Introduction to Program and Program Rationale

The primary objective of the Biophysics Program at Harvard University is to prepare investigators with diverse backgrounds for independent research careers in which the concepts and methods of physical science are applied to biological problems. The program is administered by the Committee on Higher Degrees in Biophysics , which is comprised of senior representatives from the Departments of Biological Chemistry and Molecular Pharmacology; Molecular and Cellular Biology; Chemistry and Chemical Biology; Physics; Genetics and Pathology. Owing to the interdepartmental nature of the program, research may be pursued on the Cambridge campus (in the Departments of Molecular and Cellular Biology; Chemistry and Chemical Biology; Physics; the Division of Engineering and Applied Sciences; etc.) or the Boston campus (including the Harvard Medical School, Division of Medical Sciences, and the 11 Harvard-affiliated teaching hospitals, which include the Dana-Farber Cancer Institute, Children’s Hospital, Massachusetts General Hospital, and the Beth Israel Deaconess Hospital).

The goal of the biophysics program is to nurture independent, creative scientists. To this end, the first part of the program seeks both to introduce the student to the faculty members and their research directly, enabling the student to make a considered choice of research advisor, and to involve the student in the diverse areas of biophysics through laboratory as well as coursework. This first two years provides a background for the second part of the program: the training of the student to be an independent scientist by a period of intensive research, culminating in publications and the PhD degree.

Admissions Requirements

Admissions and Scholarships

Application forms for admission and financial aid may be requested from the Admissions Office, Harvard Graduate School of Arts and Sciences, Holyoke Center, 3rd floor, 1350 Massachusetts Avenue, Cambridge, MA 02138. We encourage online submission of the application. See https://apply.embark.com/grad/Harvard/GSAS.

Applicants should state clearly in this application their desire to enter the program for the PhD degree in biophysics. All prospective students are urged to file the application forms well in advance of the deadline of December 8. Late applications will not be considered. Scores on the general Graduate Record Examination (GRE) are required except in special circumstances. GRE subject tests are recommended. Due to the early application deadline, applicants should plan to take GRE tests no later than October to ensure that original scores are received by the December 8 deadline. TOEFL is required of all foreign applicants other than those whose native language is English.

Final decisions concerning admission are made by the dean of the Graduate School of Arts and Sciences and the candidates are notified by letter from the Admissions Office.

After being accepted for admission, applicants are encouraged to make arrangements with the administrator of the Committee on Higher Degrees in Biophysics, Building C2, Room 122, 240 Longwood Avenue, Boston, MA 02115, (617-495-3360) to visit the University.

Students with the MD Degree

Students who already have the MD degree will find an opportunity to improve their knowledge of basic science in either of two ways:

1. Opportunities may be available in the several departments to engage in investigations as a research fellow under the direction of a member of the faculty. No university credit toward a degree is given for such work. Questions concerning the appointment of research fellows should be directed to the -faculty members.
2. Under special circumstances, students who have received the MD degree may become candidates for the PhD degree in biophysics, providing their qualifications for admission are approved and providing they are prepared to fulfill the normal requirements for the degree.

Combined MD-PhD Program

Students admitted to Harvard Medical School, as candidates for the MD degree, may also apply for admission to the biophysics program in order to earn a PhD degree in biophysics. This program may be of particular interest to prospective medical students with a strong background in physics and to students enrolled in the Harvard-MIT Division of Health Sciences and Technology.

Suggested Undergraduate Preparation

The large list of courses below would provide an ideal background for a student’s coursework in specialized areas of biophysics. No undergraduate major would have taken all these courses in college. Harvard course numbers are provided for further reference.

