University of California, Berkeley

About the Program

The Neuroscience Graduate Program at UC Berkeley is a unique, diverse PhD training program that offers intensive, integrated training in multiple areas of neuroscience research.

The program includes 64 faculty from different campus departments, with expertise ranging from molecular and cellular neuroscience to systems and computational neuroscience to human cognitive neuroscience.

We provide a highly interdisciplinary, intellectually dynamic training environment of coursework, research training, professional development, and mentoring, within a strong research program that produces fundamental advances in knowledge and novel techniques.

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Admission to the University

Minimum Requirements for Admission

The following minimum requirements apply to all graduate programs and will be verified by the Graduate Division:

  1. A bachelor’s degree or recognized equivalent from an accredited institution;
  2. A grade point average of B or better (3.0);
  3. If the applicant has completed a basic degree from a country or political entity (e.g., Quebec) where English is not the official language, adequate proficiency in English to do graduate work, as evidenced by a TOEFL score of at least 90 on the iBT test, 570 on the paper-and-pencil test, or an IELTS Band score of at least 7 on a 9-point scale (note that individual programs may set higher levels for any of these); and
  4. Sufficient undergraduate training to do graduate work in the given field.

Applicants Who Already Hold a Graduate Degree

The Graduate Council views academic degrees not as vocational training certificates, but as evidence of broad training in research methods, independent study, and articulation of learning. Therefore, applicants who already have academic graduate degrees should be able to pursue new subject matter at an advanced level without the need to enroll in a related or similar graduate program.

Programs may consider students for an additional academic master’s or professional master’s degree only if the additional degree is in a distinctly different field.

Applicants admitted to a doctoral program that requires a master’s degree to be earned at Berkeley as a prerequisite (even though the applicant already has a master’s degree from another institution in the same or a closely allied field of study) will be permitted to undertake the second master’s degree, despite the overlap in field.

The Graduate Division will admit students for a second doctoral degree only if they meet the following guidelines:

  1. Applicants with doctoral degrees may be admitted for an additional doctoral degree only if that degree program is in a general area of knowledge distinctly different from the field in which they earned their original degree. For example, a physics PhD could be admitted to a doctoral degree program in music or history; however, a student with a doctoral degree in mathematics would not be permitted to add a PhD in statistics.
  2. Applicants who hold the PhD degree may be admitted to a professional doctorate or professional master’s degree program if there is no duplication of training involved.

Applicants may apply only to one single degree program or one concurrent degree program per admission cycle.

Required Documents for Applications

  1. Transcripts: Applicants may upload unofficial transcripts with your application for the departmental initial review. Unofficial transcripts must contain specific information including the name of the applicant, name of the school, all courses, grades, units, & degree conferral (if applicable). 
  2. Letters of recommendation: Applicants may request online letters of recommendation through the online application system. Hard copies of recommendation letters must be sent directly to the program, by the recommender, not the Graduate Admissions.
  3. Evidence of English language proficiency: All applicants who have completed a basic degree from a country or political entity in which the official language is not English are required to submit official evidence of English language proficiency. This applies to institutions from Bangladesh, Burma, Nepal, India, Pakistan, Latin America, the Middle East, the People’s Republic of China, Taiwan, Japan, Korea, Southeast Asia, most European countries, and Quebec (Canada). However, applicants who, at the time of application, have already completed at least one year of full-time academic course work with grades of B or better at a US university may submit an official transcript from the US university to fulfill this requirement. The following courses will not fulfill this requirement:

    • courses in English as a Second Language,

    • courses conducted in a language other than English,

    • courses that will be completed after the application is submitted, and

    • courses of a non-academic nature.

Applicants who have previously applied to Berkeley must also submit new test scores that meet the current minimum requirement from one of the standardized tests. Official TOEFL score reports must be sent directly from Educational Test Services (ETS). The institution code for Berkeley is 4833 for Graduate Organizations. Official IELTS score reports must be sent electronically from the testing center to University of California, Berkeley, Graduate Division, Sproul Hall, Rm 318 MC 5900, Berkeley, CA 94720. TOEFL and IELTS score reports are only valid for two years prior to beginning the graduate program at UC Berkeley. Note: score reports can not expire before the month of June.


