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 involves more than 70 faculty from different campus departments, with expertise ranging from molecular and cellular neuroscience to developmental neuroscience, systems and computational neuroscience, and human cognitive neuroscience.

We provide a highly interdisciplinary, intellectually dynamic training environment of coursework, research training, and mentoring, within a strong research program that produces fundamental advances in knowledge and cutting-edge 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. If the applicant is admitted, then official transcripts of all college-level work will be required. Official transcripts must be in sealed envelopes as issued by the school(s) attended. If you have attended Berkeley, upload your unofficial transcript with your application for the departmental initial review. If you are admitted, an official transcript with evidence of degree conferral will not be required.
  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, not the Graduate Division.
  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.

If applicants have previously been denied admission to Berkeley on the basis of their English language proficiency, they must submit new test scores that meet the current minimum 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. 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.


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

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. Performance in rotations is evaluated and graded. Rotations also allow students to identify the laboratory in which their thesis research will be performed.  During the first-year students also take Neurosci 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.  Neurosci 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. Neurosci 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. 

Step II: Qualifying Exam

Students complete an oral qualifying exam during the spring semester of Year 2. This exam is structured around two written proposals—one in the student’s proposed area of thesis research, and the other in an area of neuroscience outside the thesis topic. During the exam, a faculty committee tests the student’s knowledge of these areas and general neuroscience. Students must demonstrate the ability to recognize important research problems, propose relevant experimental approaches, and display comprehensive knowledge of relevant 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. The students then write a dissertation based on the results of this research. On completion of the research and approval of the dissertation by the committee, the students are awarded the doctorate.

Total Normative Time

Total normative time is 5.7 years.

Time to Advancement


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 231Advanced Developmental and Stem Cell Biology4
MCELLBI 236Advanced Mammalian Physiology5
MCELLBI 240Advanced Genetic Analysis4
NEUROSC C261Cellular and Developmental Neurobiology3
2. Systems and Computational Neuroscience 
Choose one:
PSYCH 210BProseminar: Cognition, Brain, and Behavior3
VIS SCI 265Neural Computation3
NEUROSC C262Circuit and Systems Neurobiology3
3. Cognition, Brain and  Behavior 
Choose one:
PSYCH 117Human Neuropsychology3
PSYCH C127Cognitive Neuroscience3
PSYCH 210AProseminar: Cognition, Brain, and Behavior3
PSYCH 210DCourse Not Available3
PSYCH 214Functional MRI Methods3
PSYCH 240AProseminar: Biological, Cognitive, and Language Development3
PB HLTH C217DBiological and Public Health Aspects of Alzheimer's Disease3
VIS SCI 262Visual Cognitive Neuroscience3
One Graduate Elective Seminar 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 Seminar 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 290PCourse Not Available2
PSYCH 290QSeminars: Cognition2
PSYCH 290ZSeminars1-3
VIS SCI 298Group Studies, Seminars, or Group Research1-6
PSYCH 102Methods for Research in Psychological Sciences3
PSYCH 205Data Analysis3
STAT 150Stochastic Processes3
STAT 151ALinear Modelling: Theory and Applications4
STAT 153Introduction to Time Series4
STAT 204Probability for Applications4
STAT C241AStatistical Learning Theory3
STAT C241BAdvanced Topics in Learning and Decision Making3
STAT 248Analysis of Time Series4
MATH 118Fourier Analysis, Wavelets, and Signal Processing4
MATH 220Introduction to Probabilistic Methods in Mathematics and the Sciences4
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 225ACourse Not Available3
EL ENG 225BCourse Not Available3
EL ENG 226ARandom Processes in Systems4
EL ENG 229AInformation Theory and Coding3
BIO ENG C218Stem Cells and Directed Organogenesis3
BIO ENG C219Protein Engineering3
BIO ENG 231Introduction to Computational Molecular and Cellular Biology4
BIO ENG 243Computational Methods in Biology4
BIO ENG 263Principles of Molecular and Cellular Biophotonics4
BIO ENG C265Principles of Magnetic Resonance Imaging4
Vision Science
VIS SCI 212ECourse Not Available2
VIS SCI 212FCourse Not Available2
VIS SCI 212GCourse Not Available2

Other Degree Requirements 

Pedagogy course
NEUROSC 290Neuroscience First Year Research2
NEUROSC 294Neuroscience Graduate Student Presentation Seminar1
Students must also complete a 1-semester course in Applied Statistics in Neuroscience, or an equivalent approved course in statistics or quantitative analysis methods.

Time in Candidacy

Dissertation Presentation/Formal Exit Seminar

There is no formal defense of the completed dissertation. Neuroscience students are required to publicly present a thesis seminar about their dissertation research in their final year.

