Molecular, Cellular, and Developmental Biology
Kline Biology Tower, 432.3538
M.S., Ph.D.
Chair
Michael Snyder
Director of Graduate Studies
Ronald Breaker (708 KBT, 432.9389, ronald.breaker@yale.edu)
Professors
Sidney Altman, Kim Bottomly (Immunology), John Carlson, Stephen
Dellaporta, Xing-Wang Deng, Mary Helen Goldsmith, Douglas
Kankel, Michael Kashgarian (Pathology), Haig Keshishian, Perry
Miller (Anesthesiology), Mark Mooseker, Jon Morrow (Pathology),
Frederick Naftolin (Obstetrics & Gynecology), Timothy
Nelson, L. Nicholas Ornston, Thomas Pollard, Shirleen Roeder,
Joel Rosenbaum, Alanna Schepartz (Chemistry), Steven Segal
(Physiology), Michael Snyder, Robert Wyman
Associate Professors
Ronald Breaker, Craig Crews, Paul Forscher, Vivian Irish,
Archibald Perkins (Pathology)
Assistant Professors
Savithramma Dinesh-Kumar, Scott Holley, Christine Jacobs,
Frank Slack, Elke Stein, David Wells, Weimin Zhong
Fields of Study
Research in genetics and molecular biology encompasses
studies of catalytic RNAs, cell cycle regulation, chromosome
segregation, genetic recombination, mutation, transposons,
and oncogenes. Research topics in cellular and developmental
biology include structure of the cell cytoskeleton, molecular
motors, chemical biology, cell surface receptors, protein
transport, hormone action, mammalian transcription factors,
and the regulation of cell proliferation and differentiation.
Research in neurobiology focuses on sensory signal transduction,
animal color vision, growth cone motility, neural differentiation,
synaptogenesis, and the formation of topographic maps. A Special
Program in Plant Sciences provides research and training in
the molecular genetics of flowering, the developmental biology
of leaves, the physiology of hormone action, sex determination,
and the cellular and molecular biology of photomorphogenesis.
Because of the breadth of the track, students are provided
with unique opportunities for interdisciplinary studies.
To enter the Ph.D. program, students apply to an interest-based
track within the interdepartmental graduate program in the
Biological and Biomedical Sciences.
Special Admissions Requirements
Applicants should have obtained training in the structure,
development, and physiology of organisms; the structure, biochemistry,
and physiology of cells; genetics; elementary calculus; elementary
physics; inorganic and organic chemistry; statistics or advanced
mathematics. Lack of some prerequisites can be made up in
the first year of graduate study. Students having different
science training, such as degrees in chemistry, physics, or
engineering, are encouraged to apply. In addition to the GRE
General test, a Subject Test is required, preferably in Biology,
or in Biochemistry, Cell and Molecular Biology.
Special Requirements for the Ph.D. Degree
None of the fields of study has a required curriculum
of courses. Instead, with the help of a faculty committee,
each student plans a specific program that includes appropriate
courses, seminars, laboratory rotations, and independent reading
fitted to individual needs and career goals. There is no foreign-language
requirement. Late in the third term of study the student meets
with a faculty committee to decide on a preliminary topic
for dissertation work and to define the research areas in
which he or she is expected to demonstrate competence. By
the end of the second year each student prepares a dissertation
prospectus outlining the research proposed for the Ph.D. When
this is accepted by a dissertation committee of faculty members,
when the committee is satisfied that the student has demonstrated
competence in the areas necessary to conduct the proposed
work, and when the other requirements indicated above are
fulfilled, the student is admitted to candidacy for the Ph.D.
(but no later than the end of the second year of study). The
remaining requirements include completion of the dissertation
research, presentation and defense of the dissertation, and
submission of acceptable copies of the dissertation to the
Graduate School and to the Kline Science Library. All students
are required to teach in two one-term courses during their
Ph.D. study excluding the first year.
Honors Requirement
Students must meet the Graduate
School's Honors requirement by the end of the fourth term
of full-time study.