Where to Apply

Visit the Berkeley Graduate Division application page

Admission to the Program

Applicants to the program should have a bachelor's degree from a four-year college and at least one year of laboratory experience. The Graduate Record Examination (GRE) General Test is optional. For more information on our program requirements go to:

Doctoral Degree Requirements

Normative Time Requirements

Normative Time to Advancement

Step I: Lab Rotations and Presentations, First Year Classes

During the first year of graduate study, each neuroscience graduate student spends three 10-week periods performing research projects in different faculty laboratories. The goal is to expose students to different techniques and approaches in neuroscience and to provide training in experimental design, critical analysis of data, and presentation of research findings. Rotations also allow students to identify the laboratory in which their thesis research will be performed. Rotation research is graded and receives academic credit. This is accomplished by enrolling in NEUROSC 291A/B, a year-long course, during the rotation year. Also during the first-year students take NEUROSC 290A/B Methods & Career Skills Classes which introduce a broad range of modern neuroscience research methods in didactic lectures and provide advising in initial career skills.  NEUROSC 290A (Fall) includes a survey of cutting-edge research methods, advising on how to choose a thesis mentor, training in scientific rigor and reproducibility, and an introduction to the use and misuse of statistics in neuroscience research. NEUROSC 290B (Spring) includes in-depth training on how to give a top-notch scientific talk, advising on how to write effective research papers, and on scientific project management. Finally, student also enroll in MCELLBI 293C during the spring of their first year to ensure that research trainees receive ample training in Responsible Conduct in Research, and to gain an understanding of federal, state, and UC Berkeley policies and resources available to further support their research endeavors.

Step II: Qualifying Exam

Students complete an oral qualifying exam during the spring semester of Year 2. The examination has three parts: Thesis Proposal, Related Research Areas, and Foundational Questions in Neuroscience. The thesis proposal is in the form of a written, NIH-style grant proposal, which is turned in to the committee, and then defended orally. Related Research Areas are identified cooperatively by the student and his/her committee prior to the exam, and are chosen to be complementary to the main thesis research subject. These areas are examined orally. The Foundational Questions in Neuroscience are designed to test broad knowledge in Neuroscience. These are a published list of questions, the same for all students, that are available upon entry to the program. These questions are designed to test basic common knowledge of neuroscience facts and principles, and a subset of them are examined orally during the qualifying exam. During the exam, students must demonstrate the ability to recognize fundamentally important research problems, propose relevant experimental approaches, and display comprehensive knowledge of appropriate disciplinary areas and related subjects. Students must pass the qualifying examination before advancing to doctoral candidacy.

Normative Time in Candidacy

Step III: Dissertation

Students undertake research for the PhD dissertation under a four-person committee in charge of their research and dissertation. Students do original research using a wide variety of cutting-edge neuroscience methods. During this time, students must meet at least annual with their thesis committee to discuss dissertation progress, review experimental results, set goals, and ensure students are adhering to appropriate timelines to completion. The students then write a dissertation based on the results of their research. 

STEP IV: Dissertation Presentation/Formal Exit Seminar

There is no formal defense of the completed dissertation. However, Neuroscience students are required to publicly present a thesis seminar about their dissertation research in their final year. On completion of the research and approval of the dissertation by the committee, the students are awarded the doctorate.

Total Normative Time

Normative time to advancement is 2 years. Total normative time is 5.7 years.


Pedagogy, Rotations, Ethics, & Seminar Courses 

Student must take all of the following courses. Pedagogy, Rotations, and Ethics courses are taken in year 1. Brain Lunch Seminar is taken in Years 1, 2, and 4.

