Neuroscience
L-200 Sterling Hall of Medicine, 785.5932
M.S., M.Phil., Ph.D.
Directors of Graduate Studies
Haig Keshishian (Molecular, Cellular, & Developmental Biology) (KBT 640,
432.3478, haig.keshishian@yale.edu)
Charles Greer (Neurosurgery; Neurobiology) (LSOG 221, 785.4034, charles.greer@yale.edu)
Professors
George Aghajanian (Psychiatry; Pharmacology), Colin Barnstable
(Ophthalmology & Visual Science; Neurobiology), Linda
Bartoshuk (Surgery; Epidemiology; Psychology), Walter Boron
(Cellular & Molecular Physiology), Benjamin Bunney (Psychiatry;
Pharmacology), John Carlson (Molecular, Cellular & Developmental
Biology), Lawrence Cohen (Cellular & Molecular Physiology),
Nigel Daw (Ophthalmology & Visual Science; Neurobiology),
Pietro De Camilli (Cell Biology), Ronald Duman (Psychiatry;
Pharmacology), Barbara Ehrlich (Pharmacology; Cellular &
Molecular Physiology), Patricia Goldman-Rakic (Neurobiology;
Psychology), Charles Greer (Neurosurgery, Neurobiology), Susan
Hockfield (Neurobiology), Marcia Johnson (Psychology), Leonard
Kaczmarek (Pharmacology; Cellular & Molecular Physiology),
Kenneth Kidd (Genetics; Molecular, Cellular & Developmental
Biology; Psychiatry), Jeffery Kocsis (Neurology; Neurobiology),
Robert LaMotte (Anesthesiology; Neurobiology), Thomas Lentz
(Cell Biology), Laura Manuelidis (Neuropathology), David McCormick
(Neurobiology), Bita Moghaddam (Psychiatry; Neurobiology),
Mark Mooseker (Molecular, Cellular & Developmental Biology;
Cell Biology), Frederick Naftolin (Obstetrics & Gynecology;
Molecular, Cellular & Developmental Biology), Angus Nairn
(Psychiatry), Pasko Rakic (Neurobiology), J. Murdoch Ritchie
(Pharmacology), Robert Roth (Psychiatry; Pharmacology), Gary
Rudnick (Pharmacology), W. Mark Saltzman (Chemical Engineering;
Biomedical Engineering), Joseph Santos-Sacchi (Surgery; Neurobiology),
Ilsa Schwartz (Surgery; Neurobiology), Steven Segal (Epidemiology;
Cellular & Molecular Physiology), Gordon Shepherd (Neurobiology),
Frederick Sigworth (Cellular & Molecular Physiology),
Stephen Strittmatter (Neurology; Neurobiology), Allan Wagner
(Psychology), Stephen Waxman (Neurology; Pharmacology), Robert
Wyman (Molecular, Cellular & Developmental Biology), Steven
Zucker (Computer Science)
Associate Professors
Meenakshi Alreja (Psychiatry; Neurobiology), Amy Arnsten (Neurobiology),
Charles Bruce (Neurobiology), Nihal de Lanerolle (Neurosurgery;
Neurobiology), Paul Forscher (Molecular, Cellular & Developmental
Biology), James Howe (Pharmacology), Marina Picciotto (Psychiatry;
Pharmacology; Neurobiology), George Richerson (Neurology;
Cellular & Molecular Physiology), Michael Schwartz (Neurobiology),
Jane Taylor (Psychiatry), Flora Vaccarino (Child Study Center;
Neurobiology), Michael Westerveld (Neurosurgery), Anne Williamson
(Neurosurgery; Neurobiology), Tian Xu (Genetics) Assistant Professors
Patrick Allen (Psychiatry), Hal Blumenfeld (Neurology; Neurobiology),
Angélique Bordey (Neurosurgery), Wei Chen (Neurobiology),
R. Todd Constable (Diagnostic Radiology; Neurosurgery), Maria
Donoghue Velleca (Neurobiology), Reiko Maki Fitzsimonds (Cellular
& Molecular Physiology), Karyn Frick (Psychology), Lise
Heginbotham (Molecular Biophysics & Biochemistry), Anthony
Koleske (Molecular Biophysics & Biochemistry), Mark Laubach
(Neurobiology), Christy Marshuetz (Psychology), Dhasakumar
Navaratnam (Neurology; Neurobiology), Vincent Pieribone (Cellular
& Molecular Physiology), Maria Mercedes Piñango
(Linguistics), Elke Stein (Molecular, Cellular & Developmental
Biology), Ning Tian (Ophthalmology & Visual Science),
David Wells (Molecular, Cellular & Developmental Biology),
Mark Yeckel (Neurobiology), David Zenisek (Cellular &
Molecular Physiology), Weimin Zhong (Molecular, Cellular &
Developmental Biology)
Research Scientists
Joel Black (Neurology), Nicholas Carnevale (Psychology)
Fields of Study
The Interdepartmental Neuroscience Program offers
flexible but structured interdisciplinary training for independent
research and teaching in neuroscience. The goal of the program
is to ensure that degree candidates obtain a solid understanding
of cellular and molecular neurobiology, physiology and biophysics,
neural development, systems and behavior, and neural computation.