Math
Introduction to Calculus and Calculus, Series & Differential Equations (Math 1a and Math 1b)
Multivariable Calculus and Linear Algebra and Differential Equations (Math 21a
and 21b)
Advanced Calculus
Complex Analysis (Math 113)
Data Structures and Algorithms (Computer Science 124)

Physics
Introductory Mechanics and Relativity (Physics- 15a)
Introductory Electromagnetism (Physics 15b)
Wave Phenomena (Physics 15c)
Quantum Mechanics I and II (Physics 143a and 143b)
Introduction to Biophysics (Physics 140)

Chemistry
Organic Chemistry (Chem 20 and Chem 30)
Physical Chemistry and Statistical Thermodynamics (Chem 160 and 161)
Principles of Organic Chemistry (Chem 17)

Biology
Evolutionary Biology (OEB 53)
Cell Biology (MCB 54)
Biochemistry and Physical Properties of Macromolecules- (MCB 56)
Neurobiology of Behavior (MCB 80)

Biochemistry
Molecular Biology (MCB 52)
An Integrated Introduction to the Life Sciences: Genetics, Genomics, and Evolution (Life Sciences 1a)

Advice to International Students

Students from non-US countries who are continuing their studies at Harvard, or who are applying for admission to the University, should communicate with the International Office, 1350 Massachusetts Avenue, Room 851, Cambridge, MA 02138, which is especially designed to help and advise international students. The immigration and naturalization laws specify many complex legal requirements affecting the status of international students and scholars during their period of study in the United States. The office is prepared to furnish information to aliens on visa requirements, permissible employment, income tax liabilities, and many other official restrictions in effect under existing laws of the United States.

Financial Aid

All students accepted into the program are awarded full support, including a stipend, full tuition, and health fees contingent on continued satisfactory progress. Teaching fellowships are available. Second-year students must teach one term as part of their academic requirements. Students are responsible for finding their own teaching position in any one of the participating science departments.

Students are encouraged to apply for external fellowships, such as those administered by the National Science Foundation, the National Defense Science and Engineering Fellowship, and the National Institutes of Health, which are available on a competitive basis to graduate students in biophysics. Applications for those fellowships should be made directly to the appropriate agencies.

Certain limited funds are available as beneficiary aid, distributed by the deans of the Graduate School to students in the form of gifts or loans to meet unexpected financial needs. Information on loans and beneficiary aids may be obtained from the Graduate School.

Program of Study

Most graduates of the biophysics program at Harvard have been undergraduate majors in physics or physical chemistry, though a few have come from biology. Consequently, the course requirements for admission are flexible. Each student’s program of graduate study is planned in consultation with a faculty advisor. The degree program is designed to be completed in a maximum of six years.

The first year’s training in the Biophysics Program provides an introduction to five diverse areas of Biophysics: 1) Structural Molecular Biology; 2) Cell and Membrane Biophysics; 3) Molecular Genetics; 4) Physical Biochemistry; and 5) Neurosciences. The curriculum includes learning experiences in a laboratory environment as well as coursework. The program is flexible and special effort has been devoted to minimizing formal requirements.

The laboratory experience is organized as a full course, Biophysics 300. In the first 10 weeks of the fall term, faculty members associated with the biophysics program give seminars describing the current research interests of their own laboratories. Following this, a student spends six-week periods in each of three different laboratories. The list under Participating Faculty and Their Special Fields names some of our professors who currently support biophysics rotation students. In order to make sure that the student gains familiarity with several fields of biophysics, each of the three laboratory experiences usually is selected from a different one of the areas of biophysics listed above. It will also be possible to work on a suitable problem in mathematical biophysics in place of one of the three laboratory rotations.

Biophysics 101, Genomics, and Computational Biology, focuses on modern technologies with exponential growth and their impact on global quality of life through weekly updated Wiki class project (in-depth case studies on personal genomics and/or biofuels). Integrating knowledge, tools for research, and commercial decision-making concerning new aspects of bioengineering, personalized medicine, genetically modified organisms, and stem cells. Interplays of biophysical, ecological-, economic, and social/ethical modeling- will be explored through multi-disciplinary teams of students, and individual brief reports.

Biophysics 204, Structural Biology From Molecules to Cells, explores imaging of molecules and of molecular localization in cells, including x-ray and electron crystallography, electron microscopy of single molecules, and high-resolution light microscopy.

Biophysics 205, Computational and Functional Genomics, explores experimental functional genomics, computational prediction of gene function, and properties and models of complex biological systems. This course primarily involves critical reading and discussion rather then lectures.

