University of California, Berkeley

About the Program

The Biophysics Graduate Group is an interdisciplinary PhD program hosted by the California Institute for the Biosciences (QB3). Our program trains graduate students for careers at the interface of the biological and physical sciences. This interdisciplinary group provides an opportunity for interested students to receive training leading to the PhD in Biophysics. Approximately 60 faculty members are affiliated with the Biophysics Group, spanning over a dozen departments and groups at UC Berkeley. Students may work under the supervision of any faculty member belonging to the group.

Students interested in pursuing graduate work in biophysics typically acquire undergraduate training in one of the basic physical or biological sciences and during the first two years at UC Berkeley take self-selected courses in topics such as biology, physics, and chemistry to fill in any gaps in foundational knowledge.

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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 comes 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 from countries or political entities in which the official language is not English are required to submit official evidence of English language proficiency. This applies to applicants 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 mailed directly to our office from the British Council. 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 should have sufficient undergraduate training to undertake graduate work in the chosen field. This includes such undergraduate majors as: biological sciences, structural biology, physics, math, and/or chemistry. Laboratory experience is expected.

Typical students admitted to the program have demonstrated outstanding potential as a research scientist and have clear academic aptitude in multiple disciplines, as well as excellent communication skills. This is assessed based on research experience, grades, standardized exams, course selection, essays, personal background, and letters of recommendation.

Average GRE scores considered for admissions are in the 80th percentile (quantitative). We do not have set requirements for GRE scores, but generally admit students in the 80th and 90th percentile. GRE subject tests are not required, but will be reviewed if submitted with the application.

Doctoral Degree Requirements

Normative Time Requirements

Normative Time to Advancement

Normative time to advancement is two years.

Year 1

Students perform three laboratory rotations with the chief aim of identifying a research area and selecting a thesis laboratory. Students also take courses and should complete two graded courses, of their choice, in the first year.

Year 2

Students attend seminars, prepare a dissertation prospectus, and prepare for their PhD oral qualifying examination (QE). With the successful passing of the QE, students select their thesis committee and advance to candidacy for the PhD degree.

Normative Time in Candidacy

Years 3 to 5

Students undertake research for the PhD dissertation under a self-selected four-person committee in charge of their research and dissertation. Students conduct original laboratory research, and then write the 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 5-5.5/6 years.

Time to Advancement

Courses Required
BIOPHY 293AResearch Seminar: Faculty Evening Research Presentations (FERPS) and Student Evening Research Presentations (SERPS)2
BIOPHY 293BResearch Seminar: Faculty Evening Research Presentations (FERPS) and Student Evening Research Presentations (SERPS)2
Electives, as per approved study list: four semesters
MCELLBI 293CResponsible Conduct in Research1
Lab Rotations

Students conduct three 10-week laboratory rotations in the first year. The thesis lab, where dissertation research will take place, is chosen at the end of the third rotation in late April/early May.

Qualifying Examination

The qualifying examination will evaluate a student’s depth of knowledge in his or her research area, breadth of knowledge in fundamentals of biophysics, ability to formulate a research plan, and critical thinking. The QE prospectus/proposal will include a description of the specific research problem, but will serve as a framework for the QE committee members to probe the student’s foundational knowledge in the field and area of research. Proposals will be written in the manner of a NIH-style grant proposal. The prospectus must be completed and submitted to the QE chair no fewer than four weeks prior to the oral qualifying examination.

Students are expected to pass the qualifying examination by the end of the fourth semester in the program.

Time in Candidacy


After passing the qualifying exam by the end of the second year, students have until the fifth semester to select a thesis committee and submit the online advancement to candidacy form to the Graduate Division.


Primary dissertation research is conducted in years 3-5/5.5. Requirements for the dissertation are decided in consultation with the thesis adviser and thesis committee members. To this end, students are required to have yearly thesis committee meetings with the committee after advancing to candidacy.

Dissertation Presentation/Finishing Talk

There is no formal defense of the completed dissertation; however, students are expected to publicly present a talk about their dissertation research in their final year.

Required Professional Development


All biophysics students are expected to attend the annual retreat and present research talks there. They are also encouraged to attend national and international conferences to present research from the second year onwards.


Biophysics students are required to teach one semester and may teach more. The teaching requirement may be waived if the student creates and teaches a Biophysics module (student-run five-week workshops).



