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Department of Molecular Biology


Lynn W. Enquist

Associate Chair

James R. Broach

Departmental Representative

S. Jane Flint

Yibin Kang

Mark D. Rose

Thomas J. Silhavy

Director of Graduate Studies

Jean E. Schwarzbauer


Bonnie L. Bassler

David Botstein, also Lewis-Sigler Institute for Integrative Genomics

James R. Broach

Edward C. Cox

Lynn W. Enquist, also Princeton Neuroscience Institute

S. Jane Flint

Jacques R. Fresco

Elizabeth R. Gavis

Frederick M. Hughson

Mark D. Rose

Paul D. Schedl

Gertrud M. Schüpbach

Jean E. Schwarzbauer

Thomas E. Shenk

Thomas J. Silhavy

Lee M. Silver, also Woodrow Wilson School

Jeffry B. Stock

David W. Tank, also Princeton Neuroscience Institute

Saeed Tavazoie, also Lewis-Sigler Institute for Integrative Genomics

Shirley M. Tilghman

Eric F. Wieschaus, also Lewis-Sigler Institute for Integrative Genomics

Ned S. Wingreen, also Lewis-Sigler Institute for Integrative Genomics

Virginia A. Zakian

Associate Professor

Michael J. Berry, also Princeton Neuroscience Institute

Carlos D. Brody, also Princeton Neuroscience Institute

Yibin Kang

John D. Storey, also Lewis-Sigler Institute for Integrative Genomics

Samuel S. H. Wang, also Princeton Neuroscience Institute

Assistant Professor

Lisa M. Boulanger, also Princeton Neuroscience Institute

Rebecca D. Burdine

Hilary A. Coller

Ileana M. Cristea

Jonathan T. Eggenschwiler

Benjamin A. Garcia

Zemer Gitai

Manuel Llinás, also Lewis-Sigler Institute for Integrative Genomics

Coleen T. Murphy, also Lewis-Sigler Institute for Integrative Genomics

Mala Murthy, also Princeton Neuroscience Institute

Senior Lecturer

Alison E. Gammie

Alan Gelperin, also Princeton Neuroscience Institute

Heather A. Thieringer

Lecturer with Rank of Professor

Adel A. Mahmoud, also Woodrow Wilson School

Leon E. Rosenberg


Robyn E. Tanny

Associated Faculty

Peter Andolfatto, Ecology and Evolutionary Biology, Lewis-Sigler Institute for Integrative Genomics

Jannette L. Carey, Chemistry

Thomas Gregor, Physics, Lewis-Sigler Institute for Integrative Genomics

Michael H. Hecht, Chemistry

Leonid Kruglyak, Ecology and Evolutionary Biology, Lewis-Sigler Institute for Integrative Genomics

Laura F. Landweber, Ecology and Evolutionary Biology

A. James Link, Chemical and Biological Engineering

Celeste M. Nelson, Chemical and Biological Engineering

Joshua D. Rabinowitz, Chemistry, Lewis-Sigler Institute for Integrative Genomics

Joshua W. Shaevitz, Physics, Lewis-Sigler Institute for Integrative Genomics

Stanislav Y. Shvartsman, Chemical and Biological Engineering, Lewis-Sigler Institute for Integrative Genomics

Mona Singh, Computer Science, Lewis-Sigler Institute for Integrative Genomics

David L. Stern, Ecology and Evolutionary Biology

Olga G. Troyanskaya, Computer Science, Lewis-Sigler Institute for Integrative Genomics

Information and Departmental Plan of Study

At Princeton, courses in the biological sciences are offered in two departments. Students with interests in molecular, cellular, and developmental processes should enroll in the Department of Molecular Biology. Those with an evolutionary orientation and interest in organismal, population, and community processes should enroll in the Department of Ecology and Evolutionary Biology.

Every student considering majoring in the department is encouraged to attend a departmental open house that is held in the spring term to introduce students to the departmental courses, faculty, and research interests.


Prerequisites for entry into the Department of Molecular Biology are the courses MOL 214 or MOL 215.

Early Concentration

Qualified students who have been granted advanced placement credit in departmental prerequisites, and who have taken advanced courses in molecular biology (300 level and above) during the freshman and/or sophomore years may be eligible for independent work in the spring of the sophomore year and/or fall of the junior year. To qualify for early concentration status, students must have received grades of B+ or better on departmental prerequisites and advanced molecular biology courses. Early concentrators may engage in experimental research in the laboratories of members of the department and associated faculty. A limited number of early concentrators will be offered the possibility of continuing research in the laboratories of their faculty advisers during the summer following the sophomore year. Students who are interested in an early concentration should contact a departmental representative early in the sophomore year.

