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SYLLABUS for NEUROBIOLOGY 101
updated Sept. 17, 2007
Harvard College & Graduate School of Arts & Sciences
Fall Term, 2007-2008
Monday Sept 17, 2007 - January 23, 2008
GSAS Registration, Sept 12, 2007
MS Upper Class Registration, Sept 14, 2007
Neurobiology 101. Auditory Neurobiology of Language & Music
Lectures Tues & Thurs, 11:30A -1:00P
Harvard Yard, Sever Hall, Room 202
I Section per Wk (90 mins), Time & Place to be announced in class
Fatally
Mark Jude Tramo, MD, PhD
Asst. Professor of Neurology, Harvard Medical School
Steering Committee, Harvard University Mind/Brain/Behavior Interfaculty Initiative
Attending Neurologist, Massachusetts General Hospital
www.Brain'Music.org
Office hours by appoinnnenr, The Institute for Music & Brain Science, 175 Cambridge St, Suite 340, Room 382, Boston
Jonathan I. Matsui, PhD
Lecturer on Molecular & Cellular Biology, Harvard University
Neurobiology Concentration Advisor for Harvard College Undergraduates, Harvard Faculty of Arts & Sciences
Office hours by appointment, BioLabs Rm 1082a, Divinity Ave, Cambridge
Teaching Fellow
Caroline Niziolek, MS
PhD Candidate, Harvard-MIT Speech & Heating Bioscience & Technology Graduate Studies Program
Office hours by a
ointment, BioLabs Rm 1075, Divinity Ave, Cambridge
Guest Lecturers
Marc D. Hauser, PhD
Co-Director, Harvard University Mind/Brain/Behavior Interfaculty Initiative
Harvard College Professor of Psychology and of Organismic & Evolutionary Biology
Louis D. Braida, PhD
Co-Director, Harvard-M.I.T. Speech & Hearing Biosciences & Technology Graduate Studies Program
Henry Warren Ellis Professor of Electrical Engineering, M.I.T.
Sensory Communication Group, Research Laboratory of Electronics, M.I.T.
Bertrand Delgutte, PhD
Co-Director, Harvard-M.I.T. Speech & Bearing Biosciences & Technology Graduate Studies Program
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Senior Research Scientist, Research Laboratory of Electronics, M.I.T.
Eaton-Peabody laboratory• of Auditory Physiology, Dept of Otology & laryngology, Harvard Medical School
Course Description
Flow does the brain transform air-pressure waves striking our eardrums — or the words you
are reading on this page — into the sounds we hear in our heads? What parts of the hearing
brain are necessary and sufficient to understand spoken, seen, or imagined words, dance to
music, and recognize your mother's voice? When do auditory-perceptual capacities critical
to language acquisition develop?
What happens to linguistic and musical abilities if
congenital or acquired diseases alter their underlying molecular substrates and neural
circuitry?
In this 13-week lecture course, current knowledge and conceptual insights from multiple
scientific disciplines and the humanities (biology, engineering, computer science, psychology,
linguistics, musk) are brought to bear on these questions. Lectures and Sections are
organized into five topics: 1) Introduction to Auditory Neurobiology, 2) Auditory Communication•,
3) Signal Processing b. the Auditory Systenr, 4) Molecular Biology of the Auditory System, and 5)
Developmental Biology of the Auditoy System. We introduce data from experiments investigating
how the brain and ear 1) transform acoustic signals into auditory percepts; 2) discriminate
differences within and across acoustic and semantic categories of auditory percepts (e.g.,
different words, voices, musical instruments); 3) abstract perceptual constancies despite
variations in acoustic features (e.g., your roommate's voice here, there, and everywhere last
year); and 4) integrate percept formation with implicit knowledge about language and music
gained through years of acculturation. Students will be expected to master topics in systems
neurobiology, molecular neurobiology, psychophysics, and other fields of active research in
auditory neuroscience. Basic principles taught in multiple undergraduate departments will be
connected with concepts about the neurobiology of disease as they relate to deafness,
aphasia, dyslexia, and amusia; these connections will provide a basis for introducing
contextualized material in molecular pathology, systems pathophysiology, neurology, and
neuroimaging that are traditionally reserved for medical school and graduate school
education. Emerging treatment, rehabilitation, and education technologies and new ideas for
future research in the brave new worlds of neurobionics and genetic engineering will be
explored.
