Biology, Molecular and Cellular
Biology, Molecular and Cellular
The Department of Molecular and Cellular Biology (MCB) is home to an unusually diverse group of outstanding scientists. The MCB 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 MCB 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 department’s faculty and labs; research facilities and resources; admissions procedures; two PhD training programs and their degree requirements; current graduate students and graduate student life, please visit our website at http://www.mcb.harvard.edu/mcb/p/graduate-programs/ or write directly to the Graduate Office at
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or call (617) 495-3293.
Training Programs
The Department of Molecular and Cellular Biology offers two interdisciplinary training programs in the life sciences which lead to a PhD in either Biology or Biochemistry: (1) The Molecules, Cells, and Organisms (MCO) Training Program and (2) The Engineering and Physical Biology (EPB) Training Program. While all graduate students are admitted as Ph.D. degree candidates, the A.M. degree may be conferred as a non-terminal degree to mark the completion of the candidacy requirements following the Candidacy Examination.
Both the MCO and EPB training programs take 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.
Molecules, Cells and Organisms (MCO). Catherine Dulac, Program Director. Faculty participating in the Molecules, Cells, and Organisms training program come from the Departments of Molecular and Cellular Biology, Organismic and Evolutionary Biology, Chemistry and Chemical Biology, and Stem Cell and Regenerative Biology. In addition, members of the Center for Systems Biology, the Center for Brain Science, the Microbial Science Initiative, and the Harvard Stem Cell Institute are active participants in the MCO training program.
Foundational coursework in the first year prepares students for research in one of three tracks: Physical, Chemical, and Molecular Biology; Cellular, Neuro-, and Developmental Biology; and Genetics, Genomics, and Evolutionary 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 three 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.
Engineering and Physical Biology Training Program (EPB). Nancy Kleckner, Program Director. Offered in partnership with the Department of Physics and the School of Engineering and Applied Sciences, EPB trains a new generation of scientists to view living systems through the lens of physics and engineering. EPB 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. Activities take place in the intellectual community of the daVinci Group, an interdepartmental community devoted to the analysis of living systems through the lens of physics and engineering. Students accepted into the EPB training program may apply through the Department of Molecular and Cellular Biology, The Department of Physics, or the School of Engineering and Applied Sciences.
Admission Requirements and Undergraduate Preparation
Applications for admission 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 present the scores with their application. Applications without GRE scores will be considered incomplete. Students are also strongly encouraged to take a subject exam such as Biology or Biochemistry.
Online submission of the application is required and completed applications with all supporting materials, including letters of recommendation, are due online by the announced deadline to ensure consideration for the following fall. Late applications will not be considered. Well-qualified candidates are invited to campus by the department’s admissions committee in early 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 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 will not be provided beyond the fifth year without prior agreement of the student’s advisor and the MCB 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; 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.
Coursework.
MCO Training Program: First year graduate students in the MCO training program enroll in MCB 290hfr (fall and spring) and MCB, 291, 292 and 293 in the fall term. In the spring term, 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 may continue to take elective coursework in their second year. Second year MCO graduate students should also enroll in MCB 290hfr for the fall and spring of the G2 year. In addition to the required courses and electives, first-year students also enroll in MCB 300 in the fall and spring semesters of the first year (see Laboratory Rotations below).
EPB Training Program: During the first year, EPB graduate students enroll in two required courses, which include MCB 225 and ES 224, in order to explore different areas of project-based research. Additional electives are taken in consultation, giving EPB students opportunities to explore and develop diverse areas of expertise. Students choose additional elective coursework in consultation with their faculty advisor and the program director to explore and develop diverse areas of expertise. In addition to two required courses and electives, students also enroll in MCB 300 in the fall and spring semesters of the first year (see Laboratory Rotations below).
Laboratory Rotations. Students in both training programs spend their first year performing experimental research in the laboratories of faculty members. During the eight-week laboratory 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. 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.
MCO Journal Club. While reading and discussing scientific papers is an integral part of the curriculum of many courses in the MCO training program, the Journal Club provides a forum for students in the first two years of their 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 (i.e. what people were thinking before and did after this change afterwards.) These talks are advertised and open to members of the greater Life Sciences community, and an MCO faculty member may also present along with a student in any given week. Talks are held once per week 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. Journal Clubs give students a chance to speak in front of an audience of peers (without the aid of a PowerPoint presentation). 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 are encouraged to form their own Journal Club(s) which can be organized by research topic, the 3 MCO tracks, by class year, or however students would like.