Early concentrators have the option of doing more intensive experimental research as part of an independent reading course in an area of molecular biology, under the supervision of a faculty member of the department, in the spring of the sophomore year or the fall of the junior year. Independent study will take the form of an intensive reading course intended to prepare students for experimental research in molecular biology. Students will read relevant background literature for their overall field of study, as well as literature pertinent to experimental techniques. Based upon the experience of the students, and at the discretion of the adviser, students may begin experimental laboratory research. Students will meet with faculty advisers on a weekly basis to discuss assigned literature, plan experiments, and review results. At the end of each semester, students will write a term paper based on the literature and any experimental research undertaken during the semester. The paper may take the form of a grant proposal or a research report in the format of a scientific journal article, including a scholarly review of the literature along with description and discussion of the results. Students interested in an independent reading course must consult with a departmental representative early in the prior semester and obtain approval for the plan of study.

Program of Study

The following courses are requirements and, in general, should be completed before the beginning of the junior year:

General Chemistry (CHM 201 and 202) or CHM 215 or two units of CHM credit

Organic Chemistry (CHM 301 and 302) or (CHM 303 and 304)

Mathematics (MAT 101 and 102) or (MAT 103 and [MAT 104 or COS 126, PSY 251, MOL 410, MOL 436, MOL/EEB 355]). Other courses may be substituted upon approval by a departmental representative.

General Physics (PHY 101 and 102) or (PHY 103 and 104)

EB 211 (or equivalent AP plus one 300-level EEB course)

All of the above prerequisites and requirements, with the exception of MOL 214 or MOL 215, can be satisfied with advanced placement and/or summer courses at other universities (within University guidelines). MOL 214 or MOL 215 must be taken at Princeton University. If EEB 211 is not taken at Princeton (for example, because of advanced placement), the student must take one upper-level course (300 level or above) in the Department of Ecology and Evolutionary Biology, which can also count as a departmental. All 300-level and above courses, where EEB is the primary listing, will be accepted; cross-listed courses must be individually approved.

The following departmental courses are required:

Genetics (MOL 342)
Biochemistry (MOL 345)
Cell and Developmental Biology (MOL 348)
Core Lab (MOL 350)

Except under very special circumstances, these courses must be taken during the junior year or earlier. All count as departmentals and they must be successfully completed. No substitutes are allowed, except for students who wish to study abroad and who have received departmental approval. Additional departmentals can be chosen from among all 300 or higher-level courses in molecular biology, and selected upper-level courses with a strong molecular or biological component from other departments (see list). Other courses can only be taken as departmentals with the written approval of a departmental representative. All students must take at least eight, but not more than 12, departmentals. Only Princeton courses can count as departmentals; there are no exceptions to this rule.

A degree from the molecular biology department requires successful completion of the prerequisites in MOL 342, MOL 345, MOL 348, MOL 350, and four other departmentals.

No courses in biological sciences or other departmental courses may be taken pass/D/fail.

Independent Work

Junior Independent Work. In the fall semester of the junior year students participate in tutorials with postdoctoral instructors, read papers from the original literature, and prepare two short papers on assigned topics. In the spring term students carry out a second program of independent work with a faculty adviser with whom they will eventually do their senior thesis. In some instances this may include experimental work. A monograph summarizing this work is due in early May.

Senior Independent Work. During the senior year each student, with the guidance of a faculty adviser, undertakes a major research effort. This research project can be a laboratory, field, or independent study that will be written and presented as a senior thesis.

Senior Departmental Examination

Students are required to present their work to the two (non-adviser) thesis readers during an oral exam, at which the adviser is not present. The exam usually takes about one-half hour and students should be prepared to describe the background of the thesis, defend its contents, and propose future directions. The grade for the oral defense will be the average of the two from the (non-adviser) faculty members. A grading rubric will be used by the examination committee. Grades are assigned by the Undergraduate Committee with the approval of the faculty.

Study Abroad

Students concentrating in molecular biology may spend part of the junior year abroad, provided that at least one core departmental course has been taken in the sophomore year and suitable arrangements are made to complete the junior independent work. Students may take part in existing exchange programs with the University of Oxford and the Karolinska Institute or may arrange, in consultation with departmental representatives and the Office of International Programs, to participate in other approved programs abroad. Study abroad is most easily arranged for the fall semester. Students interested in study abroad should consult with departmental representatives and Dean Nancy Kanach as early as possible, preferably during the first year.