Faculty and Instructors

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


Hillel Adesnik, Assistant Professor.
Research Profile

Adam Arkin, Professor. Systems modeling.
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Eric Betzig, Professor. Physics, molecular and cell biology.
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Steven Brenner, Professor. Computational Biology, Genomics, Protein Function Prediction.
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Steve Brohawn, Assistant Professor. Molecular and cell biology.
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Carlos J. Bustamante, Professor. Nanoscience, structural characterization of nucleo-protein assemblies, single molecule fluorescence microscopy, DNA-binding molecular motors, the scanning force microscope, prokaryotes.
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Jamie Cate, Professor. Molecular basis for protein synthesis by the ribosome, RNA, antibiotics, a thermophilic bacterium, escherichia coli.
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Kathy Collins, Professor. Telomerase, non-coding RNAs, RNA silencing.
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Xavier Darzacq, Assistant Professor.
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Karen Davies, Assistant Adjunct Professor. Protein structure, bioenergentic membranes, cellualr organisation, electron cryo tomography.
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Abby Dernburg, Professor. Genomics, chromosome remodeling and reorganization during meiosis, Down syndrome, DNA.
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Michael Deweese, Associate Professor. Machine learning, computation, systems neuroscience, auditory cortex, neural coding.
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Jennifer A. Doudna, Professor. RNA machines, hepatitis C virus, RNA interference, ribosomes.
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David G. Drubin, Professor. Cellular morphogenesis, plasma membrane dynamics, microtubule cytoskeletons, cytoskeletal proteins, morphological development.
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Michael B. Eisen, Professor. Genomics, genome sequencing, bioinformatics, animal development.
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+ Dan Feldman, Professor. Neurobiology, learning, neurophysiology, sensory biology.
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Marla B. Feller, Professor. Neurophysiology, developmental neuroscience.
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Graham R. Fleming, Professor. Chemistry, proteins, chemical and biological dynamics in the condensed phase, ultrafast spectroscopy, body dynamics, liquids, solutions, glasses, photosynthetic proteins, role of solvents in chemical reactions, complex electric fields, electron transfer.
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Daniel Fletcher, Professor. Bioengineering, optical and force microscopy, microfabrication, biophysics, mechanical properties of cells.
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+ Robert J. Full, Professor. Energetics, comparative biomechanics, arthropod, adhesion, comparative physiology, locomotion, neuromechanics, biomimicry, biological inspiration, reptile, gecko, amphibian, robots, artificial muscles.
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Jack L. Gallant, Professor. Vision science, form vision, attention, fMRI, computational neuroscience, natural scene perception, brain encoding, brain decoding.
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Hernan G. Garcia, Assistant Professor.
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+ Phillip Lewis Geissler, Professor. Statistical mechanics, theoretical chemistry, microscopic behavior of complex biological and material systems, biomolecular structure and dynamics, nonlinear vibrational spectroscopy.
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Jay T. Groves, Professor. Chemistry, physical chemistry of cell membranes, molecular organization in cell membranes, receptor-ligand binding, spatial rearrangement of receptors, ligands.
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Oskar Hallatschek, Assistant Professor.
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Teresa Head-Gordon, Professor. Computational chemistry, biophysics, bioengineering, biomolecules, materials, computational science.
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James Hurley, Professor. Structural biology and biophysics of membrane remodeling in autophagy, membrane traffic, and HIV biogenesis.
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Nicholas Ingolia, Assistant Professor. Ribosome Profiling, translation, genomics.
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Ehud Y. Isacoff, Professor. Ion channel function, synaptic plasticity, neural excitability, synaptic transmission, the synapse.
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Na Ji, Associate Professor. Physics, molecular and cell biology.

