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), John Carlson (Molecular, Cellular & Developmental Biology), Marvin Chun (Psychology), Lawrence Cohen (Cellular & Molecular Physiology), R. Todd Constable (Diagnostic Radiology; Biomedical Engineering; 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), James Howe (Pharmacology), Marcia Johnson (Psychology; Psychiatry), Leonard Kaczmarek (Pharmacology; Cellular & Molecular Physiology), Haig Keshishian (Molecular, Cellular & Developmental Biology), Kenneth Kidd (Genetics; Ecology & Evolutionary Biology; Psychiatry), Jeffery Kocsis (Neurology; Neurobiology), Robert LaMotte (Anesthesiology; Neurobiology), Thomas Lentz (Cell Biology), Paul Lombroso (Child Study Center; Neurobiology), 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; Cellular & Molecular Physiology), 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), Christopher van Dyck (Psychiatry; 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), Hal Blumenfeld (Neurology; Neurobiology), Angélique Bordey (Neurosurgery; Cellular & Molecular Physiology), Charles Bruce (Neurobiology), Michael Crair (Neurobiology), Sabrina Diano (Obstetrics, Gynecology & Reproductive Services; Neurobiology), Karyn Frick (Psychology), Michael Koelle (Molecular Biophysics & Biochemistry), Anthony Koleske (Molecular Biophysics & Biochemistry; Neurobiology), Daeyeol Lee (Neurobiology), Vincent Pieribone (Neurobiology), Maria Mercedes Piñango (Linguistics), Michael Schwartz (Neurobiology), Nenad Sestan (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), David Wells (Molecular, Cellular & Developmental Biology), Anne Williamson (Neurosurgery), Mark Yeckel (Neurobiology), Weimin Zhong (Molecular, Cellular & Developmental Biology)
Assistant Professors Patrick Allen (Psychiatry), Robert Beech (Psychiatry), Thomas Biederer (Molecular Biophysics & Biochemistry), Sreeganga Chandra (Neurology; Molecular, Cellular & Developmental Biology), 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), Angeliki Louvi (Neurosurgery; Neurobiology), James Mazer (Neurobiology), Rory McCrimmon (Internal Medicine), Dhasakumar Navaratnam (Neurology; Neurobiology), Michael Nitabach (Cellular & Molecular Physiology), Christopher Pittenger (Psychiatry), Laurie Santos (Psychology), Samuel Sathyanesan (Psychiatry), Glenn Schafe (Psychology), Dana Small (Psychiatry), Elke Stein (Molecular, Cellular & Developmental Biology), James Swain (Child Study Center), Susumu Tomita (Cellular & Molecular Physiology), 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/NBIO 501a, Principles of Neuroscience Marina Picciotto, Mark Yeckel
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. WF 3:15–4:45
NSCI 502b/MCDB 730b, Cell Biology of the Neuron Elke Stein, Sreeganga Chandra
A comprehensive introduction to neuronal cell biology. Basic principles of cell biology reviewed in the context of the developing nervous system. Membrane trafficking, receptor mechanisms, neurotrophin signaling, neuronal cytoskeleton, axon guidance, and synapse formation and maintenance are discussed. TTh 7–8:15 P.M.
NSCI 504b/MCDB 735b, Seminar in Brain Development and Plasticity Weimin Zhong, Elke Stein
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. MW 2:30–3:45
[NSCI 506b, Introduction to Brain and Behavior]
NBIO 507b/NSCI 507b, Cellular and Molecular Mechanisms of Neurological Disease Dhasakumar Navaratnam, Stephen Strittmatter, Stephen Waxman
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. Web casts of the lectures and Internet-based interactive tutorials are also available. The course extends a twelve-lecture course, Neurobiologic Mechanisms of Disease, offered in the spring of 2009. The course can be taken for credit or audited. Those wishing credit will be graded on a 30-minute Internet-based final exam and a term paper. TTh 4–5
NSCI 510b/NBIO 500b, 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. 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. This course is offered to graduate and M.D./Ph.D. students only and cannot be audited.
[NSCI 514b, Neurodevelopment and Neuropsychiatric Disorders]
NSCI 519a or b, Tutorial
By arrangement with faculty and approval of DGS.
NSCI 521a/PHAR 521a, Neuroimaging in Neuropsychiatry I: Imaging Methods Julie Staley, Kelly Cosgrove, Hilary Blumberg
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. W 9–10:30
NSCI 521b, Neuroimaging in Neuropsychiatry II: Clinical Applications Julie Staley, Kelly Cosgrove
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) 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 application of state-of-the-art neuroimaging methods to research in neurologic and psychiatric disorders. W 9–10:30
NSCI 535b/NBIO 535b, History of Modern Neuroscience Gordon Shepherd
Survey of classical papers that have been the foundation for the rise of modern neuroscience since the 1950s. Areas covered range from genes and proteins through cells and systems to behavior. Classes combine overviews of different areas with discussions of selected classical papers. Emphasis is on how convergence of techniques, concepts, and personalities has been the basis for major advances. Contact course instructor for first class date and time. mW 4–6
NBIO 509b/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 to understand the neural basis of behavior.
[NSCI 540a, Introduction to Statistics]
[NSCI 571b, Neurophysiology]
[NSCI 580a, The MAP Kinase Pathway and Cognitive Disorders]
NSCI 580b, Bioethics in Neuroscience Charles Greer
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. Th 4–5:30
[NSCI 585a, Stem and Progenitor Cells in the Adult Nervous System]
[NSCI 590a, Sensory Neuroethology: Bats, Owls, Electric Fish, and Beyond]
NSCI 595a, Seminar in Visuomotor Neurophysiology Daeyeol Lee, James Mazer
Review and discussion of seminal papers in neurophysiological and computational studies of visual system. It covers papers on the receptive field physiology of neurons in the retina and central visual pathway, motor cortex, and computational theories of vision and motor control. The course largely focuses on the literature in primates, but also draws on behavioral and neurophysiological studies in other mammals, such as cats and humans. Contact course instructor for first class date and time.
[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]
[NSCI 614b, Neurobiology of Learning and Memory]
[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/MCDB 720au/NBIO 720a, Neurobiology Haig Keshishian, Paul Forscher
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. MWF 11:35–12:25
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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