Pre-requisites
MCB 80. Neurobiology of Behavior
Secondary• School Physics or Engineering
Secondary• School Algebra and Trigonometry
Not required: Ability to read or play music
Course Requirements ter Grading
Lecture & Section Participation & Homework (25% of grade)
One Mid-Term Examination (30% of grade)
Final Examination (45% of grade)
No make-up MiatTenze Examination aril/ be crud.
If you miss the Mid•Term Examination; 1) you must have a written medical excuse from University Health Services in order to pass the
count; and Z your course grade will he calculated as follows: lecture & Section = 25%; Final Examination = 75%.
Books ter Other Resources - Please see end of document*
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WEEK 1
Lecture Series I. Introduction to Auditory Neurobiology
Tuesday, Sept 18
Course Overview
The Heating Brain
Tramo
No Section in Week 1
Thursday, Sept 20
Pictures of Sound
Tramo
Friday, Sept 21, STUDY CARD DAY
5P today is the deadline for undergraduate and graduate students to complete electronic sectioning in our
course; Section time(s) and plate(es) will have been discussed in class.
WEEK 2
Lecture Series IL Auditory Communication
Tuesday, Sept 25
f?1
,11144. *Zyletfle In/Mil
Phonetics, Phonology,
Prosody in Relation to Grammar dr Semiotics in Babble, Intetjections, Expletives,
Conversation, Prose, Patty, Son& Music, Film, Puns, dr Branding (Whew!)
Tramo
Section #1
Acoustical Meant:winks: From Sound Level Meters to Oscilloscopes to Garage Band
Thursday, Sept 27
Neuroplysiolog do'Neuroanatomry of Language I
Tramo
WEEK 3
Monday, Oct 1
5P today is the deadline for undergraduates to either drop the course or change the grade from a letter-grade to
pass fail (or vice versa) without frying the Registrar a fie
Tuesday, Oct 2
Neuropsysioky
Neuroanatomry of Language II
Tramo
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Section #2
Acoustic Phonetics & Pgchoacoustics Laboratory, Demotutrations
Thursday, Oct 4
Nemwtsiolog &Neuron/Wong of Language III
Tramo
WEEK 4
Tuesday, Oct 9
Evokition of the Language Faculty
Hauser
Section #3
Human Brain Imaging
Thursday, Oct 11
Neuropkysioksy
Neuroanatewg of Music I
Tramo
WEEK 5
Monday, Oct 15
5/) today is the deadline for undergraduates to either drop the course or change the grade from a letter-grade to
passfail (or vice versa)
Tuesday, Oct 16
Neuroplysiokgy &Nesiroanatong of Music II
Tramo
Section #4
Electra- & Magneto-encephakgraply (EEG &MEG): Evoked Potentials & Event Related Potentials
(EPs & ERPs)
Thursday, Oct 18
Neuroplysiology &Me/wow:atom,. of Musk III
Tramo
WEEK 6
Tuesday, Oct 23
Comparisons Between Language & Music
Tramo
Section #5
Review of Auditory Communication
Lecture Series III: Signal Processing ky the Auditory System
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Thursday, Oct 25
The Ear and Organ of Corti: Signal Amplification, Filtering, Transduction, and Mechanical Tuning in the
Auditory Periphery
Matsui
WEEK 7
Monday, Oct 29
51) today h the deadline for graduate students to registerfortbis course
Tuesday, Oct 30
The Auditory Nerve: Representation of Vocal dr Masked Stimulus Features in the Discharge Patterns of
Auditory-Nara Fibers
Delgutte
Section #6
Phyriology & Anatomy of the Auditory System
Thursday, Nov 1
Miracle Ear. Curing Deafness via Acoustic-to-Bioeleriric Signal Transformation & Direct Electrical
Stimulation of the Cochlea Using Implantable, Programmable Devices
Braida
WEEK 8
Tuesday, Nov 6
Auditory Brainstem
Tramo
Section #7
Spike Trains & Neural Coding
Thursday, Nov 8
Auditory Brainstent &Auditory Cortex
Tramo
WEEK 9
Tuesday, Nov 13
Auditory Cortex, Multi-Modal Cortex, & Supra-Modal Cortex
Tramo
Section #8
Review of Signal Processing bi tbe Auditory System
Thursday, Nov 15