In addition to course work, graduate students participate
in a regular journal club, organize the Interdepartmental
Neuroscience Program Seminar Series, and attend other seminar
programs, named lectureships, symposia, and an annual research
retreat.
Special Admissions Requirements
Applicants to the Neuroscience Program should have
a B.S. or B.A. Most applicants have had course work in neuroscience,
psychobiology, physiological psychology, mathematics through
calculus, general physics, general biology, general chemistry,
organic chemistry, biochemistry, computer science, or engineering.
Deficiencies in these areas can be corrected through appropriate
course work in the first year of residence. Laboratory research
experience is desirable but is not a formal requirement. Scores
for the GRE (General Test required; Subject Test recommended)
or MCAT, three letters of recommendation, transcripts of undergraduate
grades, and a statement of interest must accompany the application.
To enter the Ph.D. program, students apply to an interest-based
track within the interdepartmental graduate program in the
Biological and Biomedical Sciences.
Special Requirements for the Ph.D. Degree
Each entering student is assigned a faculty advisory
committee to provide guidance. This committee is responsible
for establishing the student’s course of study and for
monitoring his or her progress. This committee will be subsequently
modified to include faculty with expertise in the student’s
emerging area of interest. Although each student’s precise
course requirements are set individually to take account of
background and educational goals, the course of study is based
on a model curriculum beginning with three core courses (Principles
of Neuroscience, Neurobiology, and Structural and Functional
Analysis of the Human Nervous System) designed to ensure broad
competence in modern neuroscience. Students are also required
to complete at least three additional courses from a broad
set of neuroscience-related courses. The Graduate School uses
grades of Honors, High Pass, Pass, and Fail and requires two
term grades of Honors during the first two years of study.
Students are expected to maintain at least a High Pass average.
A series of at least two laboratory rotations during the first
year of the program also ensures that degree candidates obtain
a solid background in systems, cellular, and molecular approaches
to neuroscience. Admission to candidacy requires passing a
qualifying examination normally given during the second year,
and submission of a dissertation prospectus (NIH grant format)
before the end of the third year. In accordance with the expectations
of the BBS program, Ph.D. students are expected to participate
in two terms (or the equivalent) of teaching. Thesis committee
meetings are required annually. Also required is the completion
and satisfactory defense of the thesis.
Requirements for M.D./Ph.D. students are the same as for
Ph.D. students with the following differences: five courses
are required (Principles of Neuroscience and Structural and
Functional Analysis of the Human Nervous System, and three
elective graduate level courses). M.D./Ph.D. students are
required to serve for one term as teaching assistants; however,
two terms of teaching are preferred.
Master's Degrees
M.Phil. See Graduate
School requirements.
M.S. Awarded only to students who are not continuing
for the Ph.D. degree but who have successfully completed one
year of the doctoral program. Students are not admitted for
this degree.
Program materials are available upon request to the Director of Graduate Studies, Neuroscience, Yale University, PO Box 208074, New Haven CT 06520-8074.
Courses
NSCI 501a, Principles of Neuroscience. Marina
Picciotto, Mark Yeckel. WF 3.15–4.45
General neuroscience seminar: lectures, readings, and
discussion of selected topics in neuroscience. Emphasis is
on how approaches at the molecular, cellular, physiological,
and organismal levels can lead to understanding of neuronal
and brain function. Also NBIO 501a.
