Department of Ecology and Evolutionary Biology
Chair
Daniel I. Rubenstein
Acting Chair
Simon A. Levin (fall)
Departmental Representative
James L. Gould
Lars O. Hedin
Professor
Jeanne Altmann
Andrew P. Dobson
James L. Gould
Lars O. Hedin, also Princeton Environmental Institute
Henry S. Horn
Leonid Kruglyak, also Lewis-Sigler Institute for Integrative Genomics
Simon A. Levin
Stephen W. Pacala
Daniel I. Rubenstein
David S. Wilcove, also Woodrow Wilson School
Visiting Professor
Robert M. May
William H. Schlesinger
Associate Professor
Laura F. Landweber
David L. Stern
Assistant Professor
Peter Andolfatto, also Lewis-Sigler Institute for Integrative Genomics
Iain D. Couzin
Lecturer with Rank of Professor
Stuart A. Altmann
Associated Faculty
Alan E. Mann, Anthropology
Ignacio Rodríguez-Iturbe, Civil and Environmental Engineering
Burton H. Singer, Woodrow Wilson School
Bess B. Ward, Geosciences
Departmental Plan of Study
Courses in the biological sciences at Princeton are offered in two departments. Students with an interest in whole-organism and large-scale processes—evolution (including molecular evolution and developmental evolution), physiology, behavior, ecology, ecosystem biology, and conservation—should enroll in the Department of Ecology and Evolutionary Biology (EEB). Those with interests in molecular, cellular, and developmental processes should enroll in the Department of Molecular Biology (MOL). Both departments provide an excellent background for medical school.
The EEB department emphasizes research and teaching from an evolutionary perspective, combining theory and empiricism and linking areas that are often treated as separate disciplines. Many of the research projects and courses are interdisciplinary. A description of these core areas, faculty interests, and unique features of the program, including details about the department’s field programs, can be found on the EEB website: www.eeb.princeton.edu/index.html.
Every student considering majoring in EEB should attend the departmental open house held in the spring term. It introduces potential majors to departmental courses, faculty, and the wide-ranging research and field-course opportunities open to our students. See the EEB website for examples of recent student research activities.
Advanced Placement. Advanced placement will be granted to students who received a 5 on the Biology AP exam. This excuses EEB majors from taking 211.
Prerequisites. To enter the department, an EEB student should take:
EEB 211 or 210, and 214 or 215.
Requirements. The following courses are required for graduation:
1. General chemistry: CHM 201 or 207, and 202;
2. Organic chemistry: CHM 301 or 303; (Students interested in mathematical biology may take one of the following courses instead of organic chemistry: PHY 203, 208; MAT 200, 201, 202, 203, 204; COS 126, 226; ORF 245, 406);
3. Molecular biology/evolution: 320, MOL 214, 215, or 459, or any other nonpolicy-oriented 300- or 400-level MOL course with permission of the instructor;
4. Mathematics/computing: MAT 101 and 102, or 103, or 104, or COS 126;
5. Physics: PHY 101, 103, 105, or 107.
Choice of Courses. Students in EEB must successfully complete at least eight upper-level departmental courses, five of which must be EEB or MOL courses. One of the eight must be a lab course; 314, 324, 417B, and all field courses count as lab courses. Students must take at least one course from each of the following three general areas:
Behavior and Physiology: 301, 311, 314, 315, 323, 350, 404, PSY 315;
Evolution and Genetics: 306, 309, 320, 360, MOL 342;
Ecology and Conservation: 308, 321, 322, 324, 328, 338, 346, 352, 380, 417.
Students may take 309, 311, 321, and 323 in the fall term of junior year to sample all three areas. Pay particular attention to the timing of the courses that are taught in alternate years.
No course taken pass/D/fail can be counted as a departmental. The minimum grade for a course to count as a departmental is C-. Only one course with a policy perspective can be counted as a departmental. Only Princeton courses can count as departmentals; the one exception is for pre-approved courses taken during a study abroad term.
Special Features of the Plan of Study. EEB offers three tropical field study programs: a semester in Panama, a semester in Kenya, and a summer in Bermuda. Details of these programs can be found on the EEB website.
Specializations. Because some EEB students are interested in specializing within the department, EEB offers several areas of focus, the most common of which are listed below with their relevant courses.