Integrated Science Sequence

An alternative path into the department is through the integrated science curriculum. ISC/CHM/COS/MOL/PHY 231-4 (a double course) can be taken in the freshman year, and ISC/CHM/COS/MOL/PHY 235-6 can be taken in the sophomore year. These courses can be substituted for CHM 203-204, PHY 103-104 or 105-6, and COS 126 in the freshman year and MOL 214, 342, and 345 in the sophomore year. For full course descriptions and more information, see the integrated science website.

Approved Courses for Departmental Credit. See the departmental website for an up-to-date list of approved departmentals. Other courses may be approved upon consideration by a departmental representative. Note: Only one non-MOL course, in which the content is primarily related to ethical, social science, or policy implications of biomedical topics, will be accepted for departmental credit.

Early Research Opportunities. Students interested in research prior to junior year should consult the faculty Web page to become familiar with the types of research being conducted in the department. Students should then meet with the freshman/sophomore adviser, who can provide information about specific research opportunities.

Program in Biophysics. The Program in Biophysics is designed for students with strong interests in molecular biology and physics who wish to combine these two subjects in their junior and senior independent work. The program offers a combination of courses and interdisciplinary research that meet the requirements of the physics or molecular biology departments, and entry requirements of graduate schools in both physics and molecular biology. Courses are chosen with the help of advisers in the Departments of Physics and Molecular Biology. A certificate in biophysics is awarded to students who successfully complete the program. Students are admitted to the program once they have chosen their field of concentration and consulted with the program director, who will assign them an adviser.

Program in Global Health and Health Policy. The Program in Global Health and Health Policy is an interdepartmental program in which undergraduates can study the determinants, consequences, and patterns of disease across societies; the role of medical technologies and interventions in health improvements; and the economic, political, and social factors that shape domestic and global public health. In addition to the core departmental courses, molecular biology concentrators would take GHP 350 by the end of junior year and GHP 351 by the end of senior year. Most upper-level MOL courses fulfill the requirements for the global health and health policy certificate.

Program in Neuroscience. The department offers the opportunity for concentrators to participate in the neuroscience program. Interested students should discuss the program with the directors and their departmental representative.

Program in Quantitative and Computational Biology. The Program in Quantitative and Computational Biology (QCB) is designed for students with a strong interest in multidisciplinary and systems-level approaches to understanding molecular, cellular, and organismal behavior. The curriculum introduces the students to experimental and analytic techniques for acquisition of large-scale quantitative observations, and the interpretation of such data in the context of appropriate models. Strong emphasis is placed on using global genome-wide measurements (e.g., microarray gene expression, sequence, phenotype) to understand physiological and evolutionary processes. At the core of the curriculum is the project lab (QCB 301), a double-credit laboratory course, taken during the fall of junior year, in which students participate in the design, execution, and analysis of experiments. The required courses provide a strong background in modern methodologies in data analysis, interpretation, and modeling. Courses are chosen with the help of advisers in molecular biology, ecology and evolutionary biology, physics, chemistry, computer science, and other related departments. A certificate in quantitative and computational biology is awarded to students who successfully complete the program requirements.


MOL 101B From DNA to Human Complexity   Fall ST

This lecture and laboratory course will acquaint nonbiology majors with the theory and practice of modern molecular biology, with a focus on biological topics of current public interest. Topics include: structure of DNA, RNA, proteins, genomes, and an overview of state-of-the-art technologies including cloning, recombinant DNA, and PCR. The course will address how recent scientific advances impact issues relevant to human biology, including understanding how genes control complex patterns of cell differentiation and the origins of mutations and inherited defects. Three lectures, one three-hour laboratory. E. Wieschaus, H. Thieringer, B. Bassler

MOL 110 Neuroscience and Everyday Life (see NEU 101)

MOL 205 Genes, Health, and Society   Spring SA

What should students know about their genes? Today, the field of human genetics is explored and debated like no other. To understand the medical applications and ethical implications of human genetics, one must grasp its scientific foundations. The course approaches these topics using lectures; textbook, journal, and newspaper readings; precept discussions; and patient interviews. It will consider: gene structure and function; the genetics and genomics of populations and of selected human disorders (cancer, mental illness, metabolic diseases); and clinical genetics (inheritance patterns, diagnosis, treatment). Two lectures, one preceptorial. L. Rosenberg

MOL 210 Evolutionary Ecology (see EEB 210)

MOL 211 The Biology of Organisms (see EEB 211)

MOL 214 Introduction to Cellular and Molecular Biology (also EEB 214)   Spring ST

