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) (FMB 412, 785.4034, charles.greer@yale.edu)
Professors
George Aghajanian (Psychiatry; Pharmacology), Amy Arnsten (Neurobiology; Psychology), Colin Barnstable (Ophthalmology & Visual Science; Neurobiology), Linda Bartoshuk (Surgery; Psychology), Walter Boron (Cellular & Molecular Physiology), Benjamin Bunney (Psychiatry; Pharmacology; Neurobiology), John Carlson (Molecular, Cellular & Developmental Biology), Marvin Chun (Psychology), Lawrence Cohen (Cellular & Molecular Physiology), R. Todd Constable (Diagnostic Radiology; Neurosurgery), Pietro De Camilli (Cell Biology), Nihal de Lanerolle (Neurosurgery; Neurobiology), Ronald Duman (Psychiatry; Pharmacology), Barbara Ehrlich (Pharmacology; Cellular & Molecular Physiology), Paul Forscher (Molecular, Cellular & Developmental Biology), Charles Greer (Neurosurgery; Neurobiology), Tamas Horvath (Comparative Medicine; Neurobiology), Marcia Johnson (Psychology; Psychiatry), Leonard Kaczmarek (Pharmacology; Cellular & Molecular Physiology), Haig Keshishian (Molecular, Cellular & Developmental Biology), Kenneth Kidd (Genetics; Molecular, Cellular & Developmental Biology; Psychiatry), Jeffery Kocsis (Neurology; Neurobiology), Robert LaMotte (Anesthesiology; Neurobiology), Thomas Lentz (Cell Biology), Paul Lombroso (Child Study Center), Laura Manuelidis (Neuropathology), Gregory McCarthy (Psychology), David McCormick (Neurobiology), Mark Mooseker (Molecular, Cellular & Developmental Biology; Cell Biology; Pathology), Angus Nairn (Psychiatry; Pharmacology), Marina Picciotto (Psychiatry; Pharmacology; Neurobiology), Pasko Rakic (Neurobiology), George Richerson (Neurology; Cellular & Molecular Physiology), Robert Roth (Psychiatry; Pharmacology), Gary Rudnick (Pharmacology), W. Mark Saltzman (Chemical Engineering; Biomedical Engineering), Joseph Santos-Sacchi (Surgery; Neurobiology), Ilsa Schwartz (Surgery; Neurobiology), Gordon Shepherd (Neurobiology), Robert Sherwin (Internal Medicine), Frederick Sigworth (Cellular & Molecular Physiology; Biomedical Engineering), Stephen Strittmatter (Neurology; Neurobiology), Allan Wagner (Psychology), Xiao-Jing Wang (Neurobiology), Stephen Waxman (Neurology; Pharmacology; Neurobiology), Robert Wyman (Molecular, Cellular & Developmental Biology), Tian Xu (Genetics), Steven Zucker (Computer Science; Electrical Engineering; Biomedical Engineering)
Associate Professors
Meenakshi Alreja (Psychiatry; Neurobiology), Hilary Blumberg (Psychiatry; Diagnostic Radiology; Biomedical Engineering), Angélique Bordey (Neurosurgery; Cellular & Molecular Physiology), Charles Bruce (Neurobiology), Michael Crair (Neurobiology), Sabrina Diano (Obstetrics, Gynecology & Reproductive Services; Neurobiology), Karyn Frick (Psychology), James Howe (Pharmacology), Michael Koelle (Molecular Biophysics & Biochemistry), Anthony Koleske (Molecular Biophysics & Biochemistry; Neurobiology), Daeyeol Lee (Neurobiology), Vincent Pieribone (Neurobiology), Maria Mercedes Piñango (Linguistics), Michael Schwartz (Neurobiology), Matthew State (Child Study Center; Genetics), Jane Taylor (Psychiatry; Psychology), Ning Tian (Ophthalmology & Visual Science; Neurobiology), Vinzenz Unger (Molecular Biophysics & Biochemistry), Flora Vaccarino (Child Study Center; Neurobiology), Michael Westerveld (Neurosurgery; Pediatrics; Child Study Center), Anne Williamson (Neurosurgery), Weimin Zhong (Molecular, Cellular & Developmental Biology)
Assistant Professors