Master's Degree
M.S. (en route to the Ph.D.) The minimum requirements
for award of the Master of Science Degree are: (1) two academic
years registered and in residence full time in the graduate
program; (2) satisfactory completion of the first two years
of study and research leading to the Ph.D.; this requirement
may be met either (a) by completing a minimum of five courses
with an average grade of High Pass, or (b) by successfully
completing an approved combination of courses and research
and passing the prospectus examination; (3) recommendation
by the department for award of the degree, subject to final
review and approval by the appropriate degree committee. No
courses that were taken prior to matriculation in the graduate
program, or in Yale College, or in summer programs may be
applied toward these requirements.
Program materials are available upon request to the Director of Graduate Studies, Department of Molecular, Cellular, and Developmental Biology, Yale University, PO Box 208103, New Haven CT 06520-8103.
Courses
MCDB 500au, Biochemistry. L. Nicholas
Ornston, Robert Macnab. MWF 9.30–10.20
An introduction to the biochemistry of animals, plants,
and microorganisms, emphasizing the relations of chemical
principles and structure to the evolution and regulation of
living systems.
MCDB 505au, Molecular Genetics of Prokaryotes. Nigel
Grindley, Charles Radding, Joann Sweasy. MW 11.30–12.45
Molecular aspects of the storage, replication, evolution,
and expression of genetic material in prokaryotes. Also
GENE 705a, MB&B 705au.
MCDB 530au, Biology of the Immune System. Kim
Bottomly and staff. MWF 9.30–10.20
The development of the immune system. Cellular and molecular
mechanisms of immune recognition. Effector responses against
pathogens; autoimmunity. Also IBIO 530a.
MCDB 550au, Physiological Systems. Steven
Segal and staff. MWF 9.30–10.20
Organ systems of the human body, emphasizing the principles
of physiological control. Biophysical properties of cells,
tissues, and organs are considered in light of homeostasis
and the regulation of body functions. Also C&MP 550a,
ENAS 550au.
[MCDB 555bu, Molecular Basis of Development.]
MCDB 560bu, Cellular and Molecular Physiology: Mollecular
Machines in Human Disease. Michael Caplan, Emile
Boulpaep, Mark Mooseker. MWF 9.30–10.20
Focus on understanding the processes that transfer molecules
across membranes. Topics also include the different classes
of molecular machines that mediate membrane transport. Emphasis
on interactions among transport proteins in determining the
physiologic behaviors of cells and tissues. Also C&MP
560b.
MCDB 570bu, Biotechnology. Michael Snyder,
Ronald Breaker, Kenneth Nelson, Joseph Wolenski. MW 11.30–12.45
The principles and applications of cellular, molecular,
and chemical techniques that advance biotechnology. Topics
include the most recent tools and strategies used by government
agencies, industrial labs, and academic research to adapt
biological and chemical compounds as medical treatments, industrial
agents, or for the further study of biological systems.
MCDB 600Lb, Advanced Biological Techniques. Michael
Snyder, Xing-Wang Deng, Scott Holley, Kenneth Nelson, Joseph
Wolenski, David Austin. MW 1–5
A laboratory course to familarize graduate students with
state-of-the-art technologies in molecular biology, genomics.
Students carry out research projects and incorporate their
own projects into the lab. The class meets for two afternoons
each week and consists of 2–3 week modules covering
the following topics: microarray analysis, plant genetic engineering,
mouse genetic engineering, imaging/microscopy, ribozyme enzymol/engineering,
phage display/ chemical biology.
MCDB 602a, Molecular Cell Biology. Sandra
Wolin, Mark Mooseker, Thomas Pollard, Graham Warren. MW 1.45–3
A comprehensive introduction to the molecular and mechanistic
aspects of cell biology for graduate students in all programs.
Emphasizes fundamental issues of cellular organization, regulation,
biogenesis, and function at the molecular level. Also
CBIO 602a, MB&B 602a.