Pedagogy courses
NEUROSC 290ANeuroscience Research Design and Analysis1
NEUROSC 290BNeuroscience Career Skills1
NEUROSC 291ANeuroscience Introduction to Research4-12
NEUROSC 291BNeuroscience Introduction to Research4-12
Ethics in Research
MCELLBI 293CResponsible Conduct in Research1
Brain Lunch Seminar
NEUROSC 294Neuroscience Graduate Student Presentation Seminar (Brain Lunch)1
All students are required to enroll in the Brain Lunch seminar for 1 semester in each of Years 1 and 2, and again in Year 4 (see "Presentations" under "Required Professional Development" below)
One Graduate Course in Each of the Following Three Foundational Areas

Students can either take one graduate-level course from each category, or three graduate level courses from two areas, plus a selected advanced undergraduate course from a third area. They are taken in years 1–2.  Courses offered will vary depending on the semester.  The courses below are samples of courses that fulfill the area requirements.

1. Cellular, Molecular and Developmental Neuroscience
Choose one:
MCELLBI 160Cellular and Molecular Neurobiology4
MCELLBI 166Biophysical Neurobiology3
MCELLBI 230Advanced Cell and Developmental Biology4
MCELLBI 231Advanced Developmental and Stem Cell Biology4
MCELLBI 240Advanced Genetic Analysis4
NEUROSC/MCELLBI C261Cellular and Developmental Neurobiology3
2. Systems and Computational Neuroscience 
Choose one:
INTEGBI 139The Neurobiology of Stress4
MCELLBI 236Advanced Mammalian Physiology5
NEUROSC/MCELLBI C262Circuit and Systems Neurobiology3
PSYCH 210BProseminar: Cognition, Brain, and Behavior3
VIS SCI 260CIntroduction to Visual Neuroscience3
VIS SCI 265Neural Computation3
3. Cognition, Brain and  Behavior 
Choose one:
PSYCH 117Human Neuropsychology3
PSYCH C127Cognitive Neuroscience3
PSYCH 210AProseminar: Cognition, Brain, and Behavior3
PSYCH 240AProseminar: Biological, Cognitive, and Language Development3
PB HLTH C217DBiological and Public Health Aspects of Alzheimer's Disease3
PB HLTH 290Health Issues Seminars (Neuroepidemiology)1-4
VIS SCI 262Visual Cognitive Neuroscience3
One course on statistical analysis or quantitative methods

Students must complete a 1-semester course in Applied Statistics in Neuroscience, or an equivalent approved course in statistics or quantitative analysis methods. This can be completed at any time prior to the semester of graduation. Students with prior appropriate coursework or whose thesis research uses substantial quantitative methods can use that prior experience to fulfill this requirement, subject to approval by the Head Graduate Adviser.

NEUROSC 299Seminars (Applied Statistics for Neuroscience)1-3
One Graduate Elective Course

Students must take one additional elective course. This can be either a graduate-level seminar or graduate-level lecture course, and can be 1 unit or more. This is typically taken in years three-four. You may also select a foundation course as an elective. Consult your thesis adviser and thesis committee to select the most appropriate course for you.