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), 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

+ Indicates this faculty member is the recipient of the Distinguished Teaching Award.


Hillel Adesnik, Assistant Professor. Neural basis of perception, neuroscience, neurobiology .
Research Profile

Martin S. Banks, Professor. Stereopsis, virtual reality, optometry, multisensory interactions, self-motion perception, vision, depth perception, displays, picture perception, visual ergonomics.
Research Profile

Helen Bateup, Assistant Professor. Molecular and cellular neuroscience, neurodevelopmental disorders, autism, epilepsy.
Research Profile

Diana Bautista, Associate Professor. Ion channels, sensory physiology, chemosensation, touch, thermosensation, somatosensory system.
Research Profile

George Bentley, Professor. Hormones and behavior, neuroendocrinology of reproduction.
Research Profile

Eric Betzig, Professor. Biophysics.

Sonia Bishop, Assistant Professor. Neural mechanisms supporting attention, emotion and their interactions; individual differences in cognitive control and emotional responsivity; neural substrate of anxiety; genetic factors modulating recruitment of cortical control and limbic affective mechanisms.
Research Profile

Kristofer Bouchard, Adjunct Professor. Biology, Engineering.

Steve Brohawn, Assistant Professor. Neurobiology.
Research Profile

Silvia Bunge, Professor. Cognition, human brain function, development.
Research Profile

Jose M. Carmena, Professor. Brain-machine interfaces, neural ensemble computation, neuroprosthetics, sensorimotor learning and control.
Research Profile

Christopher J. Chang, Professor. Chemistry, inorganic chemistry, neuroscience, bioinorganic chemistry, general physiology, organic chemistry, new chemical tools for biological imaging and proteomics, new metal complexes for energy catalysis and green chemistry, chemical biology.
Research Profile

Anne Collins, Assistant Professor. Human learning, decision-making and executive functions, Computational modeling at multiple levels (cognitive and neuroscience), Behavioral, EEG, drug and genes studies in healthy or patient populations Human learning, decision-making and executive functions, Computational modeling at multiple levels (cognitive and neuroscience), Behavioral, EEG, drug and genes studies in healthy or patient populations .
Research Profile

Emily Cooper, Assistant Professor. 3D vision, depth perception, computational neuroscience, augmented and virtual reality, perceptual graphics.
Research Profile

Mark T. D'Esposito, Professor. Cognitive neuroscience, psychology, working memory, frontal lobe function, functional MRI, neurology, brain imaging, dopamine.
Research Profile

Yang Dan, Professor. Neuronal circuits, mammalian visual system, electrophysiological, psychophysical and computational techniques, visual cortical circuits, visual neurons.
Research Profile

Michael Deweese, Assistant Professor. Machine learning, computation, systems neuroscience, auditory cortex, neural coding.
Research Profile

Andrew Dillin, Professor. Endocrinology, genetics of aging.

David Feinberg, Adjunct Professor. MRI technology development, mapping columnar and visual circuitry, modeling neurovascular coupling.
Research Profile

+ Dan Feldman, Associate Professor. Neurobiology, learning, neurophysiology, sensory biology.
Research Profile

+ Marla B. Feller, Professor. Neurophysiology, developmental neuroscience.
Research Profile

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

John Gerard Flannery, Professor. Neurobiology, optometry, vision science, cell and molecular biology of the retina in normal and diseased states.
Research Profile

David Foster, Professor. Behavioral neurophysiology of spatial learning and memory, hippocampus replay, computational models of reinforcement learning and navigationf.

Jack L. Gallant, Professor. Vision science, form vision, attention, fMRI, computational neuroscience, natural scene perception, brain encoding, brain decoding.
Research Profile

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

Ming Hsu, Assistant Professor. Cognitive neuroscience, experimental economics, behavioral economics, neuroeconomics.
Research Profile

Ehud Y. Isacoff, Professor. Ion channel function, synaptic plasticity, neural excitability, synaptic transmission, the synapse.
Research Profile

Rich Ivry, Professor. Cognitive neuroscience, behavior, cognition, brain, attention, coordination, psychology, motor and perceptual processes in normal and neurologically impaired populations, temporal processing, executive control.
Research Profile

Lucia F. Jacobs, Professor. Cognitive and brain evolution, adaptive patterns in spatial memory, spatial navigation, cognitive sex differences and decision making.
Research Profile

William J. Jagust, Professor. Neuroscience, cognition, brain aging, dementia, imaging, Alzheimerandamp,#039,s disease.
Research Profile