MCDB 603a, Seminar in Molecular Cell Biology. Sandra
Wolin, Mark Mooseker, Thomas Pollard, Graham Warren. Th 9–11
A graduate-level seminar course in modern cell biology.
The class is devoted to the reading and critical evaluation
of classical and current papers. The topics are coordinated
with the MCDB 602a lecture schedule. Thus, concurrent or previous
enrollment in MCDB 602a is required. Also CBIO 603a.
[MCDB 615bu, Genetics and Molecular Biology of Plant
Development.]
MCDB 625au, Basic Concepts of Genetic Analysis. Tian
Xu, Richard Lifton, Shirleen Roeder, Michael Stern. TTh 1–2.15
The universal principles of genetic analysis in eukaryotes
are discussed in lectures. Students also read a small selection
of primary papers illustrating the very best of genetic analysis
and dissect them in detail in the discussion sections. While
other Yale graduate molecular genetics courses emphasize molecular
biology, this course focuses on the concepts and logic underlying
modern genetic analysis. Also GENE 625a, MB&B 625au.
MCDB 630b, Biochemical and Biophysical Approaches in
Molecular and Cellular Biology. Thomas Pollard
and staff.
This graduate course introduces the theory and application
of biochemical and biophysical methods to study the structure
and function of biological macromolecules. The course considers
the basic physical chemistry required in cellular and molecular
biology but does not require a previous course in physical
chemistry. One class per week is a lecture introducing a topic.
The second class is a small group discussion of one or two
research papers utilizing those methods.
MCDB 642a, Roles of Microorganisms in the Living World.
L. Nicholas Ornston, Diane McMahon-Pratt, Robert Macnab. TTh
11.30–12.45
A topical course exploring the biology of microorganisms.
Emphasis on mechanisms underlying microbial adaptations and
how they influence biological systems. Also EMD 642a,
GENE 642a, MB&B 642a, MBIO 642a.
MCDB 660au, Structure, Function, and Development of
Vascular Plants. Graeme Berlyn. TTh 2.30–3.45
Morphogenesis and adaptation of vascular plants considered
from seed formation and germination to maturity. Physiological
and developmental processes associated with structural changes
in response to environment discussed from both a phylogenetic
and an adaptive point of view.
MCDB 670b, Advanced Seminar in Biochemistry and Genetics. Sidney
Altman, Ronald Breaker, Stephen Dellaporta. W 1.30–3.45
New aspects of the molecular biology of RNA, ribonucleoproteins,
and prions. Topics include the localization and function of
RNA and ribonucleoproteins; the role of RNA in dosage compensation,
chromosome silencing, and gene regulation; novel ribozymes
and RNA technology; prions. Discussion; involvement and attendance
are required.
MCDB 677b, Mechanisms of Development. Lynn
Cooley, Xing-Wang Deng, Scott Holley, Valerie Reinke, Frank
Slack, Michael Stern, Kevin White. M 9.45–11, F 1.30–3
An advanced course on the mechanisms of animal development
focusing on the genetic specification of cell organization
and identity during embryogenesis and somatic differentiation.
The use of evolutionarily conserved signaling pathways to
carry out developmental decisions in a range of animals is
highlighted. Course work includes student presentations and
critical analysis of primary literature. Also GENE 777b.
MCDB 685bu, Evolutionary Developmental Biology. Günter
Wagner, Vivian Irish, Kevin White. TTh 2.30–3.45
This course is an introduction into an emerging biological
discipline, evolutionary developmental biology. The course
provides an introduction to the evolutionary biology of developmental
processes as well as the developmental underpinnings of major
evolutionary transformations. Topics include the evolution
of Hox genes and other developmental genes, the origin of
multicellular organisms, the evolution of flowers, and the
origin of the arthropod and vertebrate Bauplan. The course
has a mixed lecture/seminar format and thus engages the student
to do independent study and prepare papers. Entering graduate
students are expected to complete a unique research project
and present in a lecture format to the class. Also E&EB
685bu.