Biophysics 242, Special Topics in Biophysics, invites two or more professors in the spring term to speak on new areas of their own research, thereby introducing students to new questions and methods of research that would not normally be accommodated by the regular curriculum. Past topics have included structure and function of DNA; oncogene products; two-dimensional nuclear magnetic resonance; RNA structure and function; diffusion; atomic interactions in protein-ligand interactions; analysis and design of novel protein-protein and protein-ligand interactions based on 3-D structural information; structure/ function relationships in peptide antibiotics; biological interactions at surfaces; enzyme-catalyzed redox reactions: catalysts and cofactors; molecular neurobiology and systems neurosciences; conceptual foundations and recent developments in computational biology, genomics, and macromolecular interactions; new biology through physics: molecular discoveries with light, information theory and neural systems, X-ray and electron crystallography and high-resolution light microscopy, and computational and functional genomics ; and molecular motors.

In addition to these courses, each student normally takes one course in each of the five biophysics areas mentioned previously. Most coursework is completed within the first two years. A student may also gain knowledge in a particular area by taking reading courses or by studying independently while registered for TIME.

Courses in Biophysics and
Related Fields

Courses in the following list have been divided roughly into the five areas plus mathematical biophysics. However, the list is not all-inclusive; further offerings for any given term may be found in the Harvard University list of Courses of Instruction.

Structural Molecular Biology
Genomics, and Computational Biology
(Biophysics 101)
Chemical Biology (Chemistry 270)
Molecular Biology (BCMP 200)
Proteins: Structure, Function and Catalysis (BCMP 201)
Macromolecular NMR (BCMP 228)
Crystal Symmetry, Diffraction, and Structure Analysis (Chemistry 154)
Macromolecular Structure and Function (Chemistry 280)
Structural Biology of the Flow of Information in the Cell (MCB 156)
Molecular Microbiology and Pathogenesis (Microbiology 200)
Mechanisms of Microbial Pathogenesis (Microbiology 205)
Structural Biology From Molecules to Cells (Biophysics 204)

Molecular Genetics
Molecular Genetics of Neural Development and Behavior (MCB 129)
Developmental Genetics and Genomics (MCB 150)
Molecular Mechanisms of Gene Control (MCB 155)
Principles of Genetics (Genetics 201)

Physical Biochemistry
Introduction to Biophysics (MCB 140)
Physical Chemistry (Chemistry 160)
Frontiers in Biophysics (Chemistry 163)
Topics in Biophysics (MCB 212)
Quantum Mechanics I (Chemistry 242)
Statistical Thermodynamics (Chemistry 161)
Experimental Physical Chemistry
(Chemistry 165)

Cell and Membrane Biophysics
Molecular and Cellular Immunology
(MCB 169)
Biochemistry of Membranes (MCB 176)
Molecular Biology of the Cell (Cell
Biology 201)
Growth Factors and Signal Transduction
(Cell Biology 211b)

Mathematical Biophysics
Complex and Fourier Analysis (Applied Mathematics 105a)
Ordinary and Partial Differential Equations (Applied Mathematics 105b)
Methods of Analysis (Math 115)
Physiological Systems Analysis (Eng. Sci. 145)
Nonlinear Dynamical Systems (Applied Mathematics 147)
Population Genetics (OEB 152)
Signals and Systems (Eng. Sci. 156)
Physical Mathematics I, II (Applied Mathematics 201, 202)
Fundamentals of Computational Biology (Statistics 215)
Mathematical Modeling (Applied
Mathematics 115)
Mathematics in Biology (MCB 111)

Neurosciences
Cellular Basis of Neuronal Function
(MCB 115)
Experimental Neuroscience (MCB 117)
Molecular and Developmental Neurobiology (MCB 141)
Neural Signal Processing (Eng. Sci. 148)
Introduction to Neurobiology
(Neurobiology 200)
Neurophysiology of Central Circuits
(Neurobiology 204)
Molecular Neurobiology (Neurobiology 221)
Systems Neuroscience (MCB 105)

Formal Academic Requirements

The academic requirement for the PhD degree consists of not less than two years—at least one of which must be in residence at the Harvard Graduate School of Arts and Sciences —devoted to advanced studies approved as suitable preparation for the degree by the Committee on Higher Degrees in Biophysics. In estimating the extent of a candidate’s study for the degree, the advanced work done in other graduate departments of Harvard or of other universities will be considered. A year’s work for a resident student normally consists of four courses (eight half-courses) of advanced grade. Under certain conditions summer courses taken at the Marine Biological Laboratory in Woods Hole may be counted for credit toward the degree.