Theoretical biology: 309, 320, 321, 322, and 324;
Field biology: 309, 311, 321, and a tropical field program;
Conservation biology: 308, 309, 311, 321, 323, 417, 516, and a tropical field program;
Medical biology: 309, 311, 314, 321, 323, and 328;
Human biology: 301, 306, 309, 315, PSY 330, and 336.
Note: MOL 214 is required for medical school. A full year of physics is required for medical school and by some graduate programs (especially in neurobiology). A full year of organic chemistry is required for medical school, and is a prerequisite for many molecular biology courses; the second term also counts as an EEB departmental. At least one term of organic chemistry is usually required by graduate programs in biology. Completing the organic chemistry requirement before junior year is strongly recommended.
Junior Independent Work. Students attend a weekly junior tutorial in the first half of the fall term. Faculty members present summaries of their work and research opportunities open for senior independent work. During the second half of the fall term, students work closely with one faculty member to investigate a problem using the current literature, and then write a paper. In the spring semester, students carry out a program of independent work with a faculty adviser. In some instances this may include empirical or theoretical work. Either a paper summarizing this project or a research proposal for the senior thesis is due in early May.
Senior Independent Work. During the fall or early spring of the junior year each student selects a senior thesis adviser. The adviser and the student choose a research project that the student generally pursues during the summer preceding the senior year and both terms of the senior year. The research project can involve primarily laboratory, field, data mining, theoretical, or library study that will be written and presented as a senior thesis. A one-hour oral examination, consisting of a defense of the thesis research and general questions in the biological sciences, and a poster session, will be held in May.
Tropical Field Programs. Students interested in learning about or undertaking research in the tropics have a number of options.
1. Panama. The department offers a spring term in Panama in conjunction with the Smithsonian Tropical Research Institute. Students take four intensive three-week courses in sequence, beginning with an introduction of key concepts in tropical ecology and conservation, followed by a course that incorporates extensive field trips and research projects. The program continues with an anthropology course and a final three-week course on a particular aspect of tropical biology. Prerequisite: EEB 321.
2. Kenya. This four-course program, also taught in three-week segments, takes place at Princeton University’s Mpala Research Centre in central Kenya and other sites in Kenya, in collaboration with scientists there, EEB faculty, and other appropriate faculty from Princeton University. The courses delve into conservation in Africa, restoration ecology, the natural history of mammals, and global technology and engineering. Prerequisite: EEB 321.
3. Bermuda. During June, the department, along with the Bermuda Institute of Ocean Sciences, offers Marine Biology. This four-week course focuses on the ecology, physiology, and behavior of marine organisms, particularly from the coral reef. Almost half of the labs involve field trips to the reefs or intertidal zone. During July, the Department of Geosciences offers GEO 318 Observing the Marine Environment. These courses are open to rising sophomores and juniors.
4. Other. Individual students are welcome to pursue other independent field opportunities, with scientists from the Smithsonian Institution and the Bermuda Institute of Ocean Sciences, or other research institutions, such as the School for Field Studies at the University of Cape Town, South Africa, or James Cook University in Australia.
Program in Teacher Preparation. As the need for qualified science teachers increases, some students may wish to earn a teaching certificate. Working with the departmental representative and the teacher preparation program, an appropriate course of study can be arranged.
