Department of Ecology and Evolutionary Biology
Daniel I. Rubenstein
Director of Graduate Studies
Andrew P. Dobson
Iain D. Couzin
Andrew P. Dobson
James L. Gould
Bryan T. Grenfell, also Woodrow Wilson School
Lars O. Hedin, also Princeton Environmental Institute
Laura F. Landweber
Simon A. Levin
Stephen W. Pacala
Daniel I. Rubenstein
David S. Wilcove, also Woodrow Wilson School
Peter Andolfatto, also Lewis-Sigler Institute for Integrative Genomics
Andrea L. Graham
Robert M. Pringle
Assistant Professor (continued)
Corina E. Tarnita
Lecturer with Rank of Professor
Stuart A. Altmann
Asif A. Ghazanfar, Psychology, Princeton Neuroscience Institute
Alan E. Mann, Anthropology
Ignacio Rodríguez-Iturbe, Civil and Environmental Engineering
Bess B. Ward, Geosciences
Peter R. Grant
Henry S. Horn
Graduate study in the Department of Ecology and Evolutionary Biology is designed to lead to the degree of doctor of philosophy (Ph.D.). Postdoctoral students interested in furthering their research background in these areas are also welcome. Students with an interest in behavioral neuroscience should see the Program in Neuroscience section below.
The special areas of strength in the department are evolutionary ecology, behavioral ecology, theoretical ecology, population, community, and ecosystem ecology, ecological and evolutionary genetics, molecular evolution, epidemiology of infectious diseases and conservation biology. The interests and research of faculty range widely over these areas, so that incoming students are able to select their adviser from among several professors working in the chosen discipline. Graduate students also have excellent opportunities for combining several areas for innovative interdisciplinary work.
The graduate program is designed to develop both the breadth and depth of understanding that will enable graduates of the department to respond to future advances in the field. At the same time, students acquire the detailed knowledge and techniques needed to become effective scientists. Each student is guided in developing a comprehensive but flexible course of preparation that is designed to meet their educational needs and goals.
There are only a few formal course requirements; the principal emphasis is on independent research and this begins early. Incoming students meet with the faculty within two weeks of arrival for a discussion of the general curricular and logistic needs for their chosen course of graduate study. Generally first- and second-year students take six core courses; these include a sequence of fundamental papers, the journal club, a course on professional issues and the field course on tropical ecology. Other advanced topic-related courses may be taken occasionally throughout the graduate career. In addition to course work and consultation with faculty and other graduates, students enjoy the benefits of an excellent series of seminars and colloquia throughout the year. These seminars, given by eminent visiting scientists, expand the student’s educational experience beyond the bounds of expertise that can be found in the local Princeton community, and keep the faculty and students abreast of the latest developments in their field.
There are no rigid prerequisites for admission. The program accepts graduate students with previous specializations either in biology or one of its branches, or in chemistry, engineering, the humanities, mathematics, physics, and the social sciences. A student will find it useful to have taken some basic courses in biology (including genetics) as well as calculus, organic chemistry, physics and statistics. Deficiencies in these areas may be remedied through independent reading, summer courses, or other means. If necessary, students may take basic courses during their first year.
Assessment of Academic Background
In order to determine if newly admitted students have sufficient knowledge of ecology, evolution and behavior, an advisory committee consisting of three faculty members meets with each student to discuss his or her aims and academic background. Basic knowledge in each area should be equivalent to that gained in comprehensive upperclass undergraduate courses. During the first two years, members of the committee recommend courses and are available for consultation on planning and initiating research projects.
All students are required to enroll in three departmental core courses in their first year. In addition, if they need them, students may take undergraduate courses to remedy academic deficiencies. In consultation with the director of graduate studies, new students choose a temporary adviser with whom they plan individual programs of course work and research. Students may take graduate or undergraduate courses in the department or relevant courses in any other department.
Seminars, Colloquia, and Integrated Research Seminars
Students attend colloquia related to their interests. These colloquia are offered as a weekly seminar series; they are conducted by Princeton faculty, students and outside speakers. During the academic year, many seminars are sponsored by the department and also the molecular biology, chemistry, geology, and psychology departments. Other departments and programs feature prominent outside speakers. Seminars and colloquia form an important part of the student’s graduate education and frequently attract faculty and student audiences from several departments.
Since there are many research groups that work on aligned projects, a series of Integrated Research Seminars have emerged – Theoretical Ecology Lab Tea, Integrated Behavioral Research Group, and Disease Modeling Group – which meet weekly, in addition to monthly student or postdoc colloquia in Genomics and other themes on campus.
During the first year, each student presents a seminar describing their research —often a presentation to one of the integrated research seminars. In their second and fourth years, students give more formal presentations during lunchtime seminars. In the fourth or fifth year, financial support is available to encourage students to give a seminar at a national or international meeting. All of these talks are designed to provide experience in presenting original scientific research.
New students are encouraged to begin research projects as soon as possible. In some cases the initial project becomes the thesis topic, but many students work on several smaller projects before focusing on the subject of their dissertation. To help find a thesis topic and a supervisor, students can work closely during their first two years with a series of faculty members. Graduate students are expected to continue their research and training during the summers at Princeton, in the field, or at another laboratory.