Important concepts and elements of molecular biology, biochemistry, genetics, and cell biology are examined in the context of classic experiments. During the last four weeks, the class will split into topic-based sections taught by different faculty in the following areas: cell biology, biochemistry, neuroscience, microbiology, and development. This course is strongly recommended as the introductory course for students who are considering majoring in the biological sciences. It also satisfies the biology requirement for entrance to medical school. Two 90-minute lectures, one three-hour laboratory. T. Shenk, H. Thieringer

MOL 215 Quantitative Principles in Cell and Molecular Biology (also EEB 215)   Fall ST

Central concepts and experiments in cellular, molecular, and developmental biology with an emphasis on underlying physical and engineering principles. Topics include the genetic code; energetics and cellular organization; communication, feeding, and signaling between cells; feedback loops and cellular organization; problems and solutions in development; the organization of large cellular systems, such as the nervous and immune systems. Satisfies the biology requirement for entrance into medical school. Prerequisites: AP biology, physics, and calculus. Three lectures, one three-hour laboratory. E. Cox, P. Felton, R. Tanny

MOL 231 An Integrated, Quantitative Introduction to the Natural Sciences I (see ISC 231)

MOL 232 An Integrated, Quantitative Introduction to the Natural Sciences I (see ISC 232)

MOL 233 An Integrated, Quantitative Introduction to the Natural Sciences II (see ISC 233)

MOL 234 An Integrated, Quantitative Introduction to the Natural Sciences II (see ISC 234)

MOL 235 An Integrated, Quantitative Introduction to the Natural Sciences III (see ISC 235)

MOL 236 An Integrated, Quantitative Introduction to the Natural Sciences IV (see ISC 236)

MOL 301 Experimental Project Laboratory in Quantitative and Computational Biology (see QCB 301)

MOL 320 Human Genetics, Reproduction, and Public Policy (see WWS 320)

MOL 330 Molecular Evolutionary Genetics (see EEB 320)

MOL 342 Genetics   Fall

Basic principles of genetics illustrated with examples from prokaryote and eukaryote organisms with emphasis on classic genetic techniques. The evolving conception of the gene and genome will be the primary focus of the course. Selected advanced topics will include Drosophila developmental genetics, yeast cell biology, and human disease. Two 90-minute lectures, one class. Prerequisite: 214 or 215, or permission of instructor. G. Schupbach, M. Rose

MOL 345 Biochemistry (also CHM 345)   Fall

Fundamental concepts of biomolecular structure and function will be discussed, with an emphasis on principles of thermodynamics, binding and catalysis. A major portion of the course will focus on metabolism and its logic and regulation. Three lectures, one class. Prerequisite: 214 or 215. Prerequisite or concurrent: CHM 304 and 304B. F. Hughson

MOL 348 Cell and Developmental Biology   Spring

The mechanisms that underlie development of multicellular organisms, from C. elegans to humans, will be examined using biochemical, genetic, and cell biological approaches. The course will investigate the roles that gene regulation, cell-cell communication, cell adhesion, cell motility, signal transduction, and intracellular trafficking play in the commitment, differentiation, and assembly of stem cells into specialized cell types. Two 90-minute lectures, one two-hour class. Prerequisite: 342 or 345. H. Coller, E. Gavis

MOL 350 Laboratory in Molecular Biology   Spring ST

The major objective of the course is to introduce students to a variety of tools required to perform independent research in the field of molecular biology. While conducting original research, students will employ a number of techniques that are used by molecular biologists, molecular geneticists, and biochemists. Students will gain an understanding of how, when, and why certain techniques and skills are used in a research setting. In addition, students will learn to write a research report modeled on the scientific literature. One lecture, two laboratories. Prerequisite: 342 and 345, or either course alone with permission of instructor. R. Tanny

MOL 408 Cellular and Systems Neuroscience (see NEU 408)

MOL 410 Introduction to Biological Dynamics   Fall QR

Designed for students in the biological sciences, this course focuses on the application of mathematical methods to biological problems. Intended to provide a basic grounding in mathematical modeling and data analysis for students who might not have pursued further study in mathematics. Topics include differential equations, linear algebra, difference equations, and probability. Each topic will have a lecture component and computer laboratory component. Students will work extensively with the computing package MATLAB. No previous computing experience necessary. Two 90-minute lectures, one laboratory. N. Wingreen, C. Brody

MOL 414 Genetics of Human Populations (see EEB 414)

MOL 422 Evolutionary Developmental Biology (see EEB 422)

MOL 429 Selected Topics in Molecular Biology and Human Genetics   Spring

An in-depth analysis of one area in which recent advances in molecular biology will have significant impact upon society. One three-hour seminar. Prerequisites: 342 and 348, or 214 and permission of the instructor. Staff