Courses
EEB 210 Evolutionary Ecology (also MOL 210) — Not offered this year ST
An introduction to the mechanisms and processes of population biology. Emphasis on understanding the structure, dynamics, and evolution of natural ecosystems, how plants and animals work and behave, and how genotypes and environments interact to mold evolutionary change. Counts toward requirements for entrance to medical school; premedical students as well as students planning to major in science or engineering will find 211 more appropriate. Three lectures, one class, one three-hour laboratory. D. Rubenstein, S. Pacala
EEB 211 The Biology of Organisms (also MOL 211) — Fall ST
An introduction to the biology of organisms and populations. Topics include evolution of life and biological diversity, physiology of plants and animals, neurobiology and behavior, ecology, evolution, and conservation biology. Counts toward requirements for entrance to medical school. Three lectures, one class, one three-hour laboratory. J. Gould, J. Bonner
EEB 214 Introduction to Cellular and Molecular Biology (see MOL 214)
EEB 215 Quantitative Principles in Cell and Molecular Biology (see MOL 215)
EEB 255 Life in the Universe (see GEO 255)
EEB 301 Evolution and the Behavior of the Sexes (also WOM 301) — Spring
Psychological, biological, and cross-cultural approaches to the study of sex and gender. Topics include biological components and development of sex differences; acquisitions of gender identity; social organization of key life cycle events; evolutionary considerations in the study of sex differences. One 90-minute lecture, one 90-minute class. J. Altmann
EEB 306 Human Evolution (see ANT 206)
EEB 308 Conservation Biology — Fall
A detailed application of ecological principles to the conservation of biological resources, including island biogeography, population genetics and viability, and landscape ecology. Analysis of case studies in conservation. Individual project on a conservation issue of the student’s choice. Two lectures, one preceptorial. Staff
EEB 309 Evolutionary Biology — Fall
All life on Earth has evolved and continues to evolve. This course examines how natural selection generates biological diversity, how the genome enables and constrains particular evolutionary paths, and how the unthinking apparently selfish behavior of genes leads to altruistic behavior amongst individuals. The course examines the mechanisms of speciation and the causes of extinction. Evolution provides the glue that makes all the disparate facts in biology stick together. This course will provide the basic tools to understand how and why this is true. Three lectures, one preceptorial. Prerequisite: 210 or 211. D. Stern
EEB 311 Animal Behavior — Not offered this year
An examination of the mechanisms and evolution of the behavior of human and other animals. Topics include the sensory worlds of animals, the nature of instinct, neural mechanisms of perception, comparative studies of communication, learning, cognition, mate choice, and social behavior, and the biology of human development and language acquisition. Offered in alternate years. Two 90-minute lectures, one preceptorial. J. Gould
EEB 312 Marine Biology (also ENV 312) — Summer ST
An intensive four-week course in Bermuda. Covers elements of the ecology, evolution, physiology, and behavior of marine organisms and ecosystems. Habitats examined will include the intertidal zone, seagrass beds, marshes, mangroves, and the open ocean, with special attention to coral reefs. Topics range from the physiology and behavior of individuals in the habitat, to the flow of energy, predator/prey interactions, symbioses, and population dynamics. Prerequisites: 210 or 211, ability to swim. Three hours of lectures, three hours of laboratory and field trips per day. J. Gould
EEB 314 Comparative Physiology — Spring
The study of how animals function with emphasis on the integration of physiological processes at the cellular, organ, and whole organism levels in ecological and evolutionary contexts. Comparisons among species and higher taxa are used to illustrate general physiological principles and their evolutionary correlates. Three lectures. Prerequisite: 210 or 211. Staff
EEB 315 Human Adaptation (see ANT 215)
EEB 320 Molecular Evolutionary Genetics (also MOL 330) — Spring
The use of DNA sequence comparisons to infer both micro- and macroevolutionary trends. Topics include the evolution of early life, eucaryotes, eucaryotic organelles, genome organization, protein synthesis, and intron processing. Three lectures. Prerequisite: 214. L. Landweber
EEB 321 Introduction to Population and Community Ecology — Fall
A comprehensive introduction to the ecology of populations, communities, and ecosystems. The course will first examine the dynamics of single-species populations of plants and animals. It will then examine the ecological interactions between pairs of species such as competition, mutualism, predation, parasitism, and herbivory. Finally, it will consider the dynamics and structure of whole communities of plants and animals. Two 90-minute lectures, one lab. H. Horn
EEB 322 Advanced Ecology — Not offered this year
An advanced overview of the structure of ecological communities, particularly temperate and tropical forests. Emphasis will be on factors governing species diversity and abundance on both local and global scales. Other topics will include the impact of humans on biodiversity at global scales, and the effects of biodiversity on the regulation of climate and the cycling of key elements such as carbon and nitrogen. Prerequisite: 321; one year of calculus recommended. Two 90-minute lectures, one preceptorial. S. Pacala
EEB 323 Integrative Dynamics of Animal Behavior — Fall
An exploration of the fundamental principles underlying the organization and function of animal behavior. This course will examine how complex actions emerge from simple rules. Since forces shaping behavior naturally cross scales and disciplinary boundaries, this course will draw on information from neuroscience, evolutionary biology, ecology, physiology, genetics, and the biology of complex systems. Two 90-minute lectures, one preceptorial. I. Couzin
EEB 324 Theoretical Ecology — Not offered this year QR
