisensory? Trends Cogn Sci , 2006. 10 (6): p. 278-85.
Meredith, M.A. On the neuronal basis for multisensory convergence: a brief overview. Cognitive Brain
Research , 2002. 14 : p. 31-40.
Thursday, Apr 29: Final overview and Questions