The biophysics program anticipates completion of formal course studies in the first two years.

Languages. There is no language examination but students are encouraged to gain a reading knowledge of one foreign language, preferably selected from German, Russian, and French.

Dissertation Qualifying Examination. Before beginning dissertation research, it is normally necessary for the student to fulfill the following requirements: (1) pass one Harvard course in each of the six subject areas listed above; (2) do satisfactory work in three laboratory rotations; and (3) submit and defend an original research proposal. The purpose of the oral defense of the qualifying research proposal is to ensure that the student is adequately prepared to embark on dissertation research. The exam is normally taken in the fourth term of residence, before the chair and three examiners knowledgeable in the field of the research proposal. Reexamination will be permitted. As a rule, students will not be permitted to enter the third year of graduate study unless the qualifying examination has been passed.

Dissertation. Selection of a dissertation advisor normally occurs in a student’s second year of study. Independent research on one’s dissertation technically begins once the qualifying examination is successfully completed.

The program policy is that all students who have successfully completed their qualifying exam promptly establish their Dissertation Advisory Committee (DAC). The DAC has several functions. The first will be to approve the student’s dissertation proposal. Second, the DAC, in consultation with the dissertation advisor, will periodically evaluate the progress of the student’s dissertation research. The DAC may make recommendations to the dissertation advisor as well as the student with regard to the student’s progress towards completion of the dissertation. Third, the DAC, in consultation wth the dissertation advisor, will determine at what point the student is ready to defend his or her dissertation.

It is expected that the preparation of a dissertation will usually require full-time for not less than one-and-a-half years after the qualifying examination. The dissertation must give evidence of independent original research and be clearly, logically, and carefully written in good English. The final manuscript must conform to the requirements described in The Form of the PhD Dissertation.

On receipt of the dissertation, the chair of the Committee on Higher Degrees in Biophysics will appoint a reading committee of three to judge the dissertation. The dissertation defense is comprised of two parts: the public seminar and the private defense. In the private defense the candidate will be questioned on the subject of the dissertation and its relation to the student’s special field and collateral subjects. If the reading committee is unable to agree on its recommendations, the question of the acceptance of the dissertation will be decided by the Committee on Higher Degrees in Biophysics.

Upon completion of all the requirements, the original bound dissertation, with the dissertation acceptance certificate signed by the reading committee, and one unbound copy, will be deposited at the Office of the Registrar of the Graduate School, for inspection by any member of the Faculty of Arts and Sciences.

After Commencement, the original bound copy will be deposited in the Harvard Library, open to public inspection. A second bound copy will be kept by the committee chair, and a third bound copy will be kept by the department in which the student worked.

Selected PhD Dissertation Titles

William W.L. Chen (2006). “Theoretical and Computational Studies in Protein Biophysics: Folding, Information, and Networks”

Maria A. Neimark Geffen (2006). “Encoding of Complex Stimuli in Early Sensory Systems”

Avniel S. Ghuman (2007). “Neural Dynamics and Interactions in Top-Down Facilitation of Visual Cognition”

Vidyasagar Koduri (2007). “Folding and Maturation of Lipoprotein Receptors”

Nikos B. Reppas (2007). “Genome-scale Analyses of Chromosomal Accessibility, Transcriptional Initiation, and Synthetic Crossfeeding in E.coli”

Laurence A. Shumway (2006). “Evolution of Altered Signaling in the Yeast Saccharomyces Cerevisiae”

Leo Lee Tsai (2006). “Development of a Low-field 3He MRI System to Study Posture-Dependence of Pulmonary Function”

Ellen Yeh (2006). “Enzymatic Halogenation During Natural Product Biosynthesis”

Participating Faculty and Their Special Fields

The following is a partial list of faculty members who will accept students for degree work in biophysics, with their special fields of research and instruction indicated. More than 60 members of the Harvard faculty are currently affiliated with the Biophysics program; dissertation research with other faculty members is possible by arrangement. A more complete listing is available.