Current and classical theoretical issues in ecology and evolutionary biology. Emphasis will be on theories and concepts and on mathematical approaches. Topics will include population and community ecology, epidemiology and evolutionary theory. Two lectures, one preceptorial/computer laboratory. Prerequisite: one year of calculus. S. Levin
EEB 328 Ecology and Epidemiology of Parasites and Infectious Diseases — Spring
An introduction to the biology of viruses, bacteria, fungi, protozoa, worms, arthropods, and plants that are parasitic upon other animal and plant species. The major emphasis will be on the parasites of animals and plants, with further study of the epidemiology of infectious diseases in human populations. Studies of AIDS, anthrax, and worms, and their role in human history, will be complemented by ecological and evolutionary studies of mistletoe, measles, myxomatosis, and communities of parasitic helminths. Two lectures, one preceptorial. Prerequisite: 210 or 211; one year of calculus recommended; 321. A. Dobson
EEB 332 Pre-Columbian Peoples of Tropical America and
Their Environments (also LAS 350) — Spring SA
The pre-European history of Amerind cultures and their associated environments in the New World tropics will be studied. Topics to be covered include the people of tropical America; development of hunting/gathering and agricultural economies; neotropical climate and vegetation history; and the art, symbolism, and social organization of native Americans. Field and laboratory experiences will incorporate methods and problems in field archaeology, paleoethnobotany and paleoecology, and archaeozoology. Limited to students in the Tropical Ecology Program in Panama. Prerequisite: 321. R. Cooke, D. Piperno
EEB 338 Tropical Biology (also LAS 351) — Spring ST
This intensive field course given at various sites in Panama examines the origins, maintenance, and major interactions among elements of the tropical-terrestrial biota. Study topics include identification of common orders and families of Neotropical organisms, especially plants and arthropods; tropical climate and hydrology in lowland and montane forests; biotic interactions in exemplary systems; and contemporary and historical factors in shaping tropical landscapes, with emphasis on the Isthmian Landbridge and subsequent floral and faunal interactions. Limited to students in the Tropical Ecology Program in Panama. Prerequisite: 321. Y. Basset
EEB 346 Biology of Coral Reefs — Spring ST
This field and lecture course provides an in-depth introduction to the biology of tropical coral reefs, with an emphasis on reef fish ecology and behavior. Students learn to identify fishes, corals, and invertebrates, and learn a variety of field methods including underwater censusing, mapping, videotaping, and the recording of inter-individual interactions. Each year group projects will vary depending on previous findings and the interests of the faculty. Limited to students in the Tropical Ecology Program in Panama. Prerequisite: 321. S. Pacala
EEB 350 Vertebrate Tropical Ecology — Spring
This field course addresses the life history characteristics of tropical vertebrates and the physiological traits that underlie them. Students will learn how tropical life histories differ from those in the temperate zone and will use eco-physiological techniques while conducting experiments and observations at a Smithsonian Institute field station. Limited to students in the Tropical Ecology Program in Panama. Prerequisite: 321. M. Wikelski
EEB 379 Special Topics in Public Affairs (see WWS 479)
EEB 404 Natural History of Mammals — Spring ST
Introduction to concepts, methods, and material of comparative natural history, with mammals as focal organisms. The goal is appreciation of how these vertebrates interact with diverse and potentially conflicting features of their environment. Perspectives include morphology, identification, evolution, ecology, behavior, habitat, and conservation. Original observations and experiments in field and lab, supplemented by field trips, lectures, and reading, culminate in class, group, and individual research projects. Prerequisites: 210 or 211. Limited to students in the Tropical Ecology Program in Kenya. D. Rubenstein
EEB 414 Genetics of Human Populations (also MOL 414) — Fall
This advanced seminar surveys the evolutionary history of modern humans and the genetic basis of variation in our species through reading and discussion of classic and contemporary primary literature. Topics include the evolutionary origins of modern human populations, signatures of natural selection in the human genome, and approaches for discovering genetic variants that affect disease susceptibility and variation in normal traits. Significant emphasis is placed on recent advances made possible by the human genome project. Prerequisites: 309, or 320, or MOL 342, or CHM 236. One three-hour seminar. L. Kruglyak
EEB 417A, 417B Ecosystems and Global Change (also ENV 417A, 417B) — Fall
An introduction to the concepts, approaches, and methods for studying complex ecological systems from local to global scales. Students will examine nutrient cycling, energy flow, and evolutionary processes, with emphasis on experimental approaches and comparisons between terrestrial, freshwater, and marine ecosystems. Particular attention will be on effects of human activities, including climate change, biodiversity loss, eutrophication, and acid rain. Prerequisites: 210 or 211 or equivalent; CHM 301 or equivalent. 417a: two 90-minute classes. 417b: two 90-minute classes, one three-hour laboratory. L. Hedin
EEB 419 Environmental Microbiology (see GEO 417)
EEB 422 Evolutionary Developmental Biology (also MOL 422) — Not offered this year
What makes a butterfly different from a fly and a chimpanzee different from a human? This course explores how the genetic mechanisms that control development have evolved to generate the diversity of life on earth. How can conserved genes and genetic networks produce a stable phenotype and yet evolve to generate diversity? Topics include body-plan evolution, flower developmental evolution, life-history evolution, phenotypic plasticity, allometry, social insects, canalization and developmental constraints, evolvability and evolutionary dead-ends. Prerequisites: 309 and either MOL 348 or MOL 342. Three lectures. D. Stern
Approved Courses for Departmental Credit
Ecology and Evolutionary Biology (EEB)
All 300- and 400-level courses, except 332.