After the First Year
Acceptance for Candidacy. MCB 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-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 thesis 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 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 made to the department and community as a whole; 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 and the Graduate Programs Office at least two weeks in advance of the dissertation defense. Electronic copies may be submitted. 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, students should submit one bound copy of the completed dissertation to the Graduate Office and one bound and one unbound copy to the Registrar, along with the other required exit surveys, fee for publication, and a signed Dissertation Acceptance Certificate. Detailed requirements on the dissertation are published in The Form of the PhD Dissertation, which is available online or in the Graduate Office.
Teaching. All graduate students in the Department of Molecular and Cellular Biology are required to teach at least 2 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 large 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 MCB Dissertation Titles (2011-12)
Banse, Stephen. Advisor: Craig Hunter. “Study of Chromatin Structure using Stimulated Raman Scattering Microscopy in Living Mammalian Cells”
Basu, Srinjan. Advisor: Xialong Xie. “Study of Chromatin Structure using Stimulated Raman Scattering Microscopy in Living Mammalian Cells”
Boulting, Gabriella. Advisor: Kevin Eggan. “Exploring the potential of human pluripotent stem cells as a platform for new ALS disease models”
Carter-O'Connell, Ian. Advisor: Erin O’Shea. “Structural Analysis of the CDK-Cyclin Complex of Pho85-Pho80 and Genome-Wide Characterization of the Phosphate Starvation Response in Schizosaccharomyces pombe”
Chand, Nikhilesh. Advisor: Deborah Hung. “The Two-component Sensor KinB Regulates Pseudomonas aeruginosa Virulence”
Chen, Huiyi. Advisor: Xiaolong Xie. “System-Wide Studies of Gene Expression in Escherichia coli by Fluorescence Microscopy and High Throughput Sequencing”
Cook, Kristen E. Advisor: Erin O’Shea. “Regulation of genome-wide transcriptional stress responses in Saccharomyces cerevisiae”
Daniele, Joseph. Advisor: Sam Kunes. “A Novel Proteolytic Event Controls Hedgehog Intracellular
Sorting and Transport”
Fomina Yadlin, Dina. Advisor: Stuart Schreiber. “Small molecule inducers of insulin expression in pancreatic alpha cells”
Kao, Robert. Advisor: Andrew McMahon. “Investigation of the molecular and cellular basis of patterning, morphogenesis, and tubule interconnections during mammalian kidney development”
Malca, Hadar. Advisor: Nicole Francis. “The fate of parental histones during DNA replication in vitro”
Mekhoubad, Shila. Advisor: Kevin Eggan. “Dynamics of X-Chromosome Inactivation During Human Pluripotent Stem Cell Derivation, Maintenance, and Differentiation”
Miller, Sarah B. Advisor: Victoria D’Souza. “Structural and biochemical studies of retroviral reverse transcription initiation”
Mitchell, Andrew. Advisor: Alan Saghatelian. “Discovering Bioactive Peptides and Characterizing the Molecular Pathways that Control their Activity”
Morimoto, Emiko. Advisor: Tom Maniatis. “The role of microglia in amyotrophic lateral sclerosis: Analysis of microRNAs”
Murphy, Christopher. Advisor: Matthew Michael. “Control of DNA replication by the nucleus to cytoplasm ratio”
Rizvi, Abbas H. Advisor: Erin O’Shea. “Systems level studies of nutrient homeostasis”
Rosains, Jacqueline. Advisor: Susan Mango. “Modulation of pha-4 and C. elegans foregut development by the novel gene smg-8”
Son, Esther. Advisor: Kevin Eggan. “Exploring the Plasticity of Cellular Fate Using Defined- Factor Reprogramming”
Stanton, Shaunna. Advisor: Gregory Verdine. “An Oligonucleotide Sequence Targeting the CR4-CR5 Region of the Human Telomerase RNA”
Tan, Kah Yong. Advisor: Amy Wagers. “Stem cell-based strategies to enhance muscle regeneration
through extrinsic and intrinsic regulators”
Veguilla, Rosa. Advisor: David Fisher. “Transcriptional Regulation of OCA2 and POMC by a cAMP-dependent mechanism and implications in skin pigmentation”
Wu, John. Advisor: Xiaowei Zhuang. “Singlemolecule and super-resolution fluorescence studies of the structure and function of telomerase and telomere”
Zhang, Xiaodong. Advisor: Yun Zhang. “Molecular and behavioral mechanisms of aversive olfactory learning in C. elegans”

