Neurobiology 101. Auditory Neurobiology of Language & Music

Harvard College & Graduate School of Arts & Sciences, Fall Term, 2007-2008 Academic Year

Lectures Tu & Th, 11:30am-1:00pm
1 section per week (90 min.), TuWThFri, Afternoon and Evening
Biological Laboratories, Divinity Ave, Room 1068 (note new room number)

Last Update: 8/24/07


Teaching Fellow & Guest Lecturers
Course Description
Requirements & Grading
Books: Required
Books & Other Resources: Recommended
Lecture & Section Schedule
Class Notes & Selected Lectures (PowerPoint Files)


Mark Jude Tramo, MD, PhD
Asst. Professor of Neurology, Harvard Medical School
Steering Committee, Harvard University Mind/Brain/Behavior Interfaculty Initiative
Biology Teaching Affiliate, Harvard University
Board of Psychology Honors Tutors, Harvard University
Attending Neurologist, Massachusetts General Hospital
Songwriter Member, ASCAP

Jonathan I. Matsui, PhD
Lecturer in Molecular & Cellular Biology, Harvard University
Neurobiology Concentration Advisor for Harvard College Undergraduates, Harvard Faculty of Arts & Sciences
Office hours by appointment only, BioLabs 1082a

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Teaching Fellow & Guest Lecturers

Teaching Fellow
Caroline Niziolek, MS
PhD Candidate, Harvard-MIT Speech & Hearing Bioscience & Technology Graduate     Studies Program
B.S., Brain & Cognitive Sciences, M.I.T.
Office hours: TBD

Guest Lecturers
Louis D. Braida, PhD
Henry Warren Ellis Professor of Health Sciences & Technology, Harvard Medical     School -M.I.T. Division of Health Sciences & Technology
Professor of Electrical Engineering & Computer Science, Research Laboratory of     Electronics, M.I.T.
Co-Director, Harvard-M.I.T. Speech & Hearing Biosciences & Technology Graduate     Studies Program

Marc D. Hauser, PhD
Harvard College Professor of Psychology and of Organismic & Evolutionary Biology
Co-Director, Harvard University Mind/Brain/Behavior Interfaculty Initiative

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Course Description

      How does the brain transform the 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 brain are necessary and sufficient to understand spoken, seen, or imagined words, groove to music, or 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? 
      During this 13-week lecture course, we bring knowledge and methodologies from multiple scientific and artistic disciplines (biology, engineering, computer science, psychology, linguistics, music) to bear on these questions. We introduce current concepts about 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 throughout last semester); and  4) how implicit knowledge about language and music learned after years and years of acculturation influences auditory perception.  Students will be expected to master topics in systems neurobiology, molecular and cellular neurobiology, and other disciplines.  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 graduate and medical school education.  Emerging treatment and education technologies and ideas for future research and development in the brave new worlds of neurobionics and genetic engineering will be explored.

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MCB 80. Neurobiology of Behavior
Secondary School Physics or Engineering
Secondary School Algebra and Trigonometry
Not required: Ability to read or play music

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Requirements & Grading

Biweekly Lecture Attendance & Weekly Section Participation (20% of grade)
One Mid-Term Examination (30% of grade)
Final Examination (50% of grade)
Problem Sets (to be determined)

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Books: Required

1. Webpage/Neurobio101.html - This URL contains the Syllabus, Class Notes, and Selected Lectures (in the form of PowerPoint
presentations) given by Drs Tramo, Matsui, Niziolek, Braida, and Hauser.  Parent
home page, which is likely to be of interest to many students:
2.  Fundamentals of Hearing, 5th Ed, 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 neuroimaging.
3.  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.  Mac, IBM, and other platforms are supported.

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Books & Other Resources (recommended)

