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Committee on Development, Regeneration, and Stem Cell Biology

This is an archived copy of the 2012-13 catalog. To access the most recent version of the catalog, please visit


  • Edwin L. Ferguson, Molecular Genetics & Cell Biology


  • John Cunningham, Pediatrics
  • Glyn Dawson, Pediatrics
  • Wei Du, Ben May Institute for Cancer Research
  • Richard Fehon, Molecular Genetics & Cell Biology
  • Edwin L. Ferguson, Molecular Genetics & Cell Biology
  • Michael Glotzer, Molecular Genetics and Cell Biology
  • Elizabeth Grove, Neurobiology
  • Robert Haselkorn, Molecular Genetics & Cell Biology
  • Robert K. Ho, Organismal Biology & Anatomy
  • Bruce Lahn, Human Genetics
  • Elizabeth McNally, Medicine
  • Victoria E. Prince, Organismal Biology & Anatomy
  • Ilaria Rebay, Ben May Institute for Cancer Research
  • Marsha Rosner, Ben May Institute for Cancer Research
  • Nancy B. Schwartz, Pediatrics
  • Neil H. Shubin, Organismal Biology & Anatomy
  • Kevin White, Human Genetics

Associate Professors

  • Yoav Gilad, Human Genetics
  • William Green, Neurobiology
  • Akira Imamoto, Ben May Institute for Cancer Research
  • Barbara Kee, Pathology
  • David Kovar, Molecular Genetics & Cell Biology
  • Kay MacLeod, Ben May Institute for Cancer Research
  • Jocelyn Malamy, Molecular Genetics & Cell Biology
  • Clifton Ragsdale, Neurobiology
  • Ilya Ruvinsky, Ecology & Evolution
  • Urs Schmidt Ott, Organismal Biology & Anatomy
  • Kamal Sharma, Neurobiology
  • Eric C. Svensson, Medicine

Assistant Professors

  • Jill de Jong, Pediatrics
  • James Holaska, Medicine
  • Sally Horne-Badovinac, Molecular Genetics and Cell Biology
  • Ivan Moskowitz, Pediatrics
  • Ed Munro, Molecular Genetics & Cell Biology
  • Xiaoyang Wu, Ben May Institute for Cancer Research

Emeritus Faculty

  • Martin Gross, Pathology
  • Anthony Mahowald, Molecular Genetics & Cell Biology
  • Manfred D.E. Ruddat, Ecology & Evolution

Program of Study

First Year

The first year of graduate study is spent in coursework, independent reading, and exploratory research. The number of courses constituting a full schedule for each quarter of the first year will vary, but typically includes three lecture courses or two lecture courses and a research rotation. Students are required to undertake laboratory rotations in at least two different laboratories before beginning their dissertation research. Three rotations are encouraged. These rotations can be performed during the first academic year or during the Summer Quarter.

Seminars given by invited speakers are regularly offered and students are strongly urged to attend. A separate series of meetings is presented in the fall and winter quarters by faculty to introduce students to their research. Before beginning their second year, students complete Part I of the candidacy examinations, which consists of an oral examination covering the core courses in developmental, cell, and molecular biology, and genetics.

Second year

While coursework can continue during the second year, students spend much of their time developing a research project. Students have generally chosen research advisors at the beginning of the second year. By the end of the Winter Quarter of the second year, each student’s doctoral committee is named. The student then prepares a written proposal for dissertation research and defends this proposal before the doctoral committee. This defense constitutes Part II of the candidacy examination. This examination must be completed by the end of the Spring Quarter of the second academic year.

Advanced years

After the qualifying exam, the student works full time on thesis research, although the faculty urges students to continue to take advantage of the advanced courses and seminars that are offered. Finally, each graduating student writes a dissertation describing his or her research, presents the work in a public seminar, and defends it before their doctoral committee.


Throughout their term as graduate students, students are expected to have frequent informal conversations with professors in their courses, their research advisor, and members of their doctoral committees. In this way, students can obtain frequent appraisals of their progress and constructive advice.

Formal evaluation of each student’s progress continues every academic year. In the first and second years, the evaluation is based on the student’s performance in courses, laboratory rotations and the qualifying examination. In later years, the research advisor and doctoral committee oversee the student’s dissertation research progress; a report is submitted after the yearly meeting that becomes part of the student’s permanent file and is reviewed by the Curriculum Committee. If the committee is apprised of any deficiencies in performance, the student will receive a letter describing those deficiencies and making suggestions about how to remedy them.


For information about applying to our graduate program, please visit our website at .

Requirements for the Ph.D. Degree

A Ph.D. candidate must fulfill certain formal course work requirements, pass the qualifying examination, and present a satisfactory dissertation describing the results of original research.

The committee expects a knowledge of and proficiency in contemporary developmental biology as well as auxiliary fields of molecular biology, cell biology, and genetics. This requirement will normally be met by fulfilling the formal course work listed below. However, courses taken at other institutions, in other departments, or as part of the medical school curriculum may substitute for required committee courses with the approval of the curriculum committee.

Formal Course Work

The Division of the Biological Sciences requirement of nine graded course units may be met by registering for a combination of formal courses and research credits. During the first year of graduate work students ordinarily complete one course in molecular biology, one in cell biology, one in genetics, and three courses in developmental biology.

