The Department of Molecular and Cellular Biology (MCB) is the host department of a multi-departmental Ph.D. Training Program called Molecules, Cells and Organisms (MCO). The program contains an unusually diverse group of outstanding scientists, and MCO’s unique approach to the Biological Sciences PhD cultivates versatility as well as depth of expertise by exposing its students to the full spectrum of modern biology. The program’s mission to advance biological research beyond traditional boundaries is motivated by a passion for discovery and is supported by innovative research centers and state-of-the-art facilities on Harvard’s Cambridge campus. It is this interdisciplinary and collaborative culture—motivated by a passion for scientific discovery that makes the MCO Program an exciting place to study the unsolved questions in biology. Graduate students are trained to be the next generation of life scientists: creative, independent, and productive researchers working in academia, medicine, industry, law, business, or the non-profit sector.

To learn more about the training program faculty and labs; research facilities and resources; admissions procedures; PhD training program and its degree requirements current graduate students and graduate student life, please visit our website at or write directly to Michael Lawrence in the Graduate Office at This email address is being protected from spambots. You need JavaScript enabled to view it. or call (617) 495-3293.

Training Programs

Molecules, Cells and Organisms (MCO)

Catherine Dulac, Training Program Director

MCO is a PhD program, although it is possible to receive the AM degree to signify the completion of requirements following the Candidacy Examination. The MCO program takes full advantage of the university’s outstanding faculty and extensive laboratory resources to provide pre-doctoral students with a solid foundation in the concepts and scientific approaches used in laboratories today to prepare them for a future at the forefront of life sciences.

Faculty participating in the MCO Training Program come from the Departments of Molecular and Cellular Biology, Organismic and Evolutionary Biology, Chemistry and Chemical Biology, Stem Cell and Regenerative Biology, and Physics. In addition, members of the FAS Center for Systems Biology, the Center for Brain Science, the Microbial Science Initiative, and the Harvard Stem Cell Institute are active participants in MCO.

Foundational coursework in the first year prepares students for research in one of four tracks: Genetics, Genomics, and Evolutionary Biology; Cellular, Neuro-, and Developmental Biology; Biochemistry, Chemical and Structural Biology; and Engineering and Physical Biology. MCO trainees spend the first year exploring a broad sweep of fundamental problems at every level through a set of core courses representing the program tracks, followed by deep immersion in focused areas. The objective of the MCO training program is to prepare students for a future in science that will require interdisciplinary breadth, as well as depth in specific disciplines.

The EPB track trains a new generation of scientists to view living systems through the lens of physics and engineering. Students work comfortably in both the life sciences and the physical sciences, and applicants may have their primary undergraduate training in either area. Program components combine flexibility with rigor, place a priority on independence and imagination, and emphasize extensive individual faculty-student interactions.

Admission Requirements and Undergraduate Preparation

Applications for admission to the MCO PhD program are accepted from students who have received a bachelor’s degree or equivalent training. Entering students should have a record of introductory courses in chemistry, biology, physics, and mathematics. While the following courses should not be regarded as prerequisites for admission to graduate study, most admitted students have completed these courses as undergraduates:

1. Biology (at least one general course in biology and two terms of biology at a more advanced level)
2. Biochemistry
3. Organic Chemistry
4. Physical Chemistry
5. Physics (a general course in physics)
6. Mathematics (a basic knowledge of differential and integral calculus). Competence in elementary programming is also desirable.
7. Laboratory in Biology, Biochemistry, or Instrumental Analysis.

Applicants are required to take the General Graduate Record Examination (GRE) and to include these scores with their application. Students are also encouraged to take a subject exam such as Biology or Biochemistry, although the subject test is not a requirement for the application. Applications without the general GRE test scores will be considered incomplete and will not be reviewed.

Online submission of the application is required and completed applications with all supporting materials, including 3 letters of recommendation, are due online by the announced deadline to ensure consideration for the following fall. Late or incomplete applications will not be considered. Well-qualified candidates are invited to campus (or interviewed by telephone or Skype if overseas) by the department’s admissions committee in January and February. These visits bring potential candidates to campus for 2–3 days to meet with both faculty and students.

Financial Support

The department of Molecular and Cellular Biology and Program in Molecules, Cells and Organisms guarantees full financial support for five years to all PhD candidates while they are making satisfactory progress toward the PhD degree. Students are expected to complete graduate work to obtain the PhD within five years. Ordinarily, financial support is only provided beyond the fifth year with prior agreement of the student’s advisor and Graduate Committee.