Neuro-Related Courses
EL ENG 290AAdvanced Topics in Electrical Engineering: Advanced Topics in Computer-Aided Design1-3
EL ENG 290BAdvanced Topics in Electrical Engineering: Advanced Topics in Solid State Devices1-3
EL ENG 290CAdvanced Topics in Electrical Engineering: Advanced Topics in Circuit Design1-3
EL ENG 290DAdvanced Topics in Electrical Engineering: Advanced Topics in Semiconductor Technology1-3
EL ENG 290FAdvanced Topics in Electrical Engineering: Advanced Topics in Photonics1-3
EL ENG 290NAdvanced Topics in Electrical Engineering: Advanced Topics in System Theory1-3
EL ENG 290OAdvanced Topics in Electrical Engineering: Advanced Topics in Control1-3
EL ENG 290PAdvanced Topics in Electrical Engineering: Advanced Topics in Bioelectronics1-3
EL ENG 290QAdvanced Topics in Electrical Engineering: Advanced Topics in Communication Networks1-3
EL ENG 290SAdvanced Topics in Electrical Engineering: Advanced Topics in Communications and Information Theory1-3
EL ENG 290TAdvanced Topics in Electrical Engineering: Advanced Topics in Signal Processing1-3
EL ENG 290YAdvanced Topics in Electrical Engineering: Organic Materials in Electronics3
LINGUIS 290ATopics in Linguistic Theory: Syntax3
LINGUIS 290BTopics in Linguistic Theory: Semantics3
LINGUIS 290DTopics in Linguistic Theory: Pragmatics3
LINGUIS 290ETopics in Linguistic Theory: Phonology3
LINGUIS 290FTopics in Linguistic Theory: Diachronic Linguistics3
LINGUIS 290HTopics in Linguistic Theory: Linguistic Reconstruction3
LINGUIS 290LAdditional Seminar on Special Topics to Be Announced3
LINGUIS 290MTopics in Linguistic Theory: Psycholinguistics3
MCELLBI 290Graduate Seminar1
PSYCH 290BSeminars: Biological2
PSYCH 290ESeminars: Perception2
PSYCH 290HSeminars: Developmental2
PSYCH 290ISeminars: Personality2
PSYCH 290JSeminars: Social2
PSYCH 290KSeminars: Clinical2
PSYCH 290QSeminars: Cognition2
PSYCH 290ZSeminars1-3
VIS SCI 298Group Studies, Seminars, or Group Research1-6
NEUROSC 299Seminars1-3
PSYCH 102Methods for Research in Psychological Sciences3
PSYCH 111Human Neuroanatomy3
PSYCH 115Introduction to Brain Imaging Analysis Methods3
PSYCH 125The Developing Brain3
PSYCH 205Data Analysis3
PSYCH 208Methods in Computational Modeling for Cognitive Science3
STAT 150Stochastic Processes3
STAT 151ALinear Modelling: Theory and Applications4
STAT 153Introduction to Time Series4
STAT 158Experimental Design4
STAT C241AStatistical Learning Theory3
STAT C241BAdvanced Topics in Learning and Decision Making3
STAT 248Analysis of Time Series4
MATH 118Fourier Analysis, Wavelets, and Signal Processing4
Computer Science and Programming
COMPSCI C280Computer Vision3
Electrical Engineering
EL ENG 120Signals and Systems4
EL ENG 123Digital Signal Processing4
EL ENG 126Probability and Random Processes4
EL ENG 221ALinear System Theory4
EL ENG 226ARandom Processes in Systems4
EL ENG C227CConvex Optimization and Approximation3
EL ENG 229AInformation Theory and Coding3
BIO ENG 231Introduction to Computational Molecular and Cellular Biology4
BIO ENG C265/EL ENG C225EPrinciples of Magnetic Resonance Imaging4
Vision Science
VIS SCI 260AOptical and Neural Limits to Vision3
VIS SCI 260DSeeing in Time, Space and Color3
Public Health
PB HLTH 245Introduction to Multivariate Statistics4

Required Professional Development


During their fourth year of study, students are required to make a presentation on the progress of their thesis work while enrolling in NEUROSC 294 (Neuroscience Graduate Student Presentation Seminar, also known as "Brain Lunch"), a journal club, for a letter grade.


Neuroscience students are required to serve as graduate student instructors (GSIs) for two semesters. Whenever possible, GSI assignments are determined with an eye toward student research interests. Teaching occurs during fall semester of the second year and spring semester of the third. Teaching affords students supervised experience in a variety of educational situations, including labs, discussion sections, and demonstrations. GSIs also participate in record-keeping, grading, advising, and student consultations.

GSIs are evaluated by both supervising faculty and the students they teach. These evaluations become a permanent part of the student file. Deserving GSIs are nominated for the Outstanding Graduate Student Instructor Award.



Faculty and Instructors


Hillel Adesnik, Associate Professor. Neural basis of sensation, perception, and action.
Research Profile

Helen Bateup, Associate Professor. Molecular basis of synapse and circuit changes associated with epilepsy and autism.
Research Profile

Diana Bautista, Professor. Molecular mechanisms underlying the sensations of itch, touch, and pain.
Research Profile

Eric Betzig, Professor. Development of new tools for imaging and image analysis to enable biological discovery.
Research Profile

Sonia Bishop, Associate Professor. Neural basis of attention, emotion, and anxiety. Individual differences in cognitive control and emotional responsivity.
Research Profile