Developmental Biology Courses

DVBI 35600. Vertebrate Developmental Biology. 100 Units.

This advanced-level course combines lectures, student presentations, and discussion sessions.  It covers major topics on the developmental biology of embryos (e.g. formation of the germ line, gastrulation, segmentation, nervous system development, limb pattering, organogenesis).  We make extensive use of the primary literature and emphasize experimental approaches (e.g. classical embryology, genetics, molecular genetics).

Instructor(s): V. Prince, K. Sharma     Terms Offered: Spring
Prerequisite(s): BIOS 20180s or 20190, or AP 5 sequence
Equivalent Course(s): BIOS 21356,MGCB 35600

DVBI 36100. Plant Development and Molecular Genetics. 100 Units.

Genetic approaches to central problems in plant development will be discussed.  Emphasis will be placed on embryonic pattern formation, meristem structure and function, reproduction, and the role of hormones and environmental signals in development.  Lectures will be drawn from the current literature; experimental approaches (genetic, cell biological, biochemical) used to discern developmental mechanisms will be emphasized. Graduate students will present a research proposal in oral and written form; undergraduate students will present and analyze data from the primary literature, and will be responsible for a final paper. 

Instructor(s): J. Greenberg     Terms Offered: Spring
Prerequisite(s): Completion of the general education requirement in the biological sciences
Equivalent Course(s): BIOS 23299,ECEV 32900,MGCB 36100

DVBI 36400. Developmental Mechanisms. 100 Units.

This course provides an overview of the fundamental questions of developmental biology, with particular emphasis on the genetic, molecular and cell biological experiments that have been employed to reach mechanistic answers to these questions.  Topics covered will include formation of the primary body axes, the role of local signaling interactions in regulating cell fate and proliferation, the cellular basis of morphogenesis, and stem cells.  The discussion section covers selected papers from the literature and focuses on the critical evaluation of experimental evidence.

Instructor(s): E. Ferguson, R. Fehon     Terms Offered: Winter
Prerequisite(s): BIOS 20182, 20192, 20187, or 20235
Equivalent Course(s): BIOS 21237,MGCB 36400

 Distribution Courses

DVBI 31200. Molecular Biology-I. 100 Units.

Nucleic acid structure and DNA topology; methodology; nucleic-acid protein interactions; mechanisms and regulation of transcription in eubacteria, and of replication in eubacteria and eukaryotes; mechanisms of genome and plasmid segregation in eubacteria.

Instructor(s): L. Rothman-Denes     Terms Offered: Winter Quarter
Equivalent Course(s): MGCB 31200,BCMB 31200

DVBI 31300. Molecular Biology-II. 100 Units.

The content of this course covers the mechanisms and regulation of eukaryotic gene expression at the transcriptional and post-transcriptional levels. Our goal is to explore research frontiers and evolving methodologies. Rather than focusing on the elemental aspects of a topic, the lectures and discussions highlight the most significant recent developments, their implications and future directions.

Instructor(s): J. Staley, J. Holaska, A. Ruthenburg     Terms Offered: Spring Quarter
Equivalent Course(s): MGCB 31300,BCMB 31300

DVBI 31400. Genetic Analysis of Model Organisms. 100 Units.

Fundamental principles of genetics discussed in the context of current approaches to mapping and functional characterization of genes.  The relative strengths and weaknesses of leading model organisms are emphasized via problem-solving and critical reading of original literature.

Instructor(s): A. Palmer, D. Bishop, E. Ferguson, J. Malamy     Terms Offered: Autumn
Equivalent Course(s): BCMB 31400,HGEN 31400,MGCB 31400

DVBI 31500. Genetic Mechanisms. 100 Units.

Advanced coverage of mechanisms involved in promoting genome stability and genome evolution.  A variety of experimental systems are explored from bacteriophage to humans.  Topics include the genetics and biochemistry of DNA repair, homologous and site-specific recombination, transposition and genome rearrangement.  Two of three weekly meetings are lecture and the third student led discussion of recent papers from the primary literature. The course emphasizes experimental design and interpretation of primary data.

Instructor(s): D. Bishop     Terms Offered: Spring Quarter
Equivalent Course(s): MGCB 31500

DVBI 31600. Cell Biology I. 100 Units.

Eukaryotic protein traffic and related topics, including molecular motors and cytoskeletal dynamics, organelle architecture and biogenesis, protein translocation and sorting, compartmentalization in the secretory pathway, endocytosis and exocytosis,and mechanisms and regulation of membrane fusion.

Instructor(s): A. Turkewitz, B. Glick     Terms Offered: Autumn Quarter
Equivalent Course(s): MGCB 31600,BCMB 31600

DVBI 31700. Cell Biology II. 100 Units.

This course covers the mechanisms with which cells execute fundamental behaviors. Topics include signal transduction, cell cycle progression, cell growth, cell death, cancer biology, cytoskeletal polymers and motors, cell motility, cytoskeletal diseases, and cell polarity. Each lecture will conclude with a dissection of primary literature with input from the students. Students will write and present a short research proposal, providing excellent preparation for preliminary exams.

Instructor(s): M. Glotzer, D. Kovar     Terms Offered: Winter
Equivalent Course(s): MGCB 31700