Neurobiology 204: Neurophysiology of Central Circuits


Spring Term 2008


Co-Directors: Rick Born and Rachel Wilson

(rborn[at], 432-1307; rachel_wilson[at], 432-5571)


Faculty: John Assad, Margaret Livingstone, John Maunsell, Clay Reid

TA: Andreas Liu (andreasliu[at]


Time: Mondays and Wednesdays from 10:00 a.m. to 12:00 p.m.

Location: Warren Alpert Building, Rm. 336.


This course serves as an introduction to the physiology of circuits in the vertebrate central nervous system. Topics include the visual, auditory, olfactory, and somatosensory systems, the neural control of eye movements, reward, and perception. The behavior of these systems will be analyzed at three levels: the electrophysiological properties of single neurons, synaptic interactions between neurons in vitro, and the behavior of the circuits in vivo.



Neuro-200/HST-130 (“Introduction to Neuroscience”) is recommended but not strictly required for students who are not Ph.D. candidates in the Harvard Neuroscience Graduate program. If you have not taken this course, please let the instructors know. We can recommend general review articles or other materials that will help give you the background that most of the other students will have.



Wednesday classes will generally consist of a two-hour lecture. On Mondays we will discuss one or more papers. The schedule below is also available as a Google Calendar.  You can either click on the link in the upper left corner or search “NB204” under public calendars at Google Calendar.  Discussions of papers in class should be very interactive. Each student should be prepared to discuss the following for each of the assigned papers: 1) The big picture (abstract and introduction), 2) methods, 3) each figure, 4) main conclusions.








Jan. 30

Concept: Receptive fields & neural codes (Born/Wilson)

Handout, Zador paper, Wang paper


Feb. 4

Discussion (Born/Wilson)

Feb. 6

Vision: Retina to V1 (Reid)

Handout, Ohki review, Wickersham paper


Feb. 11

Discussion (Reid)

Feb. 13

Concept: Circuits & connectivity (Reid)

Handout, Denk review


Feb. 18

Holiday: President's day

Feb. 20

Somatosensory system (Wilson)



Feb. 25

Discussion (Wilson)

Feb. 27

Olfactory system (Wilson)



Mar. 3

Discussion (Wilson)

Mar. 5

Auditory system (Assad)

Handout, Carr & Konishi 1990


Mar. 10

Discussion (Assad)

Mar. 12

Concept: Cortical Maps (Born)

Handout,  Barlow review, Chenn & Walsh 2002, Fukuchi-Shimogori & Grove 2001


Mar. 17

Discussion (Born)

Mar. 19

Sensory integration (Born)

Handout, Livingstone & Hubel 1988, Nadler & DeAngelis 2008 (supplemental materials)


Mar. 24


Mar. 26



Mar. 31

Discussion (Born)

Apr. 2

Vision: Direction selectivity (Livingstone)

Handout, Barlow & Levick 1965, Fried et al. 2002


Apr. 7

Discussion (Livingstone)

Apr. 9

Concept: Neurons & perception (Maunsell)

Handout, Parker & Newsome 1998, Heuer & Britten 2004


Apr. 14

Discussion (Maunsell)

Apr. 16

Basal ganglia & reward (Assad)

Handout, Schultz 2006, Hollerman & Schultz 1998, Platt & Glimcher 1999


Apr. 21

Discussion (Assad)

Apr. 23

Cerebellum & motor learning (Born)

Handout, Raymond et al. 1996, De Zeeuw et al. 1998


Apr. 28

Discussion (Born)

Apr. 30

Hand out Final Exam




The grade will be computed as follows:

homework - 40%

final exam - 40%

class participation - 20%

Grades will be lowered for repeated absenteeism or arriving late to class.  



Homework assignments are due before class Monday.   Please e-mail homework to andreasliu[at] by 10:00am Monday.  Name your file nb204surnameX.doc, where X is the week of the assignment (e.g. nb204smith3.doc in week 3) and please put “nb204” in the subject line. Homework assignments will be based on the papers we discuss on class on Mondays. The faculty member teaching that week may have special instructions for your written assignment, so check the mini-syllabus before starting to read and write.  In general, you will be instructed to write either a referee’s report or a commentary.  Make sure you know which type is being assigned each week. Occasionally a different type of written assignment might be specified; in this case, the instructor will give you instructions about what they want to see.  A detailed description of the writing curriculum can be found here.


Final exam

Your final exam will be to write one referee’s report and one commentary, each on a different paper assigned by the faculty at the end of the course. Use the feedback you receive on your homework to polish your writing skills within these two specific formats. Before starting to write, please review the big ideas presented in the four “concept lectures” during the semester. Think about what major concepts covered during the course this paper pertains to, and use your understanding of these concepts as you write your referee’s report and your commentary.



In previous years, class members have requested additional background on some of the statistical and analytical methods we encounter in the course.  To meet this need, we have added two new features.


1. Special Sessions. We are lucky to have Andy Liu as our teaching assistant this spring.  Andy is a mathematician who has spent time thinking seriously about neurobiology while a member of Wade Regehr's lab.  Andy will be offering four additional evening sessions on selected topics for students who need or want more mathematical background.  The topics are:


1) basic probability and distributions

2) basic information theory and coding with ensembles

3) reverse correlation, white noise analysis

4) signal detection, ROC


The meeting times for these sessions will be established at the first class meeting on Jan. 30, 2008.


2. On-line Primers.  Background reading on the above topics can be found here.