NSCI 504b, Seminar in Brain Development and Plasticity. Weimin
Zhong. MW 2.30–3.45
Weekly seminars (Monday) and discussion sessions (Wednesday)
to explore recent advances in our understanding of brain development
and plasticity, including neuronal determination, axon guidance,
synaptogenesis, and developmental plasticity. Also MCDB
735bu.
[NSCI 506b, Introduction to Brain and Behavior.]
NSCI 507b, Cellular and Molecular Mechanisms of Neurological
Disease. Dhasakumar Navaratnam, Stephen Strittmatter,
Stephen Waxman.
Focuses on those diseases in which modern neuroscience
has advanced mechanistic explanations for clinical conditions.
The application of pathophysiologic understanding to therapeutics
is considered.
[NSCI 508a, Functional Properties of Cortical Neurons
and Circuits.]
NSCI 510b, Structural and Functional Organization of
the Human Nervous System. Michael Schwartz, Pasko Rakic.
An integrative overview of the structure and function
of the human brain pertaining to major neurological and psychiatric
disorders. Also NBIO 500b.
[NSCI 514a, The Regulation of Cell Fate during CNS Development.]
NSCI 519a/b, Tutorial.
By arrangement with faculty and approval of the director
of graduate studies.
[NSCI 521a, Neuroimaging in Neuropsychiatry.]
NSCI 539b, Synaptic Organization of the Nervous
System. Gordon Shepherd, Anne Williamson, Michael
Hines.
An integrative introduction to the principles underlying
the organization of neural systems. The focus is on the best-understood
systems, including spinal cord, olfactory bulb, retina, cerebellum,
thalamus, basal ganglia, and cerebral cortex. Students integrate
experimental findings from anatomy, electrophysiology, and
neuropharmacology with computational models at the cellular
and circuit level.
[NSCI 540a, Introduction to Statistics in Psychology.]
NSCI 600a, Experimental Methods in Neuroscience. R.
Todd Constable. TTh 2–4
This course examines the experimental techniques currently
available for the neuroscientist. It explores the kinds of
information obtainable in studying phenomena ranging from
electrophysiological recordings of individual neurons to metabolic
processes, from ensembles of neurons to behavioral output.
Techniques covered include microscopic methods (light, electron),
electrophysiology (extracellular/intracellular single cell
recordings, multiple cell recording methods, brain slices),
macroscopic methods (ERP, MEG, TMR), metabolic measures (microdialisis,
biosensors, MR spectroscopy), imaging approaches (optical
tomography, PET, SPECT, functional MRI), and interventional
techniques (lesions, cortical stimulation, knockout genetics,
surgery, drugs). The knowledge gained from each of these approaches,
the limitations of the methods, and future developments are
considered.
[NSCI 605b, Pathways of Discovery in Neuroscience.]
NSCI 614a, Neurobiology of Learning and Memory. Thomas
Brown. M 11.30–2
This seminar integrates hypotheses and research methods
used to elucidate the neurobiological mechanisms underlying
learning and memory. Levels of analysis range from molecular
and cellular to systems and behavioral, with a primary focus
on cellular and systems neurophysiology. Discussion includes
the philosophy and rationale underlying some of the more successful
and interesting methods. A goal is to evaluate critically
how one might connect synaptic phenomena such as long-term
potentiation and depression to behavioral changes such as
acquisition and extinction. Focus is on combining in vitro
and in vivo methods that offer the possibility for yielding
quantitative theoretical or computational models. Also
PSYC 572a.
[NSCI 646, Advances in Cognitive Neuroscience: Prefrontal
Cortex and Memory.]
NSCI 654b, Sensory Processes. Lawrence Marks,
Joseph Stevens.
A course on the senses, emphasizing functional properties
of human vision, hearing, taste, smell, and skin senses.
NSCI 720a, Neurobiology. Haig Keshishian,
Paul Forscher. MWF 11.30–12.20
Examination of the excitability of the nerve cell membrane
provides a starting point for the study of molecular, cellular,
and intracellular mechanisms underlying the generation and
control of behavior. Also MCDB 720au, NBIO 720a.
The following course is also of particular value to students
in Neuroscience:
MCDB 721Lau, Laboratory for Neurobiology. Haig
Keshishian, Robert Wyman.
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