James M. Hogle, PhD, Edward S. Harkness Professor of Biological Chemistry and Molecular Pharmacology, Harvard Medical School; Chair, Committee on Higher Degrees in Biophysics.Structure and function of viruses and virus related proteins; x-ray crystallography.

John Assad, PhD, Associate Professor of Neurobiology, Harvard Medical School. Mechanisms of visual processing in the visual cortex of awake behaving monkeys.

Frederick Michael Ausubel, PhD, Professor of Genetics, Harvard Medical School. Molecular biology of bacterial pathogenesis in plants and animals.

Brian J. Bacskai, PhD, Associate Professor of Neurology, Harvard Medical School. Multiphoton microscopy in imaging Alzheimer’s disease.

Howard C. Berg, PhD, Herchel Smith Professor of Physics and Professor of Molecular and Cellular Biology. Motile behavior of bacteria.

Stephen C. Blacklow, MD, PhD, Associate Professor of Pathology, Harvard Medical School. Biophysics of receptor-ligand interactions.

William Hines Bossert, PhD, David B. Arnold, Jr. Professor of Science. Theoretical Population and Community Biology, models of parasitic disease processes.

Martha L. Bulyk, SB, PhD, Assistant Professor of Medicine and Health Sciences and Technology, Harvard Medical School. Functional and computational genomics studies of transcription factors and Cis regulatory elements.

Lewis Cantley, PhD, Professor of Systems Biology, Harvard Medical School. Structural basis for specificity in eukaryotic signal transduction pathways.

James J. Chou, PhD, Assistant Professor of Biological Chemistry and Molecular Pharmacology, Harvard Medical School. NMR spectroscopy on membrane-associated proteins and peptides.

George M. Church, PhD, Professor of Genetics, Harvard Medical School. Human and microbial functional genomics, genotyping, and gene expression regulatory network models.

David E. Clapham, MD, PhD, Aldo R. Castaneda Professor of Cardiovascular Research and Professor of Neurobiology, Harvard Medical School. Imaging, optics and biology.

Jon C. Clardy, PhD, Professor of Biological Chemistry and Molecular Pharmacology. Small molecule signaling, biosynthesis and drug discovery.

Jonathan B. Cohen, PhD, Professor of Neurobiology, Harvard Medical School.
Structure and function of ligand-gated ion channels.

R. John Collier, PhD, Maude and Lillian Presley Professor of Microbiology and Molecular Genetics, Harvard Medical School. Structure and activity of bacterial toxins.

David P. Corey, PhD, Professor of Neurobiology, Harvard Medical School. Biophysics of Mechanosensation.

Bruce Francis Demple, PhD, Professor of Toxicology in the Faculty of Public Health. Redox Signaling and Repair of Oxidative DNA Damage.

John Elliot Dowling, PhD, Gordon and Llura Gund Professor of Neurosciences and Professor of Opthalmology. Biophysical aspects of the visual system.

Michael J. Eck, MD, PhD, Associate Professor of Biological Chemistry and Molecular Pharmacology, Harvard Medical School. Structural studies of proteins involved in -signal transduction pathways.

Florian Engert, MA, PhD, Associate Professor in Molecular and Cellular Biology. Synaptic plasticity and neuronal networks.

Raymond Leo Erikson, PhD, American Cancer Society Professor of Cellular and Developmental Biology. Protein kinases, reversible protein phosphorylation.

Rachelle Gaudet, PhD, Associate Professor in Molecular and Cellular Biology. Structural studies of the stereochemistry of signaling and transport through biological membranes.

David Golan, MD, PhD, Professor of Biological Chemistry and Molecular Pharmacology and Medicine, Harvard Medical School. Membrane dynamics; membrane structure; cellular adhesion.

Jene A. Golovchenko, PhD, Gordon McKay Professor of Molecular and Cellular Biology. Probing polymers with nanopores, experimental condensed matter physics.

Edward E. Harlow, PhD, Professor of Biological Chemistry and Molecular Pharmacology, Harvard Medical School. Tumor suppressor genes; cell cycle control.