[MCDB 692a, Advanced Seminar in Cell Biology: Mechanisms
of Signal Transduction.]
MCDB 720au, Neurobiology. Haig Keshishian,
Paul Forscher. MWF 11.30–12.20
Examination of the excitability of the nerve cell membrane
as a starting point for the study of molecular, cellular,
and intercellular mechanisms underlying the generation and
control of behavior. Also NBIO 720a, NSCI 720a.
MCDB 721Lau, Laboratory for Neurobiology. Haig
Keshishian, Robert Wyman. T or W 1.30–6
Optional laboratory. Introduction to the neurosciences.
Projects include the study of neuronal excitability, sensory
transduction, CNS function, synaptic physiology, and neuroanatomy.
MCDB 735bu, Seminar in Brain Development and Plasticity. Weimin
Zhong. MW 2.30–3.45
Interpretation of primary literature including recent
reviews and basic research papers in the areas of neuron generation
and regeneration, neuron phenotype determination, axon guidance
systems, and the role of activity in organizing and increasing
the efficiency of synaptic connections. Also NSCI 504b.
MCDB 750b, Core Topics in Biomedical Informatics. Perry
Miller and staff. T 1.30–3.20
Introduction to common unifying themes that serve as
the foundation for different areas of biomedical informatics,
including clinical, neuro-, and genome informatics. Emphasis
is on understanding basic principles underlying informatics
approaches to biomedical data modeling, interoperation among
biomedical databases and software tools, standardized biomedical
vocabularies and ontologies, and other topics of interest.
The course involves lectures, class discussions, student presentations,
and significant programming assignments. Prerequisite: previous
computer programming experience and permission of the instructor.
MCDB 752au, Genomics and Bioinformatics. Dieter
Söll, Mark Gerstein, Michael Snyder. MW 1–2.15
Genomics describes the determination of the nucleotide
sequence and many further analyses to discover functional
and structural information on all the genes of an organism.
Topics include the methods and results of functional and structural
gene analysis on a genome-wide scale as well as a discussion
of the implications of this research. Bioinformatics describes
the computational analysis of genomes and macromolecular structures
on a large scale. Topics include sequence alignment, biological
database design, geometric analysis of protein structure,
and macromolecular simulation. Also CPSC 752au, MB&B
752au.
MCDB 861bu, Global Problems of Population Growth. Robert
Wyman. TTh 2.30–3.45
The worldwide population explosion in its human, environmental,
and economic dimensions. Sociobiological bases of reproductive
behavior. Population history and the cause of demographic
change. Interactions of population growth with economic development
and environmental alteration. Political, religious, and ethical
issues surrounding fertility; human rights; and the status
of women.
MCDB 900a and 901b, First-Year Introduction to Research. Susan
Ferro-Novick, Ronald Breaker, Michael Stern.
Lab rotations, grant writing, and ethics for Molecular
Cell Biology, Genetics, and Development track students. Also
CBIO 900a, 901b; GENE 900a, 901b.
MCDB 950a and 951b, Second-Year Research.
By arrangement with faculty.
The following courses are required for students in the
joint B.S./M.S. program with Yale College:
MCDB 585b, Research in MCDB for B.S./M.S. Candidates.
A two-credit course taken in the third-to-last term (typically
the second term of the junior year). At the end of this course,
students complete a detailed prospectus describing their thesis
project, and the work completed thus far. An oral and written
presentation of this prospectus is evaluated by the adviser
and two faculty members; the evaluation will determine whether
the student may continue in the program.
MCDB 595, Intensive Research in MCDB for B.S./M.S. Candidates.
A four-credit course (two credits each term) that is
similar to MCDB 495 and spans the last two terms (i.e., typically
the senior year). During this course, students give an oral
presentation describing their work. At the end, a comprehensive
thesis is turned in and evaluated by the adviser and two other
faculty members. Students must earn a B grade or higher in
this course in order to receive the M.S. degree.
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