Required Courses


  1. BBS230: Analysis of the Biological Literature Critical analysis of original research articles in intensive small group discussions. Analyze range of papers in biochemistry, genetics, microbiology, and cell and developmental biology in terms of context, hypotheses, methods, results and future experiments.
  2. CB 201:  Molecular Biology of the Cell Molecular basis of cellular compartmentalization, protein trafficking, cytoskeleton dynamics, mitosis, cell locomotion, cell cycle regulation, signal transduction, cell-cell interaction, and cellular/biochemical basis of diseases. Methods covered: mass spectrometry, microscopy, and quantitative approaches to Cell Biology.
  3. BBS330: Critical Thinking and Research Proposal Writing BBS 330 provides a thorough coverage of the essential elements of research proposal writing, a skill that is required throughout one's scientific career but for which formal training is frequently lacking.  After an introductory lecture that provides a general overview of the conceptualization and writing of a research proposal, students will be guided through a series of exercises designed to develop these critical skills
  4. One course in statistics/quantitative biology (see options below)
  5. Two advanced electives in cell biology (see options below)


Students are also encouraged to take selected Nanocourses, which are offered throughout the year. These courses feature typically 3 lecture hours from leaders in the topic under consideration, followed by a second 2 hour period assessment period, involving a student driven project related to the topic under study. The Harvard faculty have developed dozens of Nanocourses, which are given periodically. For more information, see:


Statistics/Quantitative Courses


Neuro 303qc:  Tools for Statistical Inference in Experimental Science
An expanded quarter course providing an introduction to the statistical treatment of experimental data.  It will cover basic topics in statistics, including probability distributions, sampling, hypothesis testing, Baye’s Theorem, t tests, confidence intervals, and ANOVA and related tests.  Discusses the appropriate choice of statistical test, the underlying assumptions, and how to draw appropriate inferences from test results.


Biophysics 170:  Quantitative Genomics
Introduction to quantitative modeling and analysis of genome evolution, functional and structural genomic data. Strong emphasis on evolutionary understanding and analysis.
The course provide foundation in the following four areas: Molecular evolutionary and Population Genetics, Comparative Genomics,  Functional Genomics, and  Structural Genomics.


Courses through the Biostatistics Department at the Harvard School of Public Health may also be applicable.

Advanced Electives


Cell Biology 207. Developmental Biology: Molecular Mechanisms of Vertebrate Development
Course Faculty: Andrew Lassar, Patricia D’Amore, Alan Davidson, John Flanagan, Xi He, Jordan Kreidberg, Richard Maas, Clifford Tabin, Malcolm Whitman  Analyzes the developmental programs of frog, chick, zebrafish, and mouse embryos, emphasizing experimental strategies for understanding the responsible molecular mechanisms that pattern the vertebrate embryo.


Cell Biology 211. Molecular and Systems Level Cancer Cell Biology
Course Faculty: Piotr Sicinski, Jarrod Marto, Marc Vidal  Examines the molecular and systems basis of cancer including alterations in signal transduction, cell cycle, apoptosis and DNA repair.


Cell Biology 212. Molecular Mechanisms of Cancer
Course Faculty: James DeCaprio and Myles Brown  Examines the molecular basis of human cancer, including lung, breast, prostate, melanoma and leukemia. Concepts including stem cells, senescence, genomic instability, angiogenesis, oncogenes, tumor suppressors and viruses in human cancer will be examined.


Cell Biology 214. Developmental Biology and Genetics: Molecular Mechanism of Invertebrate Biology
Course Faculty: David Van Vactor, Spyros Artavanis-Tsakonas, Jarema Malicki The course will explore genetic tools for the analysis of developmental phenomena in flies, worms, and mice. We cover a continuum from pattern formation, cell growth, and cell fate to cell differentiation and morphogenesis.
Cell Biology 226. Concepts in Development, Self-Renewal and Repair
Course Faculty: Laurel Raftery, Alan Davidson, Iain Drummond, Niels Geijsen, N. Nanda Nanthakumar, Lizabeth Perkins, David Scadden Explores developmental mechanisms through the life cycle, contrasting pluripotency and cell fate restriction in embryos and adult tissues. In depth analysis of in vivo approaches, with emphasis on adult stem cells, tissue repair and self-renewal.


Cell Biology XXX: Experimental Approaches to Cell Biology
Course Faculty: David Van Vactor, Wade Harper, and Cell Biology program faculty.  This introductory level course will provide a rapid survey of major topics and themes in cell biology in parallel with hands-on exposure to a variety of experimental approaches, technologies and model systems, including microscopy, proteomics, and biochemical machines.  Registration is limited to eight students; preference given for CB program students.


DRB 330: Experimental Approaches to Developmental Biology
Course Faculty: David Van Vactor, Amy Wagers, and members of the Division  This introductory level course will provide a rapid survey of major topics and themes in developmental biology in parallel with hands-on exposure to a variety of experimental
approaches, technologies and model systems (Drosophilia, C. elegans, Xenopus, chick & mouse).


Quarter Courses

(This list is frequently updated. Check the Nanos and Quarters website for current offerings).


Quantitative Optical Microscopy for Cell Biology
Course Faculty: Tomas Kirchhausen  This five-day total immersion course aims to bring together cell biologists and light microscopy specialists to explore requirements and opportunities offered by advance fluorescence light microscopy techniques applied to the quantitative observation in real-time of single molecules and molecular assemblies in living cells and in isolation. The course will combine lectures with daily hands-on laboratory practices followed by discussion. This advanced imaging course is directed towards students, post-doctoral fellows and faculty with previous working experience using fluorescent light microscopy.