Stephen Coplan Harrison, PhD, Professor of Biological Chemistry and Molecular Pharmacology. Structure of viruses and viral membranes; protein-DNA interactions; X-ray diffraction.

John Woodland Hastings, PhD, Paul C. Mangelsdorf Professor of Natural Sciences. Molecular mechanisms in bioluminescent reactions and circadian rhythms.

Donald E. Ingber, MD, PhD, Judah Folkman Professor Vascular Biology, Harvard Medical School. Research in integrin signaling, cytoskeleton, and control of angiogenesis.

David Jeruzalmi, BS, M.Phil, PhD, Associate Professor in Molecular and Cellular Biology. Structural studies of nucleo-protein assemblies.

Tomas Kirchhausen, PhD, Professor of Cell Biology, Harvard Medical School. Molecular mechanisms of membrane traffic, x-ray crystallography; chemical genetics.

Roy Kishony, PhD, Assistant Professor of Systems Biology, Harvard Medical School. System-level Genetic Networks.

Nancy Elizabeth Kleckner, PhD, Herchel Smith Professor of Molecular Biology. Molecular and Mechanical Analysis of Chromosomes.

Roberto G. Kolter, PhD, Professor of Microbiology and Molecular Genetics, Harvard Medical School. DNA protection from oxidative damage, cell-cell communication in biofilms, microbial evolution.

Galit Lahav, PhD, Assistant Professor of Systems Biology, Harvard Medical School. Dynamics of network motifs in single living human cells.

David R. Liu, PhD, Professor of Chemistry and Chemical Biology. Amplifiable and evolvable unnatural molecules; natural molecule evolution; engineering of synthetic and biological molecules.

Jun Liu, BS, PhD, Professor of Statistics. Statistical theory and inference for stochastic processes: with applications to bioinformatics.

Gavin MacBeath, BS, PhD, Associate Professor of Chemistry and Chemical Biology. Chemical biology and systems biology.

Tom P. Maniatis, PhD, Thomas H. Lee Professor of Molecular and Cellular Biology. Eukaryotic gene expression.

Jarrod A. Marto, PhD, Assistant Professor of Biological Chemistry and Molecular Pharmacology, Harvard Medical School. Quantitative proteomics of cancer progression.

Markus Meister, PhD, Jeff C. Tarr Professor of Molecular and Cellular Biology, Department of Molecular and Cellular Biology. Function of neuronal circuits.

Keith Wyatt Miller, DrPhil, Mallinckrodt Professor of Pharmacology, Department of Anaesthesia, Harvard Medical School. Characterization of lipid-protein interactions; regulatory confirmation changes and molecular mechanisms of drug action on ion channels from nerve membranes using rapid kinetics and spectroscopy (EPR, NMR, and FTIR).

Timothy Mitchison, PhD, Hasib Sabbagh Professor of Systems Biology, Harvard
Medical School. Cytoskeleton dynamics; mechanism of mitosis and cell locomotion; small molecule inhibitors.

Andrew W. Murray, PhD, Herchel Smith Professor of Molecular Genetics. Regulation of mitosis.

Venkatesh N. Murthy, PhD, Morris Kahn Associate Professor of Molecular and Cell Biology. Mechanisms of synaptic transmission and plasticity.

Erin K. O’Shea, PhD, Professor of Molecular and Cellular Biology. Systems level and molecular analysis of signaling pathways, transcriptional regulation, and methods for expressing and assaying the proteins derived from an organism.

David S. Pellman, MD, Associate Professor of Pediatrics, Harvard Medical School. The mechanics and regulation of mitosis.

Mara Prentiss, PhD, Mallinckrodt Professor of Physics. Exploitation of optical manipulation to measure adhesion properties, including virus cell binding.

Tom A. Rapoport, PhD, Professor of Cell Biology, Harvard Medical School. Mechanism of how proteins are transported across the endoplasmic reticulum membrane.

Frederick P. Roth, PhD, Assistant Professor of Biological Chemistry and Molecular Pharmacology, Harvard Medical School. Computational molecular biology.

Gary Ruvkun, PhD, Professor of Genetics, Harvard Medical School. Genetic control of developmental timing, neurogenesis, and neural function.