Auditory Neurobiology
      The Central Auditory System, Gunter Ehret, Raymond Romand, Oxford Press, NY 1996.  This is one of the few auditory CNS textbooks, albeit edited, 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.  Although this book is out of print, there appears to be a sufficient number of copies available via to expect the Harvard Coop can avail 10 for sale; we will also request that one be placed on reserve at Cabot Library.
      Anatomy and Physiology of Hearing for Audiologists, William Clark & Kevin Ohlemiller, Thomson Delmar, CliftonPark, NY, 2007.  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 went 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 in the first part of the course with what you learn about hearing loss in a later part of the course.  20 copies to be placed on sale at the Harvard Coop.
      Spikes: Exploring the Neural Code, Fred Rieke, David Warland, Rob
deRuytervanSteveninck, William Bialek, M.I.T. Press, 1999.  Relevant to our lecture series on Signal Processing in the Auditory Nervous System, and likely to be of particular interest for students wishing to explore computational neuroscience, bioengineering, biophysics, mathematical modeling, probability and statistics, neural coding, and single-unit neurophysiology.
     Colorful Introduction to the Anatomy of the Human Brain, A: A Brain and Psychology Coloring Book,
John P.J. Pinel, Maggie E. Edwards, Boston: Allyn & Bacon, Inc., 1997. Neuroanatomy always presents a special challenge - short, books with good pictures are hard to find, and atlases are expensive and include way too much material.  Students often overestimate how difficult this topic is and how much you need to know for a course like this — partly because anatomists stubbornly invent their own nomenclatures rather than share each others’, partly because we aren’t clear about what is important to learn and what isn’t.  Consider this: there are two  hemispheres; each of which has only four lobes and about two dozen lobules/gyri (same Greek root as the sandwich “gyros”).  You need to know the names of these lobes, lobules, and gyri, and how they differ/relate to other designations (e.g., Brodmann’s numbers).  At least you will then know more than many neuroscientists, even some of your own professors and TFs, who have been caught from time to time using these terms incorrectly!  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.  You don’t need to learn about nerves and the spinal cord for this course.  The Talairach Atlas of the cerebral hemispheres (below) is commonly in widespread use at present.
      Co-Planar Stereotaxic Atlas of the Human Brain: 3-Dimensional Proportional System : An Approach to Cerebral Imaging, Jean Talairach and Pierre Tournoux - 3 requested for sale at Harvard Coop (>$200 ea!); one requested for reserve at Cabot Library

Auditory Perception & Cognition
      Acoustical Society of America Auditory Demonstration CD.  Talking about auditory experiments on pitch, loudness, harmony, etc is as frustrating as talking directions with 20 turns.  Often, words just don’t do it.  It’s way better to listen to the experimental tasks on this CD (some of which we plan to play) than to hear us talk or read prose explaining them. After you hear the stimuli and understand what the experiments were about, reading Yost, Moore (see below), and the explanations that come with the CD make more sense.  We have not ordered any via Harvard Coop or for the library since the CD is easy to get - Here’s how:
Send orders for “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): Price: ASA members $23; Nonmembers $31. Postage and handling charges are: U.S. 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 U.S. bank or by Mastercard, Visa, or American Express credit cards.
     Introduction to the Psychology of Hearing, 5th Edition, Moore B, Cambridge Univ, 2003 - Currently the “psychophysics of hearing” book by the world’s leading psychoacoustics, this is a surprisingly small and digestible text that one can read cover to cover or use a reference text.  Good for getting at some of the details of gnarly issues like: which harmonics in a harmonic series, and how many, are needed to make the pitch of a missing first harmonic (a.k.a. “the missing fundamental frequency”) strong?  Although “Psychology” is in the book’s title, don’t be misled: little about the cognitive psychology of hearing can be found in these pages.

Language & Music
      Introducing Phonetic Science, M Ashby & J Maidment , Cambridge U Press, NY, 2005.  We could have recommended many books for language and music - after all, they are such interesting topics!  But our focus in this course is on their underlying neural mechanisms as they relate to perception and development, so we are keeping the book list limited to dictionaries and an encyclopedia, except for this book by Ashby & Maidment, which does a superb job explaining and illustrating the key points you need to know about speech perception and production.  We’ve requested that 20 be put on sale at the Harvard Coop. 
      The Harvard Dictionary of Music, 4th Ed, D. Randle (Ed), Harvard Press, Cambridge, MA, 2003.  20 on sale at Harvard Coop.  We’ll ask that the copy in the Music Library be put on reserve there.
      Dictionary of Linguistics and Phonetics, 5th Ed, David Crystal, Blackwell, Boston, 2003.  Ten on sale at Harvard Coop.  One on reserve at Cabot Library.
      Cambridge Encyclopedia of Language, 2nd Ed, David Crystal, Cambridge Univ Press, 1997.  Ten on sale at Harvard Coop.  One on reserve at Cabot Library.

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Lecture and Section Schedule
Tuesday, Sept. 18 Course Overview; The Hearing Brain Tramo  
  No Section in Week 1    
Thursday, Sept. 20 Pictures of Sound Tramo  
Friday, Sept. 21

Study Card Day

Students need to complete electronic sectioning (if needed) by Friday at 5:00 PM.