Mid-Tern Examination: Auditory Communication & Signal Processing by the Aucktog System
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Saturday, Nov 17
Yale Game, at Yale
WEEK 10
Tuesday, Nov 20
Review of Mid-Term Examination
Tramo
Thursday, Nov 22
Thanksgiving
No Chars, No Section
WEEK 11
Lecture Series IV: Developmental Biology of the Auditory System
Tuesday, Nov 27
Development oft& Ear & Penpberal Auditory Nervous System
Matsui
Section #9
'The (Neuro)Bionic Woman": Prime Time for Bioengineering the Future
Thursday, Nov 29
Development of the Central Auditory Newous System
Matsui
WEEK 12
Tuesday, Dec 4
Language Development in Infants & Children
Matsui
Section #10
ita" Pt. te, Critical Periods, & Sign Language
Lecture Series V. Molecular Biology of the Auditory System
Thursday, Dec 6
Molecular Pathology & Pathopbesiology of Genetic Diseases I
Matsui
WEEK 13
Tuesday, Dec 11
Molecular Pathology
Pathornsiology of Genetic Diseases II
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Matsui
Section #11
Congenital Hearing Loss a' the Search for Mutations
Thursday, Dec 13
Social Prychology of Developmental Communication Disorders
Matsui
WEEK 14
Tuesday, Dec 18
Review of Mokmkr ier Developmental Biology of the Auditory System
Matsui
No Section in Week 14
Wednesday, Dec 19
First Day of Winter Recess
READING PERIOD
Section #12
Date will have been discussed and announced in class
FINAL EXAM
Date will have been annomwed in class
Exam Group 13: Mon., Jan 14, 2008
Exam Group 14: Thu., Jars 17, 2008
In selecting courses, students should understand that final exams for two courses that fall within the same
Exam Group or Exam Group Set (see below) will be scheduled on the same day. Students who want to avoid
having two exams on one day should not enroll in two courses with the same Exam Group or Exam Group
Set. All courses assigned to the following Exam Group Sets are scheduled to have their final exams on the
same day:
Exam Groups I, 7, 14
Exam Groups 2, 15, 16
Exam Groups 6, 10
Exam Groups 8, 9, II, 12, 17, 18
Occasionally, the Registrar may need to assign an Exam Group that does not correspond to the meeting days
and times of a course - students are therefore advised that they should not make any travel plans until the
official final examination schedule is published.
*REQUIRED BOOK efr FREEWARE
1. Fundamentals of Hearing, 5" Ec4 by William Yost. Academic Press, NY, 2007.
Available at the Harvard Coop. Excellent figures, appendices, and supplements re: topics we cover in
acoustics, psychophysics, audiology, and signal processing in the peripheral auditory nervous system.
Lecture notes and recommended reading will provide additional information on these topics and
most, if not all, of the written material about signal processing in the central auditory system,
molecular and cellular neurobiology, systems neurobiology, neurobiology of disease, and
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neuroimaging. At Coop; On Resent
2. http://www.praat.org.
Paul Boersma and David Weenink's website has Praat freeware ("praat" is Dutch for "gossip" or
"speak") — a digital sound recording, editing, and analysis program. We use Praat in our course
demonstrations, and it is in widespread use in speech and hearing research laboratories around the
world. Kudos to Pal and David! Mac, IBM, and other platforms are supported. One of the best
ways to learn about sound perception is to make sounds and listen how different manipulations affect
how what you make hears. Just go to the website and download the freeware to a disk that is available
to you; run the program and play with sounds while listening on headphones or good loudspeakers to
optimize perception. Relevant to material and exercises covered during Lecture Series I though Ill and in
Sections. Have fun with it!