Bernardo L. Sabatini, BS, PhD, MD, Associate Professor of Neurobiology, Harvard Medical School. Regulation of synaptic transmission and dendritic function in the mammalian brain.

Aravinthan D.T. Samuel, PhD, Assistant Professor of Physics. Behavioral neuroscience and neurophysiology.

Stuart Schreiber, PhD, Morris Loeb Professor of Chemistry and Chemical Biology. Forward and reverse chemical genetics: using small molecules to explore biology.

Brian Seed, PhD, Professor of Genetics, Harvard Medical School. Genetic analysis of signal transduction in the immune system.

Jagesh V. Shah, PhD, Assistant Professor of Systems Biology, Assistant Professor of Health Sciences and Technology, and Assistant Professor of Medicine, Harvard Medical School. Quantitative models of cellular behavior to investigate protein function.

Eugene Shakhnovich, PhD, Professor of Chemistry and Chemical Biology. Theory and experiments in protein folding and design; theory of molecular evolution; rational drug design and physical chemistry of protein-ligand interactions; theory of complex systems.

William M. Shih, PhD, Assistant Professor of Biological Chemistry and Molecular Pharmacology, Harvard Medical School. Biomolecular nanotechnology.

Steven E. Shoelson, MD, PhD, Professor of Medicine, Harvard Medical School. Structural and cellular biology of insulin -signal transduction-.

Pamela Silver, PhD, Professor of Systems Biology, Harvard Medical School. Dynamics of the nucleus, regulation of nuclear transport and partioning of nuclear components during cell division. Systems cell biology.

Timothy A. Springer, PhD, Latham Family Professor of Pathology, Harvard Medical School. Biophysics of cell adhesion and vascular shear flow.

Shamil R. Sunyaev, PhD, Assistant Professor of Medicine and Health Sciences and Technology, Harvard Medical School. Computational methods in genetics, genomics, and proteomics-.

Jack W. Szostak, PhD, Professor of Genetics, Harvard Medical School. Directed evolution; information content and molecular function; self-replicating systems.

Naoshige Uchida, PhD, Assistant Professor of Molecular and Cellular Biology. Sensory Information in Neuronal Processes.

Antoine van Oijen, PhD, Assistant Professor of Biological Chemistry and Molecular Pharmacology, Harvard Medical School. Single-molecule novel fluorescence and nano-manipulation studies of protein-protein and protein-nucleic acid interactions.

Gregory Verdine, PhD, Erving Professor of Chemistry and Chemical Biology. Protein-nucleic acid interactions; transcriptional regulation; x-ray crystallography.

Gerhard Wagner, PhD, Elkan Blout Professor of Biological Chemistry and Molecular
Pharmacology, Harvard Medical School. Protein and nucleic acid structure, interaction and mobility; NMR spectroscopy.

John R. Wakeley, BS, MS, PhD, Professor of Organismic and Evolutionary Biology. Theoretical population genetics.

Christopher T. Walsh, PhD, Hamilton Kuhn Professor of Biological Chemistry and Molecular Pharmacology, Harvard Medical School. Enzymatic reaction and antibiotic synthesis mechanisms.

Ronald L. Walsworth, PhD, Senior Lecturer on Physics. Experimental atomic physics, biophysics, and soft matter physics.

Thomas Walz, PhD, Professor of Cell Biology, Harvard Medical School. High-resolution electron microscopy.

George Whitesides, PhD, Woodford L. and Ann A. Flowers University Professor. Rational drug design; biomaterials science and biophysics.

Kai W. Wucherpfennig, MD, PhD, Professor of Neurology, Harvard Medical School. Basic mechanisms of T cell mediated autoimmune diseases.

Xiaoliang Sunney Xie, PhD, Professor of Chemistry and Chemical Biology. Single molecule spectroscopy and dynamics and molecular interaction and chemical -dynamics in biological systems.

Gary I. Yellen, PhD, Professor of Neurobiology, Harvard Medical School. Molecular physiology of ion channels: functional motions, drug interactions, and electrophysiological mechanisms.

Xiaowei Zhuang, BS, MS, PhD, Professor of Chemistry and Chemical Biology and of Physics. Single-molecule biophysics.