MOL 431 Advanced Topics in Developmental Neurobiology   Fall

Contemporary approaches to the study of neural development, emphasizing genetic and molecular techniques. Topics include generation, patterning, differentiation, migration and survival of neurons and glia, axon growth and guidance, target selection, synapse formation/elimination, activity-dependent remodeling of connectivity, and the relationship between neural development and behavior. Two 90-minute classes. Prerequisites: 342, 348, or permission of instructor. J. Eggenschwiler

MOL 434 Macromolecular Structure and Mechanism in Disease   Not offered this year

This course examines structure-function relationships for a number of proteins involved in human diseases. Topics will include oncogenesis, signal-transduction and apoptosis, as well as protein folding, mis-folding, and trafficking. Classes will involve a mixture of lecture and discussion of original scientific papers, with emphasis given to developing an understanding of how to examine and evaluate primary literature. Prerequisite: 345 or permission of instructor. Two 90-minute lectures. Staff

MOL 435 Pathogenesis and Bacterial Diversity   Not offered this year

An examination of current topics exploring the microbial world with emphasis on signal transduction, and the molecular basis for bacterial diversity and their roles in bacterial pathogenesis. Topics will include the regulation of cell division and sporulation, quorum sensing, mechanisms of microbial differentiation, evolution of communicable diseases, molecular mechanisms of pathogenesis, and identification of virulence factor and immunization. Two lectures, one preceptorial. Prerequisites: 214, 215, or permission of instructor. A. Newton

MOL 437 Computational Neuroscience (see NEU 437)

MOL 448 Chemistry, Structure, and Structure-Function Relations of Nucleic Acids (also CHM 448)   Spring

The chemistry and structure of mononucleotides, oligonucleotides, and polynucleotides and their helical complexes as a basis for understanding and predicting the structures and structure-function relations of naturally occurring DNAs and RNAs. Related functions may include fidelity of DNA replication, mutagenic mechanisms, molecular evolution, telomeres, recently discovered RNA functions, structure of the genetic code. Prerequisite: general chemistry and one semester of organic chemistry. Two 90-minute lectures. J. Fresco

MOL 450 Stem Cells and Cell Fate Decision Processes in the Genomic Era   Not offered this year

Focuses on the current state of stem cell research and the future directions for this field. Stem cell research has great promise for the future of regenerative medicine. Very little is known about the molecular biology that underlies stem cell fate determination. The completion of the human and mouse genome sequences, together with novel technologies to observe global gene expression, offer unique opportunities to unravel stem cell regulatory mechanisms. Explores parallels to other, more mature biological systems. Two lectures, one preceptorial. Prerequisite: 342 and 348, or instructor's permission. Staff

MOL 455 Introduction to Genomics and Computational Molecular Biology (also COS 455)   Fall

Introduction to computational and genomic approaches used to study molecular systems. Topics include computational approaches to sequence similarity and alignment, phylogenetic inference, gene expression analysis, structure prediction, comparative genome analysis, and high-throughput technologies for mapping genetic networks. Two lectures, one preceptorial. M. Singh, S. Tavazoie

MOL 457 Computational Aspects of Molecular Biology   Fall

The applications of computers to research in molecular biology with emphasis on the acquisition of competence in using available tools. Topics include: nucleic acid sequence analysis, secondary structure prediction, sequence homology, the protein folding problem, computer-aided molecular design, and the use of genetic databases. Prerequisites: one 300-level course in molecular biology, chemistry, or biochemistry. Three lectures. J. Welsh

MOL 459 Viruses: Strategy and Tactics   Not offered this year

Viruses are unique parasites of living cells and may be the most abundant, highest evolved life forms on the planet. The general strategies encoded by all known viral genomes are discussed using selected viruses as examples. The course covers the molecular biology (the tactics) inherent in these strategies. It also introduces the biology of engagement of viruses with host defenses, what happens when viral infection leads to disease, vaccines and antiviral drugs, and the evolution of infectious agents and emergence of new viruses. Three lectures, one two-hour preceptorial. Prerequisite: 342 and 348, or instructor's permission. L. Enquist

MOL 460 Diseases in Children: Causes, Costs, and Choices (also STC 460)   Spring

Within a broader context of historical, social, and ethical concerns, a survey of normal childhood development and selected disorders from the perspectives of the physician and the scientist. Emphasis on the complex relationship between genetic and acquired causes of disease, medical practice, social conditions, and cultural values. The course features visits from children with some of the conditions discussed, site visits, and readings from the original medical and scientific literature. Prerequisite: 214 or 215. Two 90-minute classes. D. Notterman