RECOMMENDED BOOKS Or CD
• hnp://www.brainmusic.orWMBB91 Webpage/Neurobiol0l.html This page on the Institute for Music &
Brain Science website contains the Course Syllabus, selected lecture slides and images (in the form of
powerpoint presentations), section material, pdf's of relevant literature, other types of files, and more — the sum
total of which will exceed the space of our course site on the FAS server. The home page,
www.Brain'Music.org, has several links that are likely to be of interest to students in the course.
• Anatomy and &jollity of Hearing for Andiokgists, William Clark & Kevin Ohlemiller, Thomson Delmar,
ClifionPark, NY, 2007. At Coop and On Resent. One way to learn how something works is to find out how it
breaks. This book does a good job of teaching health professionals who specialize in the diagnostic evaluation
of patients with hearing loss what nwnt wrong with the basic biological mechanisms in their patients. So, if you
read this book (or at least parts of it), you'll make connections between what you learn about the neurobiology
of normal hearing with what you learn about the pathophysiology and pathology of hearing loss. Excellent
source for current knowledge about the peripheral auditory system, but not the central auditory system;
consequently, won't help you gain insights into neural mechanisms underling percept formation and auditory
cognition in language and music.
• The Central Auditory ,S:ystem, Gunter Ehret & Raymond Remand (eds), Oxford Press, NY 1996. At Coop and On
Resent This is one of the few auditory central nervous system textbooks that covers the single- and multi-unit
physiology of auditory cortex, thalamus, midbrain, and medulla in the kind of detail appropriate for a high-level
auditory neurobiology course. However, it is an edited book, so it is not surprising that it lacks a central theme
to tie the chapters together, other than the fact they are about mkrophysiology, microanatomy, and signal
processing. That said, the much of the material covered in Lertan Series III: Signal Processing In, tbe Auditory
Mims System, will draw from literature covered in this book.
• Spikes: F-xploring the Neural Code, Fred Rieke, David Worland, Rob deRuytervanSteveninck, William Biakk,
M.I.T. Press, 1999. At Coop and On Resent. This book is of special interest for students wishing to explore
perceptual-physiological correlates from the perspectives of systems neurobiology, computational neuroscience,
biophysics, mathematical modeling, and probability and statistics. Relevant to ',trine Series III: .figna/Proassing
Iry the Auditory Nervous System.
• Cambridge Engdopedia of Language, 7 d Ed, David Crystal, Cambridge Univ Press, 1997. At Coop and On Resenr.
This large book treats the term "language" in its broadest sense across multiple disciplines. The sections on
phonetics, phonology, prosody, evolution, and development are relevant to I_Inimre Series II: Amditog
Communication and 1-mturr and briars Series IV: Developmental Biology of the Auditory System. The figures are
excellent and the condensed format expedites covering the bare essentials. You will need to go beyond this
encyclopedia to get a fuller and more accurate picture of each topic.
• Harvard Dictionary of Music, C Ed, D. Randle (ed), Harvard Press, Cambridge, MA, 2003. At Coop and On
Resent. Although coursework and performance in music is not a pre-requisite for this course, you will be
required to read about the acoustics, phonology, and grammar of popular Western music (e.g., rock, classical)
during the course. Entries in music dictionaries (e.g., "harmonic analysis", "triad") can be quite informative,
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especially after some basic knowledge has been acquired in lectures and sections. At Coop 6' On Resent. If you
buy the Dictionary. at Harvard Coop, Harvard Univ Press, or elsewhere, please make sun you get the e edition.