Patrick Allen (Psychiatry), Robert Beech (Psychiatry), Thomas Biederer (Molecular Biophysics & Biochemistry), Hal Blumenfeld (Neurology; Neurobiology), Sreeganga Chandra (Neurology; Molecular, Cellular & Developmental Biology), Wei Chen (Neurobiology), Ralph DiLeone (Psychiatry; Neurobiology), Jeremy Gray (Psychology), Elizabeth Jonas (Internal Medicine; Neurobiology), Sven-Eric Jordt (Pharmacology), Hür Köser (Electrical Engineering), Mark Laubach (Neurobiology), David LaVan (Mechanical Engineering), Erin Lavik (Biomedical Engineering), Michael Levene (Biomedical Engineering), James Mazer (Neruobiology), Rory McCrimmon (Internal Medicine), Dhasakumar Navaratnam (Neurology; Neurobiology), Michael Nitabach (Cellular & Molecular Physiology), Laurie Santos (Psychology), Samuel Sathyanesan (Psychiatry), Glenn Schafe (Psychology), Nenad Sestan (Neurobiology), Dana Small (Psychology; Surgery), Elke Stein (Molecular, Cellular & Developmental Biology), James Swain (Child Study Center), Susumu Tomita (Cellular & Molecular Physiology), David Wells (Molecular, Cellular & Developmental Biology), Mark Yeckel (Neurobiology), David Zenisek (Cellular & Molecular Physiology), Yufeng Zhou (Cellular & Molecular Physiology)
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 (BBS).
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 four core courses (Principles of Neuroscience, Neurobiology, Bioethics in Neuroscience, and Structural and Functional Organization 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 Organization 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 Degree 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. The minimum requirement for this is a passing grade in at least four courses, including two Honors grades, and two successful laboratory rotations. 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.
TTh 4–5
Focuses on those diseases (Alzheimer’s, Parkinson’s, ALS, and other neurodegenerative diseases, Triplet Repeat induced diseases, multiple sclerosis, epilepsy, etc.) in which modern neuroscience has advanced mechanistic explanations for clinical conditions. The course highlights recent molecular, electrophysiological, and imaging experiments in parsing disease mechanisms. The application of pathophysiologic understanding to therapeutics is considered. Also NBIO 507b.
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 as it pertains to major neurological and psychiatric disorders. Neuroanatomy, neurophysiology, and clinical correlations are interrelated to provide essential background in the neurosciences. Lectures in neurocytology and neuroanatomy survey neuronal organization in the human brain, with emphasis on long fiber tracts related to clinical neurology. Weekly three-hour laboratory sessions devoted to neuroanatomy in which students dissect the human brain and examine histological sections in close collaboration with faculty members. Lectures in neurophysiology cover various aspects of neural function at the cellular level, with a strong emphasis on the mammalian nervous system. Each student may participate in a weekly physiology conference with a faculty member, covering such topics as vision, sensory physiology, motor systems, simple nervous systems, or general neurophysiology. Clinical correlations consist of five sessions given by one or two faculty members representing both basic and clinical sciences. These sessions relate neurological symptoms to cellular processes in various diseases of the brain. Variable class schedule; contact course instructors. Also NBIO 500b.
[NSCI 514b, Neurodevelopment and Neuropsychiatric Disorders.]
NSCI 519a/b, Tutorial
By arrangement with faculty and approval of DGS.
NSCI 521a, Neuroimaging in Neuropsychiatry I: Imaging Methods. Julie Staley, Kelly Cosgrove.