Tuesday, Sept. 25 Avada Kedavra! Or, Listening to "Prozac" Phonetics, Phonology, & Prosody in relation to Grammar, Semantics, & Semiotics in Babble, Interjections, Conversation, Prose, Poetry, Song, Music, Film, Puns, Branding, & Clanging Tramo  
  Section #1: Pictures of Sound & Acoustical Measurements Niziolek  
Thursday, Sept. 27 Neurophysiology & Neuroanatomy of Language I Tramo  
Monday, Oct. 1 Last day for Undergraduates to drop this course or change status or course grade (letter-grade vs. pass-fail) without having to pay the Registrar a fee    
Tuesday, Oct. 2 Neurophysiology & Neuroanatomy of Language II Tramo  
  Section #2: Acoustic Phonetics & Psychoacoustics Laboratory Demonstrations Niziolek  
Thursday, Oct. 4 Neurophysiology & Neuroanatomy of Language III Tramo  
Tuesday, Oct. 9 Evolution of the Language Faculty Hauser  
  Section #3: Human Brain Imaging Niziolek  
Thursday, Oct. 11 Neurophysiology & Neuroanatomy of Music I Tramo  
Monday, Oct. 15 Fifth Monday: Last day for Undergraduates to drop this course or change status
of course grade (letter-grade vs. pass-fail)
Tuesday, Oct. 16 Neurophysiology & Neuroanatomy of Music II Tramo  
  Section #4: Electro- & Magnetoencephalography (EEG & MEG): Evoked Potentials & Event Related Potentials (EPs & EPRs) Niziolek  
Thursday, Oct. 18 Neurophysiology & Neuroanatomy of Music III Tramo  
Tuesday, Oct. 23 Comparisons Between Language & Music Tramo  
  Section #5: Review of Auditory Communication Niziolek  
Thursday, Oct. 25 Acoustic Signal Processing: Ear & Hair Cell Sensory Receptors Matsui  
Monday, Oct. 29 Last day for Graduate Students to Add/Register for this course    
Tuesday, Oct. 30 Spiral Ganglion Cells & Auditory Nerve Tramo  
  Section #6: Physiology & Anatomy of the Auditory System Niziolek  
Thursday, Nov. 1

Acoustic Signal Transformation & Acoustic-Bioelectrical Transduction: Hearing Aids & Direct Electrical Stimulation of the Auditory Nerve to Treat Hearing Loss in Babies & Adults

Tuesday, Nov. 6 Auditory Brainstem Tramo  
  Section #7: Spike Trains and Neural Coding Niziolek  
Thursday, Nov. 8 Auditory Cortex I Tramo  
Tuesday, Nov. 13 Auditory Cortex II Tramo  
  Section #8: Review of Auditory Communication & Signal Processing by the Auditory System Niziolek  
Thursday, Nov. 15 Mid-term Examination: Auditory Communication & Signal Processing by the Auditory System  
Saturday, Nov. 17 Yale Game, at Yale    
Tuesday, Nov. 20 Review of Mid-Term Examination Tramo, Matsui, Niziolek  
Thursday, Nov. 22 Thanksgiving - NO CLASS, NO SECTION    
Tuesday, Nov. 27 Development of the Ear & Peripheral Auditory Nervous System Matsui  
  Section #9: "The (Neuro)Bionic Woman": Bioengineering the Future Niziolek  
Thursday, Nov. 29 Development of the Central Auditory Nervous System Matsui  
Tuesday, Dec. 4 Language Development in Infants & Children Matsui  
  Section #10: "Ra" vs. "La," Critical Periods, & Sign Language Niziolek  
Thursday, Dec. 6 Molecular Pathology & Pathophysiology of Genetic Diseases I Matsui  
Tuesday, Dec. 11 Molecular Pathology & Pathophysiology of Genetic Diseases II Matsui  
  Section #11: Congenital Hearing Loss & the Search for Mutations Niziolek  
Thursday, Dec. 13 Social Psychology of Developmental Communication Disorders Matsui  
Tuesday, Dec. 18

Review of Lectures Series IV & V: Molecular & Developmental Biology of the Auditory System

  No Section in Week 14    
  Section #12 (date TBD): Course Review Niziolek  


Exam Group 13: Mon., Jan 14, 2008
Exam Group 14: Thu., Jan 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 1, 7, 14
    Exam Groups 2, 15, 16
    Exam Groups 6, 10
    Exam Groups 8, 9, 11, 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.


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