Kristofer Bouchard, Adjunct Assistant Professor. Functional organization and dynamic coordination of sensorimotor networks underlying learned, skilled behaviors.
Research Profile

Steve Brohawn, Assistant Professor. Molecular basis of sensory transduction and electrical signaling, especially mechanosensation.
Research Profile

Silvia Bunge, Professor. Neural mechanisms, development, and plasticity of higher cognitive functions in humans.
Research Profile

Jose M. Carmena, Adjunct Professor. Neural basis of motor skill learning. Application to neural prostheses and development of neural dust technology.
Research Profile

Christopher J. Chang, Professor. Chemical tools for imaging and optogenetics in neurobiology.
Research Profile

Anne Collins, Associate Professor. Computational modeling of human learning, decision-making, and executive functions.
Research Profile

Emily Cooper, Assistant Professor. Computational modeling of visual perception.
Research Profile

Mark T. D'Esposito, Professor. Neural basis of high-level cognitive processes such as working memory and executive control.
Research Profile

Yang Dan, Professor. Neural circuits controlling sleep; mechanisms of executive control. .
Research Profile

Michael Deweese, Associate Professor. Neural mechanisms underlying auditory processing and selective attention in the cerebral cortex and artificial neural networks.
Research Profile

Andrew Dillin, Professor. Genetic and molecular mechanisms regulating aging and aging-related disease.
Research Profile

* Dan Feldman, Professor. Sensory processing and plasticity in the somatosensory cortex.
Research Profile

* Marla B. Feller, Professor. Functional development and organization of neural circuits in the retina.
Research Profile

Yvette Fisher, Assistant Professor. Flexibility of neural circuits for spatial navigation.
Research Profile

John Gerard Flannery, Professor. Gene therapies for inherited retinal degenerations.
Research Profile

David Foster, Associate Professor. Encoding of spatial memory and navigation toward rewards by neural ensembles in the hippocampus.
Research Profile

Jack L. Gallant, Professor. Identifying cortical maps to discover how the brain represents information about the world and its own mental states.
Research Profile

Andrea Gomez, Assistant Professor. Instructive cues for neural form and function.
Research Profile

Ming Hsu, Associate Professor. Neural basis of economic and consumer decision-making.
Research Profile

Ehud Y. Isacoff, Professor. Mechanisms of ion channel function, synapse development, plasticity, and neural circuit function.
Research Profile

Richard Ivry, Professor. Cognition and action, with an emphasis on how people select actions, learn skills, and produce coordinated .
Research Profile

William J. Jagust, Professor. Anatomic, biochemical, and neurochemical bases of brain aging and dementia.
Research Profile

Na Ji, Associate Professor. Novel imaging methods to understand the brain.
Research Profile

Daniela Kaufer, Acting Associate Dean and Professor. Molecular mechanisms of brain plasticity in response to stress and neurological insults.
Research Profile

Robert Thomas Knight, Professor. Novel chemical reagents for non-invasive optical sensing and manipulation of ion channels and synapses.
Research Profile

Richard H. Kramer, Professor. Novel chemical reagents for non-invasive optical sensing and manipulation of ion channels and synapses.
Research Profile

+ Lance Kriegsfeld, Professor . Brain and endocrine regulation of circadian rhythms.
Research Profile

Stephan Lammel, Associate Professor. Midbrain dopamine circuits in reward-based behaviors and pathological changes in addiction, depression and schizophrenia.
Research Profile

Markita Landry, Assistant Professor. Exploiting nanomaterials to probe and characterize complex biological systems at the nano-scale; nanosensors for brain chemistry.
Research Profile

Lexin Li, Professor. Statistical neuroimaging analysis, brain connectivity analysis, imaging causal inference, multimodal and longitudinal imaging analysis, and imaging tensor regression.
Research Profile

Chunlei Liu, Associate Professor. MRI technology development for the study of neural circuits and modulation.
Research Profile

Ellen Lumpkin, Professor. To elucidate force transduction mechanisms that initiate the senses of touch and pain.
Research Profile

Michel Maharbiz, Adjunct Professor. Building micro- and nano- scale machine interfaces to cells and organisms, including development of neural dust technology.
Research Profile