W 9–10
Neuroimaging methodologies including Positron Emission Tomography (PET), Single Photon Emission Computed Tomography (SPECT), Magnetic Resonance Imaging (MRI), functional Magnetic Resonance Imaging (fMRI), Magnetic Resonance Spectroscopy (MRS), and gene array imaging (GAI) are rapidly evolving tools used to study the living human brain. Neuroimaging has unprecedented implications for routine clinical diagnosis, for assessment of drug efficacy; for determination of psychotropic drug occupancy; and for the study of pathophysiological mechanisms underlying neurologic and psychiatric disorders. This course is designed to provide an overview of the theory and current state of development of the different neuroimaging modalities. A second course, offered in the spring, focuses on applications. Also PHAR 521a.
NSCI 522b Neuroimaging in Neuropsychiatry II: Clinical Applications. Julie Staley, Kelly Cosgrove.
W 9–10
See description for NSCI 521a. This spring-term course focuses on applications.
[NSCI 535b, History of Modern Neuroscience.]
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. Also NBIO 509b.
[NSCI 540a, Introduction to Statistics.]
[NSCI 571b, Neurophysiology.]
NSCI 580a, The MAP Kinase Pathway and Cognitive Disorders. Paul Lombroso.
M 10–11.30
This seminar begins with a review of the classical MAP kinase pathway. Over the last decade, it has become clear that the function of this pathway in the brain differs from its function elsewhere in the body in that it plays a key role in learning and memory. The course begins with readings of seminal papers establishing the importance of MAP kinase in the induction, expression, and maintenance of LTP. Given the importance of the MAP kinase pathway in the establishment and consolidation of long-term memories, it should not be surprising that disruptions in this pathway lead to cognitive deficits. The seminar moves on to the molecular bases of several developmental disorders where mutations have been found in key players of the MAP kinase pathway.Some of the disorders covered include neurofibromatosis, fragile X syndrome, Rubenstein-Taybi syndrome, and Coffin-Lowry syndrome. Students are assigned papers and lead the discussions at each of the meetings.
NSCI 580b, Bioethics in Neuroscience. Charles Greer.
Th 4–5.30
This course is an introduction to ethics and ethical decision making in the neurosciences. Format for the course is an informal discussion. Each week, we are joined by members of the Yale faculty and community who can share their experiences and expertise as it relates to the topic of the week. This course is mandatory for first-year graduate students in the Interdepartmental Neuroscience Program (INP). Grading is Satisfactory/Unsatisfactory and is based on attendance/participation, weekly reaction papers, and a final term paper.
[NSCI 590a, Sensory Neuroethology: Bats, Owls, Electric Fish and Beyond.]
[NSCI 600a, Experimental Methods in Neuroscience.]
[NSCI 605b, Pathways of Discovery in Neuroscience.]
[NSCI 611a, Stem Cells and Approaches to Repair in the Nervous System.]
NSCI 612b, Molecular Transport and Intervention in the Brain. Mark Saltzman, Richard Carson.
This is a graduate-level seminar on mechanisms and rates of movement of molecules in the brain and the design of novel drug delivery systems. Topics include mathematical methods for modeling diffusion and flow processes, diffusion in the brain interstitium, fluid flows in the brain and spinal cord, the blood-brain barrier, microdialysis measurements, controlled release systems, microfluidic approaches for drug delivery. Weekly readings are assigned from neuroscience and engineering texts; current papers from the literature are used to guide discussion each week. Also ENAS 812b.
NSCI 614b, Neurobiology of Learning and Memory. Thomas Brown.
T 1.30–3.20
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 a 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 C&MP 675a, PSYC 572a.
[NSCI 645a, Foundations of Behavioral Neuroscience.]
[NSCI 646b, Advances in Cognitive Neuroscience: Prefrontal Cortex and Memory.]
[NSCI 648b, Cellular Analysis of Learning and Memory: Vertebrate Model Systems.]
[NSCI 654b, Sensory Processes.]
NSCI 720a, Neurobiology. Haig Keshishian, Paul Forscher.
MWF 11.35–12.25
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 720a, 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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