Prospective students are encouraged to apply for outside funding from agencies such as the National Science Foundation at the time of their application and international students should apply for outside funding before coming to the United States.

Degree Requirements

The First Year

Academic Residence

The Graduate School of Arts and Sciences requires a minimum of two years of full-time study in residence. The Graduate School of Arts and Sciences Handbook describes the regulations and rules that apply to students in the Graduate School of Arts and Sciences.


The standard course load for all MCO first year graduate students is five courses in each of the fall and spring
semesters.  In both semesters, all students enroll in MCB290hfr
(Graduate Student Journal Club) and MCB300 (Laboratory Rotation).  In the fall term, students in alll tracks except for Engineering and Physical Biology (EPB) enroll in MCB 291, 292 and 293.  Students in the EPB track enroll in MCB
294 plus two additional courses that are decided in consultation with the EPB Track Leaders.  These additional courses may include foundation MCO courses and/or other courses as deemed appropriate to the intellectual goals of the student.

In the spring term, in addition to MCB 290hfr and MCB 300, each student enrolls in a quantitative methods course (ordinarily MCB 111) along with two elective courses selected from their chosen track, in consultation with their advisor or track head. Students with
advanced quantitative skills can substitute another course with a mathematical or computational focus for MCB111.

Students may continue to take elective
coursework in their second year.

Laboratory Rotations. Students spend their first year performing experimental research in the laboratories of faculty members in at least three 8-week lab rotations. During the rotations, students interact with individual faculty members and explore possible subjects for future dissertation research. The laboratory rotations do not coincide with the semester start and end dates, but all first year students should register for MCB 300 in the fall and again in the spring to indicate the laboratory rotation course. Some students choose to carry out an additional rotation during the summer preceding their first year, or, if they have not decided upon a home lab following the spring term, may opt for an additional rotation in May and June of the first year. Each student arranges for a permanent faculty dissertation advisor and begins dissertation research by the end of the first year.

Outside Fellowship Application. All prospective students are encouraged to apply for outside funding from agencies such as the National Science Foundation at the time of application to the program. If a student has not procured a fellowship upon admission, first year students are asked to submit a research proposal to a nationally recognized funding agency. International students are asked to apply for funding opportunities from their home country at the time of application for admission, as many foreign fellowships must be procured before the student matriculates in the United States. A fellowship writing workshop is conducted early every fall to aid students in how to put together a compelling proposal.

Ethics Workshop in the Responsible Conduct of Research (RCR). In addition to academic coursework, all MCB PhD candidates must complete a workshop in the responsible conduct of research by the end of the first year of study. The workshop is sponsored and conducted by members of the faculty. Instruction in RCR must be undertaken at least once during each career stage, and at a frequency of no less than once every four years, so students in the fifth year are required to enroll in a refresher course in the responsible conduct of research.

MCO Student-Faculty Journal Club. While reading and discussing scientific papers is an integral part of the curriculum of many courses, the Journal Club provides a forum for students in the first two years of the PhD program to be coached on presenting papers in a way that should engage even a non-specialist. The coaching trains students how to contextualize the findings of a previously published paper in relation to their own place in history. The weekly talks are advertised and open to members of the Life Sciences community, and an MCO faculty member may also present along with a student in any given week. Talks are held weekly during each semester, and each year at least one G1 and G2 will have the opportunity to be coached, engage in a Journal Club presentation, and receive follow-up feedback. For the first month of the fall semester, G1s attend presentations by G2s and faculty, and are thereafter incorporated into the weekly schedule. Following the G2 year, students form their own Journal or Data Club(s) which can be organized by research topic, class year, or however students would like.

After the First Year

Acceptance for Candidacy

MCO students are evaluated in the spring of their second year by a faculty exam committee that meets with students to discuss their dissertation proposal. The Candidacy (Qualifying) Examination demonstrates a student’s qualifications for advanced research. Typically it is a one- to two-hour presentation of the dissertation research proposal made to members of an Examination Committee, which is chosen by the student in consultation with the dissertation advisor. In addition, students may be examined on course work, readings, and other required knowledge in the field.

Progress Meetings/Reports

Students accepted for candidacy arrange to meet at least once annually with their dissertation advisory committee (DAC). At these progress meetings, students should summarize the status of their dissertation research, detailing their accomplishments for the past year and goals for the coming year and the period until completion. The progress reports ensure that students, their advisors, and the advisory committee have the same understanding of students’ progress toward the PhD degree.