Sung-Hou Kim, Professor. Computational genomics, Structural Biology, drug discovery, disease genomics.
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Mimi A. R. Koehl, Professor. Biomechanics, insects, invertebrate functional morphology, fluid dynamics, aerodynamics, marine animals, filtration, gliding vertebrates.
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Richard H. Kramer, Professor. Cells, synaptic transmission, chemical signaling between neurons, ion channels, electrical signals, chemical reagents, synapses.
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Sanjay Kumar, Professor. Biomaterials, molecular and cellular bioengineering, stem cells, cancer biology, translational medicine.
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John Kuriyan, Professor. Structural and functional studies of signal transduction, DNA replication, cancer therapies, phosphorylation.
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Markita Landry, Assistant Professor.
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Luke Lee, Professor. Biophotonics, biophysics, bionanoscience, molecular imaging, single cell analysis, bio-nano interfaces, integrated microfluidic devices (iMD) for diagnostics and preventive personalized medicine.
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Polina Lishko, Assistant Professor. Reproductive and Developmental Biology, ion channels, Physiology of Fertilization and Early Embryo Development.
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Kranthi K. Mandadapu, Assistant Professor. Statistical Mechanics, Continuum Mechanics — Polycrystalline Materials, Biological Membranes, Bacterial Motility.
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Michael A. Marletta, Professor. Chemical biology, molecular biology, structure/function relationships in proteins, catalytic and biological properties of enzymes, cellular signaling, nitric oxide synthase, soluble guanylate cyclase, gas sensing, cellulose degradation, polysaccharide monooxygenases.
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Susan Marqusee, Professor. Amino acids, determinants of protein structure and folding, biophysical, structural and computational techniques, translocation, protein synthesis.
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Andreas Martin, Associate Professor. Proteasome.
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Richard Mathies, Professor of the Graduate School. Genomics, biophysical, bioanalytical, physical chemistry, laser spectroscopy, resonance Raman, excited-state reaction dynamics photoactive proteins, rhodopsins, microfabricated chemical biochemical analysis devices, forensics, infectious disease detection.
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Barbara J. Meyer, Professor. Developmental biology, gene expression, genetic determination of sex, regulatory genes, chromosome dynamics, X-chromosome.
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Evan W. Miller, Assistant Professor. Biophysical, bioanalytical and physical chemistry.
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Mohammad Mofrad, Professor. Nuclear pore complex and nucleocytoplasmic transport, mechanobiology of disease, cellular mechanotransduction, integrin-mediated focal adhesions.
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Priya Moorjani, Assistant Professor. Human population genetics and evolutionary biology.
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Eva Nogales, Professor. Biochemistry, complex biological assemblies, structure and regulation of the cytoskeleton, microtubule dynamics, human transcriptional initiation machinery, biophysics.
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Eunyong Park, Assistant Professor. Structure and mechanism of membrane transport proteins, cryo-EM, biochemistry, biophysics.
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Daniel S. Rokhsar, Professor. Biology, collective phenomena and ordering in condensed matter and biological systems, theoretical modeling, computational modeling, behavior of quantum fluids, cold atomic gases, high temperature superconductors, Fermi and Bose systems.
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David Savage, Assistant Professor. Synthetic biology and metabolism.
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Aaron Streets, Assistant Professor. Genomics and evolution of aging and cellular and organismal diversity.
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Peter Sudmant, Assistant Professor. Genomics, genetics, computational biology, structural variation, RNA, diversity, aging, population genetics.
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Frederic Theunissen, Professor. Behavior, cognition, brain, psychology, birdsong, vocal learning, audition, neurophysiology, speech perception, computational neuroscience, theoretical neuroscience.
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Evan Williams, Professor. Spectroscopy, molecular structure and dynamics, analytical chemistry, biophysical chemistry, structure and reactivity of biomolecules and biomolecule/water interactions, mass spectrometry, separations, protein conformation, protein and DNA sequencing.
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Ke Xu, Assistant Professor. Biophysical chemistry, cell biology at the nanoscale, super-resolution microscopy, single-molecule spectroscopy.
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Ahmet Yildiz, Associate Professor. Single molecule biophysics, molecular motors, telomeres.
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Roberto Zoncu, Assistant Professor. Biophysics and Structural Biology, MCB.
Research Profile

Contact Information

Biophysics Graduate Group

574 Stanley Hall

Phone: 510-642-0379

Fax: 510-666-3399

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Program Chair (as of 7/1/19)

Ahmet Yildiz, PhD (Physics)

Phone: 510-666-3792

Head Graduate Adviser

Susan Marqusee, MD, PhD (Molecular and Cell Biology)

Phone: 510-642-7678

Graduate Program Coordinator

Kate Chase

574 Stanley Hall, MC 3220

Phone: 510-642-0379

Program Chair (up to 7/1/19)

James Hurley, PhD (Molecular and Cell Biology)

Phone: 510-643-9483

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