By the early spring of their second year of graduate enrollment, students must submit a research report and thesis proposal to stand for the general examination.
The general examination is normally taken in the spring of the second year of study. There is no prejudice against taking the examination after only one year of study if a student feels prepared.
The general examination consists of an oral examination, about three hours in length, conducted by the student’s dissertation committee, which is chosen in advance by the student and the adviser. The dissertation committee is normally composed of four or five Princeton faculty members; it may include members of other departments within the University. Should the student’s area of study warrant it, faculty members from other institutions with special competence are invited to serve on the dissertation committee.
The student submits a written review of background information relevant to the thesis topic as well as a thesis proposal detailing research objectives, preliminary progress and future plans. Questions asked during the oral examination focus on the thesis topic but also cover all scientific areas relevant to the topic of the thesis.
After passing the general examination, a student is awarded an incidental master’s degree (M.A.).
Dissertation and Final Public Oral Examination
After the general examination, the dissertation committee continues to meet with the graduate student at least once a year to discuss the student’s research. The student is expected to prepare a written summary of his or her work before each of these meetings. As the final step in the graduate program, the committee evaluates the dissertation and once the committee approves the thesis, the candidate has two weeks to prepare and present to the department the final public oral examination for the doctoral degree. Normally, degree requirements are completed in five years.
Each graduate student usually teaches for two terms in an undergraduate laboratory course. This provides valuable mentoring experience to the student and provides a helpful foretaste of an essential aspect of an academic career.
All graduate students entering the program are supported in their first year by a first-year fellowship or the prestigious Centennial Fellowship provided by the Graduate School. From the second year onward, support derives from fellowships provided by the department, training grants and personal fellowships won by students and assistantships for teaching in undergraduate laboratories and precepts. In order to help the department spread its financial resources, prospective graduate students who are U.S. citizens or permanent residents are requested to apply for national fellowships that provide stipend and tuition support. Information on such fellowships is available from the financial aid or career counseling offices as well as the National Science Foundation.
Foreign students are encouraged to apply for financial support from their native countries. Fellowships and assistantships include a stipend for 12 months and a full tuition grant. Students in good standing will continue to receive support during subsequent years of graduate study.
Program in Neuroscience
Interactions among neuroscience faculty in the departments of ecology and evolutionary biology, molecular biology, and psychology are formally recognized by the Program in Neuroscience, which makes possible a Ph.D. in biology and neuroscience for students enrolled in these departments. Please see the program description.
The Princeton Environmental Institute (PEI) offers, in affiliation with the Woodrow Wilson School of Public and International Affairs, an opportunity in its program for Science, Technology, and Environmental Policy (STEP) for graduate students to explore the environmental policy dimensions of their doctoral research. Through the PEI-STEP program, PEI offers half financial support for two years (stipend and tuition). Support is generally awarded in the second and third years of the student's Ph.D. tenure. PEI-STEP Fellows also receive a $3,500 award to support their graduate research.
Ecology and Evoluntionary Biology
EEB 503 Fundamental Concepts in Evolution
Peter Andolfatto, Andrea Graham, Bridgett vonHoldt
An advanced foundation in evolution. Topics include evolutionary and quantitative genetics, molecular evolution, and evolutionary stable strategies.
EEB 504 Fundamental Concepts in Ecology, Evolution, and Behavior II
Peter Andolfatto, Iain Couzin, James Gould, Andrea Graham, Bryan Grenfell, Lars Hedin, Laura Landweber, Simon Levin, Daniel Rubenstein, Bridgett vonHoldt, and David Wilcove
Advanced discussions of ecology, evolution, and behavior focus on 50 fundamental papers. Ecological topics include dynamics and structure of populations, communities, ecosystems, and conservation biology. Behavioral topics include instinct and learning, social behavior, physiological ecology, and the evolution of behavior. Evolutionary topics include speciation, evolutionary and quantitative genetics, molecular evolution, evolutionary stable strategies, and evolution of development. (This is a core course.)
EEB 506 Responsible Conduct in Research
A six-week course satisfying the training mandate for the responsible conduct of research (RCR). Each meeting is led by one or two faculty members using the text: “On Being a Scientist: A Guide to Responsible Conduct in Research.” Each meeting covers two chapter-topics, except for the trio of subjects—authorship, intellectual property and competing interests. Key points on each topic, a list of questions, and at least one case study are discussed by the group. Supplemental case studies are provided by the faculty member leading each class. (This is a core course.)
EEB 507 Recent Research in Population Biology
Henry Horn and Corina Tarnita
Systematic reviews of recent literature in areas of ecology, evolution, and animal behavior are made. The general survey of literature is supplemented with detailed discussion of selected research papers of unusual importance and significance. (This is a core course.)