Dissertation Defense

Four to five years of full-time research is usually required for completion of the PhD degree. Completed research is presented for approval as a written dissertation. Granting of the degree requires the approval of a faculty advisory committee that reviews the dissertation on its contents. The candidate will also be called upon to demonstrate the ability to formulate and defend original ideas on scientific topics not directly related to the subject of the dissertation.

The dissertation defense is comprised of two components: the first is a public presentation; the second is a private defense and examination before the student’s dissertation advisory committee.

The candidate must provide copies of the completed (unbound) dissertation to members of their committee at least two weeks in advance of the dissertation defense. Electronic copies may be submitted upon committee approval. The dissertation should include an abstract of not more than 350 words, stating the purpose, main results and conclusions of the dissertation research. Upon successful completion of the public and private defense, the student will receive a signed Dissertation Acceptance Certificate. Students should submit an electronic copy of the dissertation, which must include a copy of the signed Dissertation Acceptance Certificate to the Registrar as well as along with the other required exit surveys, fee for publication, and a signed original hard-copy Dissertation Acceptance Certificate.The Dissertation Acceptance Certificate may be obtained from the Graduate Office following successful completion of the dissertation. Detailed requirements on the dissertation are published in The Form of the PhD Dissertation.A copy of the dissertation should be submitted to the MCO Graduate Programs Office upon final completion


All graduate students in the Department of Molecular and Cellular Biology and Training Program in Molecules, Cells and Organisms are required to teach at least two classes during their time in the PhD program. This requirement must be completed by the end of the G3 year. The intent is to make sure students receive two kinds of experience in teaching: A introductory undergraduate course with a good amount of independent section teaching and another course that is more discussion-based, containing more advanced material. Typically, students will teach one class in the fall of the G2 year and one class during the fall or spring of the G3 year, but it is up to the student in which semesters the requirement is fulfilled.

Recent Dissertation Titles (2014-2015)

  • November 2014

    Ashrafi, Ghazaleh.  Advisor:  Tom Schwarz, Harvard Medical School.  "Trafficking and Turnover in Neuronal Axons"

    Hsu, Patrick.  Advisor:  Feng Zhang (Broad Institute) and Xiaowei Zhuang (Chemistry and Chemical Biology).  "Development of the CRISPR nuclease Cas9 for high precision mammalian genome engineering"

    Oak, Youbean.  Advisor:  Gaudenz Danuser, Harvard Medical School.  "Filopodia-independent roles of the actin bundling protein fascin in promoting cell motility"

    Sandoe, Jackson.  Advisor:  Kevin Eggan (Stem Cell and Regenerative Biology).  "Developing Human Stem Cell Derived Motor Neuron Models of Amyotrophic Lateral Sclerosis"

    March 2015

    Liu, Jessica.  Advisor:  Timur Yusufzai, Harvard Medical School.  "Biochemical Characterization of the Domain Architecture of Chromatin Assembly Motor Proteins Human CHD1 and CHD2"

    Rogers, Katherine.  Advisor:  Alex Schier (Molecular and Cellular Biology).  "Mesendoderm patterning by the Nodal/Lefty activator/inhibitor system"

    May 2015

    Chew, Guo-Liang.  Advisor:  Alex Schier (Molecular and Cellular Biology).  "Non-canonical translation in vertebrates"

    Grama, Abhinav.  Advisor:  Florian Engert (Molecular and Cellular Biology).  "Motion processing in the larval zebrafish tectum"

    Hacisuleyman, Fatma Ezgi.  Advisor:  John Rinn (Stem Cell and Regenerative Biology).  "The discovery and characterization of the lncRNA Firre"

    Lacoste, Alix.  Advisor:  Alex Schier (Molecular and Cellular Biology).  "Convergent excitatory pathways mediate the zebrafish escape behavior"

    Leiman, Sara.  Advisor:  Richard Losick (Molecular and Cellular Biology).  "Genetics and Regulation of Bacterial Biofilms"

    Perez, Julio.  Advisor:  Catherine Dulac (Molecular and Cellular Biology).  "Genomic Imprinting in the Brain: the Persistent Influences from Mom and Dad"

    Yilmaz Balban, Melis.  Advisor:  Markus Meister (Molecular and Cellular Biology / California Institute of Technology).  "Roles of retinal circuits in the innate visual behaviors of mice"
    May 2015

Faculty List