500-level courses, with permission of the instructor.
Molecular Biology (MOL)
All 300- and 400-level courses.
320 see WWS 320**
342 Genetics
345 Biochemistry
348 Cell and Developmental Biology
350 Laboratory in Molecular Biology
408 Cellular and Systems Neuroscience
410 Introduction to Biological Dynamics
429 Selected Topics in Molecular Biology and Human Genetics
431 Advanced Topics in Developmental Neurobiology
434 Macromolecular Structure and Mechanism in Disease
435 Pathogenesis and Bacterial Diversity
437 Computational Neuroscience
450 Stem Cells and Cell Fate Decision Processes in the Genomic Era**
457 Computational Aspects of Molecular Biology
459 Viruses: Strategy and Tactics
460 Diseases in Children: Causes, Costs, and Choices**
Most graduate-level MOL courses, with permission of the instructor.
Anthropology (ANT)
*431 Biomedical Anthropology
Chemical Engineering (CHE)
447 Biochemical Engineering
Chemistry (CHM)
301 Organic Chemistry I
302 Organic Chemistry II
303 Organic Chemistry I: Biological Emphasis
304 Organic Chemistry II: Biological Emphasis
306 Physical Chemistry: Thermodynamics and Kinetics
331 see GEO 331
333 Oil to Ozone: Chemistry of the Environment (also ENV 333)
542 Principles of Macromolecular Structure
Civil and Environmental Engineering (CEE)
417 see GEO 417
471 see GEO 471
Computer Science (COS)
126 General Computer Science
226 Algorithms and Data Structures
Economics (ECO)
329 Environmental Economics**
Environmental Studies (ENV)
322 see GEO 322
333 see CHM 333
399 see GEO 399**
401 Environmental Policy Workshop**
Geosciences (GEO)
322 Biogeochemical Cycles and Global Change (also ENV 322)
331 Introduction to
Environmental Geochemistry: Chemistry of the Natural Systems (also CHM 331)
399 Environmental Decision Making (also ENV 399)**
417 Environmental Microbiology (also CEE 417)
428 Biological Oceanography
471 Introduction to Water Pollution Technology (also CEE 471)
Mathematics (MAT)
200 Linear Algebra with Multivariable Calculus for Economists
201 Multivariable Calculus
202 Linear Algebra with Applications
203 Advanced Multivariable Calculus
204 Advanced Linear Algebra with Applications
217 Honors Linear Algebra
Operations Research and Financial Engineering (ORF)
245 Fundamentals of Engineering Statistics
406 Statistical Design of Experiments
Physics (PHY)
203 Classical Mechanics A
208 Principles of Quantum Mechanics
Psychology (PSY)
330 Introduction to Connectionist Models: Bridging between Brain and Mind
*336 The Diversity of Brains
Woodrow Wilson School (WWS)
320 Human Genetics, Reproduction, and Public Policy (also MOL 320)**
327 Pharmaceutical Research and Health Policy**
Other courses offered by the University may be accepted with the approval of the departmental representative.
*One-time-only course or topic
**Policy course