EEB 511/APC 511 Seminar in Mathematical Biology
Applications of reaction-diffusion equations in mathematical biology. Specific examples of Fisher-Kolmogorov-Petrovski-Piskunov equation and FitzHugh-Hagumo system of equations. Discussion of modeling principles, as well as methods of mathematical analysis, the latter to include phase plane analysis for ordinary differential equations, maximum principle for parabolic PDE's, methods for singularly perturbed systems, and matched asymptotic expansions.
EEB 514/APC 514 Biological Dynamics
Introduction to the mathematical description of quantitative phenomena in living systems; Hodgkin Huxley equations of nerve membranes; the generation of spatial patterns in development, single cells, and colonies of cells; chemotaxis; the population dynamics of disease; dynamics activity of networks of neurons; intracellular chemical and gene-networks. Emphasis on formulation and experimental basis for the equations, and their relationship to significant biological issues.
EEB 515 Conservation Biology
Introductory lectures on ecological principles underlying biodiversity; the dynamics of populations and ecosystems; the origin, maintenance, and loss of genetic variation; and the causes of extinction and speciation. These and other topics are developed in subsequent discussions and presentations. Examples include consequences of habitat fragmentation, theory and methodology of breeding of endangered species, and design criteria for nature preserves.
EEB 516/WWS 586B Topics in Science Technology and Environmental Policy
David Wilcove and Lars Hedin
These are courses intended to help students develop and apply skills in the application of scientific, technological, and environmental analyses to problems of policy interest. Fall courses are numbered 585, spring courses are numbered 586.
EEB 517/MOL 515 Method and Logic in Quantitative Biology
Ned Wingreen and David Botstein
Close reading of published papers illustrating the principles, achievements, and difficulties that lie at the interface of theory and experiment in biology. Two important papers, read in advance by all students, will be considered each week; emphasis will be on student discussion, not formal lectures. Topics include: cooperativity, robust adaptation, kinetic proofreading, sequence analysis, clustering, phylogenetics, analysis of fluctuations, maximum likelihood methods.
EEB 519 Theoretical Ecology
The investigation of the ecological and evolution of natural communities through the use of mathematical models. Topics include complexity and stability, patterns of species abundance and diversity, theory of the niche, spatial and temporal heterogeneity, and the dynamics of natural populations.
EEB 521 Tropical Ecology
Intensive three-week field course undertaken in January in a suitable tropical locality. There are readings, discussions, and individual projects. The content and location are varied to suit the needs of the participants. Students provide their own travel funds and faculty teach this on rotation. (This is a core course.)
EEB 522 Colloquium on the Biology of Populations
Lars Hedin and Simon Levin
Discussion of the central problems of population biology and approaches that have proved fruitful. Topics ranging throughout ecology, evolution, biogeography, and population genetics are usually related to presentations by visiting speakers and students. (This is a core course.)
EEB 524 Topics in Evolution
Laura Landweber and Bridgett vonHoldt
Discussion of current topics in evolutionary biology. Topics vary from year to year, depending on current controversial issues.
EEB 525 Quantitative Field Ecology
Techniques for physical measurement, photography, mapping, and recording data. Development of simple but specialized statistics. Strategies for handling spatial and individual variations. Constructive criticism of statistics in the literature of ecology and behavior. Specific exercises are adapted to students' needs. One afternoon per week there is a lecture, seminar, demonstration, and/or fieldwork. Prerequisite: a working knowledge of elementary statistics.
EEB 526/ENV 526 Environmental Issues Seminar
Ignacio Rodriguez-Iturbe and Daniel Rubenstein
Course examines the links between hydrology, landscape and wildlife ecology and socio-political aspects of African arid lands and explores how understanding these dynamics shape such landscapes and how an understanding of these dynamics can be used to create an integrated foundation for conserving resources while enhancing livelihoods.
EEB 529/PSY 510 Topics in Neuroscience and Behavior
Lectures on selected topics of current interest in neuroscience, and behavior. Emphasis is on areas of active current research; content reflects the interests of participating students.
EEB 530/PSY 518 Introduction to Neurochemistry and Neuropharmacology
Basic research methods and concepts in neurochemistry and neuropharmacology. The emphasis is on chemical neurotransmission. Topics include brain metabolism; structure and function of cell membranes; neuronal excitation and conduction; neurotransmitter synthesis; packaging, release, reuptake, and catabolism; neuropeptides, second messengers, role of chemical neurotransmission in physiology, and behavior; and the mechanisms of action of psychoactive drugs.
EEB 531 Advanced Vertebrate Biology
Topics vary from year to year depending on the instructor's and the students' special interests and may include patterns of reproduction relating to breeding seasons, delayed births, parental care, mating, ovulatory cycles, age of sexual maturity, and viviparity.
EEB 532 Topics in Animal Behavior
Topics vary from year to year but cover areas of behavioral ecology, sociobiology, the ethology of learning, and the evolution of intelligence. Lectures delineating the fundamental issues surrounding each topic are followed by discussions of the relevant literature or by a small-group research project.
EEB 533 Topics in Ecology
Discussion of the growing points in population and community ecology is held. Topics vary from year to year and include such subjects as the regulation of population numbers, the organization of communities, conservation, and the variation in ecological processes at different spatial and temporal scales.