Na Ji, Associate Professor. Biophysics.
Research Profile

Daniela Kaufer, Professor. Neuroscience, stress, neural stem cells, epilepsy, traumatic brain injury, blood brain barrier, prosocial behavior.
Research Profile

Stanley A. Klein, Professor. Optometry, vision science, spatial vision modeling, psychophysical methods and vision test design, corneal topography and contact lens design, source localization of evoked potentials, fMRI, amblyopia.
Research Profile

Robert Thomas Knight, Professor. Cognitive neuroscience, language, physiology, memory, attention, psychology, working memory, neuropsychology, human prefrontal cortex, neural mechanisms of cognitive processing, sensory gating, sustained attention, ad novelty detection.
Research Profile

Richard H. Kramer, Professor. Cells, synaptic transmission, chemical signaling between neurons, ion channels, electrical signals, chemical reagents, synapses.
Research Profile

+ Lance Kriegsfeld, Associate Professor. NeuroendocrinologyCircadian Biology, Neuroimmunology, cancer biology, animal behavior.
Research Profile

Stephan Lammel, Assistant Professor. Neuroscience, Optogenetics, dopamine, addiction, depression.
Research Profile

Markita Landry, Assistant Professor. Chemistry.
Research Profile

Dennis M. Levi, Professor. Optometry, vision science, pattern vision, abnormal visual development.
Research Profile

Lexin Li, Professor. Neuroimaging data analysis, networks data analysis, personalized recommendation, statistical genetics, computational biology, dimension reduction, variable selection, high dimensional regressions, statistical machine learning, data mining, computational statistics.
Research Profile

Chunlei Liu, Professor. Brain imaging, magnetic resonance imaging, remote neural modulation, biosystems and computational biology, signal processing.

Ellen Lumpkin, Professor. Cell and Developmental Biology, research focus: to elucidate force transduction mechanisms that initiate the senses of touch and pain.
Research Profile

Michel Maharbiz, Professor. Electrical Engineering and Computer Sciences. Research Focus: Building micro- and nano- scale machine interfaces to cells and organisms, including development of neural dust technology.
Research Profile

Evan W. Miller, Assistant Professor. Chemical biology, organic chemistry, fluorescence microscopy, neuroscience, imaging .

Bruno Olshausen, Professor. Visual perception, computational neuroscience, computational vision.
Research Profile

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

Teresa Puthussery, Assistant Professor. Retinal Neurobiology and Neurophysiology.
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. Molecular Cell Biology.

David Schaffer, Professor. Neuroscience, biomolecular engineering, bioengineering, stem cell biology, gene therapy.
Research Profile

Kristin Scott, Professor. Nerve cell connectivity in developing nervous systems, taste perception in the fruit fly, taste neural circuits, sensory maps in the brain.
Research Profile

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

Michael Silver, Associate Professor. Cognitive neuroscience, pharmacology, learning, attention, visual perception, neuroimaging.
Research Profile

Fritz Sommer, Adjunct Professor. Bayesian methods, information theory, memory, sensory processing, visual system.
Research Profile

Mark A. Tanouye, Professor. Genetics, neuroanatomy, electrophysiology, mechanisms of nervous system structure and function, drosophila mutants.
Research Profile

W. Rowland Taylor, Professor. Retinal circuit function, neural architecture, immunohisochemical studies.
Research Profile

Frederic Theunissen, Professor. Behavior, cognition, brain, psychology, birdsong, vocal learning, audition, neurophysiology, speech perception, computational neuroscience, theoretical neuroscience.
Research Profile

Doris Tsao, Professor. Neurobiology, Howard Hughes Medical Institute Investigator. The Tsao lab seeks to understand how the brain builds a model of the visual world.
Research Profile

Matthew P. Walker, Professor. Plasticity, learning, memory, fMRI, emotion, sleep, EEG.
Research Profile

Jonathan David Wallis, Professor. Prefrontal cortex, neurophysiology, executive control, decision making.
Research Profile

Kevin Weiner, Assistant Professor. Psychology.
Research Profile

David Whitney, Professor. Cognitive neuroscience, cognition, attention, visual perception, vision, visually guided action.
Research Profile

Linda Wilbrecht, Assistant Professor. Neuroscience, addiction, early life adversity, adolescence.
Research Profile

Ke Xu, Assistant Professor. Biophysical chemistry, cell biology at the nanoscale, super-resolution microscopy, single-molecule spectroscopy.
Research Profile

Michael Yartsev, Assistant Professor. Neuroscience, engineering.
Research Profile

Contact Information

Neuroscience Graduate Group

444 Li Ka Shing Center

Phone: 510-642-8915

Fax: 510-642-4966

Visit Group Website

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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