• Co-Planar Stereotaxic Atlas oft& Human Brain: 3-Dimensional Proportional System: An Approach to Cerebral Imaging,
Jean Talairach and Pierre Toumoux. On Resent. Neuroanatomy always presents a special challenge — short
books with good pictures — and labels - are hard to find. Atlases are very expensive and include way too much
material. Students often overestimate how difficult this topic is and how much you need to know for a count
like this — partly because we aren't clear about what is important to learn and what isn't You need to learn
the anatomy of the cerebral cortex for Lecture Series II and III. When it comes to the cerebral cortex, consider
this: there are only two hemispheres; each of which has only four lobes and about two dozen gyri (same Greek
root as the sandwich "gyros"). You need to know the names of these lobes and gyri. That's the base
knowledge. Then, get some sense of how they relate to other, more indirect or abstract designations (e.g.,
Brodmann's "area 22", "Wemicke's area", "auditory" cortex ). There are several books at Cabot Library and
the Medical School Library (Countway) that are excellent resources - just use "Human Neuroanatomy" as a
Hollis search term. There are also several resources available on the Internet. You don't need to learn about
the the spinal cord, spinal nerves, or cranial nerves (except the "auditory nerve", a.k.a., "cranial nerve VIII",
"vestibulocochlear nerve') for this course. The Talairach Atlas of the cerebral hemispheres is in widespread
use in current neuroimaging research.
• Colorful Introduction to the Anatonry of the Human Brain, A: A Brain and Rrydrology Coloring Book, John P.J. Pined,
Maggie E. Edwards, Boston: Allyn & Bacon, Inc., 1997. On Resent. Interactive learning, the old-fashioned way.
• Introduction to the Porbologe of Hearing 0 Edition, Moore B, Cambridge Univ, 2003. On Resent. The most
popular "psychology of hearing" book for over two decades, this small paperback is single-authored by one of
the world's leading psychoacousticians. It has more psychology and less physiology and anatomy than Yost's
book. Although "psychology of hearing" is the title, it's really an introduction to the subdiscipline,
psychophysics of hearing, for which it can be used as an abridged reference text; it does not cover the
subdiscipline, cognitive psychology of hearing, which our course also covers.
• IntroducingPbonetic Science, M Ashby & J Alaidment , Cambridge U Press, NY, 2005. On Resent. The authors do
an excellent job explaining and illustrating some of the key points about phonetics and phonology that are
introduced in 1-seturr Series 11: Auditory Communication.
• Dictionary off-intrust/is and Phonetics, 0 Ed, David Crystal, Blackwell, Boston, 2003. On Resent. Precise and
sometimes in-depth definitions of terms we introduce in Lecture Series 11: Auditory. Communication.
• The Audio Dirtionag, 7 4 Ed, Glenn White, U Wash, Seattle, WA, 1991. Another resource for clarifying the
meaning of recondite tents — in this instance, terms used to describe different types of acoustic signals
described in Lecture Series II and III. We didn't put this on reserve.
• Acoustical Society of Amenra Auditory Demonstration CD. Trying to understand, no less talk about, auditory
experiments on pitch, harmony, timbre, etc. is as frustrating as trying to follow circuitous directions involving
twists and turns around streets with names you've never heard before — and can't even pronounce! Words
just don't do it It's better to listen to the stimuli used in psychoacoustic experiments. Houtsma and Rossing
made this "Best of" compilation CD for that reason. If you hear the stimuli and understand what the
experiments were about (concise explanations of each track and related experiments are included), your reading
of Yost and related recommended books will be clearer. We will use some of the tracks in Lecture Sena II:
/Irak°, Communication. The CD is published by the Acoustic Society of America. Send orders for the "ASA
Auditory Demonstrations on Compact Disc" to Acoustical Society of America (ASA) Publications, P.O. Box
1020, Sewickley, PA 15143-9998; Tel: 412-741-1979; Fax: 412-741-0609; Email (for inquiries):
asapubs@abdintl.com. Price: ASA members $23; Nonmembers $31. Postage and handling charges are: US.
orders - $6.00 for first title; $2.00 for each additional title; Prepay by check or money order in U.S. funds
drawn on a US. bank or by Mastercard, Visa, or American Express credit cards.
END 9/15/07
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