Belief Options for a Practicing Scientist
Course Faculty: Andrius Kazlauskas  While scientific approaches effectively explain the natural world, they fail to resolve moral, ethical and social questions.  These latter issues are part of the scientist’s world and at times even one’s research project.  This quarter course will compare and contrast common belief options to grapple with theological questions that face scientists.  We will read excerpts from a number of authors (such as F. Collins, C.S. Lewis, S. Freud, R. Dawkins, S. Harris) that address how one comes to faith and what that faith can look like.  We will also discuss issues that are typically at the interface between religion and faith such as evolution and the use of stem cells in research.  There will be a total of 6 meetings that will consist of student-led discussion based on the types of readings described above.  While there will not be any exams, successful completion of the course will include writing a final paper/thesis making a case for which of the belief options is best suited for a scientist. 

Histology for Graduate Students
Course Faculty: Gerald Greenhouse and Adrian Salic  A 6-session quarter course for graduate students on histology--the study of structure, and therefore function, in cells and tissues. The class will include a session on each of the major tissue types in mammals--connective, muscle, nerve and epithelial. Each session will include an introductory lecture followed by shared observation of slides using the medical school’s 12-headed light microscope. In the last session, students will do tissue staining with recently developed compounds in the lab of Adrian Salic. This class is recommended for any graduate student whose thesis work will benefit from a strong working knowledge of cell structure and tissue architecture. Students examining cell changes in carcinogenesis, development or regeneration and those characterizing mammalian phenotypes are especially encouraged to participate. Class size is limited to 11.
Experimental Design for Biologists
Course Faculty: David Glass and Randall King  This course will focus on both the theory and practice of experimental design, asking when, how, and whether hypotheses or questions should be used to frame experiments, and how these frameworks may perturb experimental design and interpretation.  A thorough discussion of the various types of experimental controls will be provided.  This will go beyond a standard exposition of negative and positive controls, and will include elements like system design, system controls, controls for underlying assumptions, experimentalist controls. The students will be asked to read some philosophy, a few chapters from Dr. Glass' book on Experimental Design, and then to apply what was learned to their own projects, by giving presentations on their own experimental designs, and flow-charts of their projects.  This is not a statistics course - rather it covers those elements of design that usually precede and follow statistical analysis.


Stem Cells: Properties and Applications
Course Faculty: George Q. Daley and Amy Wagers  This quarter course will discuss scientific problems in both ES and adult stem cell research, providing an historical context for stem cell research as well as a discussion of the basic biological properties and applications of these cells and the questions and controversies that currently drive the field.  We will conclude with a discussion of the societal and ethical implications of stem cell research and the impact of federal guidelines on research.  Students will be responsible for reading and critically discussing 3-5 relevant papers each week.  Papers for discussion will be taken from both classic and current literature. Class participation is required and enrollment is limited to 16 students.


Advanced Topics in Cell Biology: CB309qc
Course Faculty: David Van Vactor and Cell Biology Faculty (eight meetings per semester).  This quarter course covers cutting edge research across a broad range of topics from angstrom resolution to organismal physiology, from basic mechanisms underlying fundamental cellular processes to the molecular etiology of human disease.  The course is designed to provide students with intimate intellectual access to our faculty in a setting where all questions are welcome, where new experimental ideas can be explored, and where the horizon-line of each field can be considered.  One primary research article and one review article will be identified by each lecturer; follow up reading will also be recommended for students that wish to deepen their knowledge in a given topic.  Registration limited to eight students; preference offered to CB program students.



Nanocourses are a relatively new course format designed to teach specific subjects at an advanced level and in a condensed format. These courses are aimed at bringing students, post-docs and faculty up to date on a particular field, to provide insight into the current problems in that field, and to define a solid basis for further study in that field, all within a short time period. Each course meets for a minimum of 6 hours over 2 days. The first session is lecture-based and taught by 1 to 3 faculty who present an advanced level of knowledge on current research areas, specific experimental approaches and new technologies. The second-session is discussion-based or hands-on and gives students an opportunity to explore the topic in depth in a small-group setting.

The cell biology training program offers a variety of nanocourses each semester. Check the Nanos and Quarters website for current course offerings. (link to )

Microscopy Education

The Nikon Imaging Center (NIC) is a core facility affiliated with the Cell Biology, Systems Biology, and Biochemistry and Molecular Pharmacology Departments. Trainees in the cell biology program are invited to participate in a number of workshop courses offered by the NIC each year to gain experience in basic and advanced light microscopy techniques.

Additional BBS Core Courses

BCMP 200:  Molecular Biology
An advanced treatment of molecular biology’s Central Dogma. Considers the molecular basis of information transfer from DNA to RNA to protein, using examples from eukaryotic and prokaryotic systems. Lectures, discussion groups, and research seminars.

BCMP 201:  Proteins:  Structure, Function, and Catalysis
Protein biochemistry with emphasis on the interrelated roles of protein structure, catalytic activity, and macromolecular interactions in biological processes. Course provides the core background and the perspective required to consider and dissect biological problems at a mechanistic, molecular level.
Genetics 201: Principles of Genetics 
An in-depth survey of genetics, beginning with basic principles and extending to modern approaches and special topics. We will draw on examples from various systems, including yeast, Drosophila, C. elegans, mouse, human and bacteria.

Micro 201: Molecular Biology of the Bacterial Cell
This course is devoted to bacterial structure, physiology, genetics, and regulatory mechanisms. The class consists of lectures and group discussions emphasizing methods, results, and interpretations of classic and contemporary literature.





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