Evan W. Miller, Associate Professor. Development and application of molecular tools for studying neuroscience.
Research Profile

Bruno Olshausen, Professor. Computational models of sensory coding and visual perception.
Research Profile

Steven Piantadosi, Assistant Professor. Understanding what computational processes support language acquisition, math learning, and general cognition.
Research Profile

Teresa Puthussery, Assistant Professor. Processing of visual signals in the healthy and diseased retina.
Research Profile

Michael Rape, Professor. Molecular Mechanisms of Cell Fate Decisions in Development and Disease.
Research Profile

Austin Roorda, Professor. Vision Science and Optometry, development of adaptive optics to track, measure, and correct the eye's imperfection.
Research Profile

Kaoru Saijo, Assistant Professor. Microglial cell maintenance of homeostasis in the brain and sex dimorphism in diseases.
Research Profile

David Schaffer, Professor. Engineering stem cell and gene therapeutics.
Research Profile

Randy Schekman, Professor. Membrane assembly, vesicular transport, and membrane fusion among organelles of the secretory pathway.
Research Profile

Kristin Scott, Professor. Taste detection, processing, and behaviors.
Research Profile

Karthik Shekhar, Assistant Professor. Single-cell genomics and statistical inference.
Research Profile

Michael Silver, Professor. Neurophysiological and neurochemical substrates of human visual perception, attention, and learning.
Research Profile

Friedrich Sommer, Adjunct Professor. Theoretical principles of learning and perception.
Research Profile

Rowland Taylor, Professor. Structure and function of neural circuits in the retina.
Research Profile

Frederic Theunissen, Professor. Perception of complex sounds.
Research Profile

Doris Tsao, Professor. The Tsao lab seeks to understand how the brain builds a model of the visual world.
Research Profile

Matthew P. Walker, Professor. The impact of sleep on human health and disease.
Research Profile

Joni Wallis, Professor. Neuronal mechanisms underlying high-level cognitive and behavioral processes.
Research Profile

Kevin Weiner, Associate Professor. Models to explain how brain structure and function contribute to measurable behaviors (e.g. face perception).
Research Profile

David Whitney, Professor. Visual perception and attention.
Research Profile

Linda Wilbrecht, Associate Professor. Experience dependent plasticity and the development of circuits involved in value based decision making.
Research Profile

Ke Xu, Associate Professor. Super-resolution fluorescence microscopy to interrogate cellular processes at the nanoscale.
Research Profile

Michael Yartsev, Associate Professor. Neural basis of complex spatial and acoustic behaviors.
Research Profile

Emeritus Faculty

Martin S. Banks, Professor Emeritus. Visual space perception and sensory combination.
Research Profile

John Clarke, Professor of the Graduate School of Physics. Superconducting Quantum Interference Devices such as ultralow-frequency MRI.
Research Profile

Howard Fields, Adjunct Emeritus Professor. Mesolimbic circuits involved in goal directed behaviors, opioid regulation of synaptic function.
Research Profile

Ralph D. Freeman, Professor Emeritus. Central Visual Pathways: Systems and Computational Neuroscience.
Research Profile

John Ngai, Professor Emeritus. Understanding the molecular and cellular mechanisms underlying the function, development and regeneration of the vertebrate olfactory system.
Research Profile

Geoff Owen, Professor Emeritus. Comprehensive theory of retinal image processing.
Research Profile

Mu-Ming Poo, Professor Emeritus. Formation and plasticity of synapses, and activity-dependent modification of neural circuits.
Research Profile

Gerald Westheimer, Professor of the Graduate School Division of Neurobiology. Perceptual learning in spatial visual tasks.
Research Profile

Bob Zucker, Professor of the Graduate School Division of Neurobiology. Mechanisms underlying regulation of synaptic transmission.
Research Profile

Contact Information

Neuroscience Graduate Group

444 Li Ka Shing Center

Phone: 510-642-8915

Fax: 510-642-4966

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Program Chair/Head Graduate Advisor

Michael Silver, PhD

Phone: 510-642-3130

Graduate Program Manager

Leleña Avila

444 Li Ka Shing Center

Phone: 510-642-8915

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