Department of Ecology and Evolutionary Biology
Daniel I. Rubenstein
James L. Gould
Laura F. Landweber
Director of Graduate Studies
Andrew P. Dobson
Andrew P. Dobson
James L. Gould
Bryan T. Grenfell, also Woodrow Wilson School
Lars O. Hedin, also Princeton Environmental Institute
Leonid Kruglyak, also Lewis-Sigler Institute for Integrative Genomics
Laura F. Landweber
Simon A. Levin
Stephen W. Pacala
Daniel I. Rubenstein
David S. Wilcove, also Woodrow Wilson School
Robert M. May
William H. Schlesinger
Peter Andolfatto, also Lewis-Sigler Institute for Integrative Genomics
Iain D. Couzin
Andrea L. Graham
Asif A. Ghazanfar, Psychology, Princeton Neuroscience Institute
Alan E. Mann, Anthropology
Ignacio Rodríguez-Iturbe, Civil and Environmental Engineering
Bess B. Ward, Geosciences
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, disease, behavior, neuroscience, ecology, ecosystem biology, conservation, and climate change--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 department's website.
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 students. See the department's website for examples of recent student research activities.
Advanced placement will be granted to students who received a 5 on the Biology AP exam. This excuses EEB majors from taking 211.
The Department of Ecology and Evolutionary Biology requires the following prerequisites, which should normally be completed by the end of sophomore year. (Two exceptions, physics and statistics, are noted below.)
- Two terms of introductory biology (an AP score of 5 can substitute for EEB 211, but not the second introductory course chosen, either EEB/MOL 214 or EEB/MOL 215);
- Mastery of calculus to the level of MAT 102 or 103, or advanced placement (an AP score of 5 on the AB test or an AP score of 4 on the BC test);
- Two terms of introductory chemistry (or equivalent, or an AP score of 5);
- The first term of introductory physics (or the equivalent, or an AP score of 5). [Note that medical schools require two terms of physics. Physics can be delayed to the junior or senior year if necessary but preferably should be taken earlier].
- An introductory statistics course (EEB 355 is preferred and counts as a departmental; other appropriate, non-departmental courses include: ECO 202, ORF 245, POL 345, PSY 251, or SOC 301). The statistics requirement must be fulfilled by the end of fall term senior year. Because EEB 355 is offered in the spring, students planning to participate in the EEB spring program in either Panama or Kenya need to take EEB 355 in their spring of sophomore year or avail themselves of one of the non-departmental alternatives offered in the fall.
Students who are passionate about solving problems in the areas of ecology, evolution, behavior, conservation, and disease may apply for early concentration in ecology and evolutionary biology. Students having advanced placement in biology and having taken at least one upper-level EEB course, in addition to having completed many of the department's required pre- and co-requisite courses with grades of at least B+, are eligible. Early concentrators will begin independent work during the spring of their sophomore year. Students interested in early concentration should contact a departmental representative at the end of the freshman year or at the start of the sophomore year.
Requirements. The following courses are required for graduation:
1. Students in ecology and evolutionary biology 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 an EEB lab course; 314, 324, 417B, or MOL 350 and all field courses count as lab courses.
Choice of Courses. Students must take at least one course from each of the following three general areas:
Behavior and Physiology: 301, 311, 314, 323, 350, 404, PSY 315;
Evolution and Genetics: 309, 320, 414;
Ecology and Conservation: 308, 312, 321, 322, 324, 328, 338, 346, 352, 380, 417.
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; MAT 200, 201, 202, 203, 204; COS 126, 226); or two upper-level courses in an aligned area with permission of the departmental representative);
3. Integrated Biology Sequence: Students who complete the two-year sequence need only take EEB 211.
Students may take 309, 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 department's website.
Specializations. Students interested in specializing within the department should enroll in the courses listed below.
Theoretical biology: 309, 320, 321, 322, and 324
Field biology: 312, 321, and a tropical field program
Conservation biology: 308, 417, and a tropical field program
Medical biology: 314, 327, and 328
Human biology: 301, 306, 315, PSY 330, and 336
Note: MOL 214 or 215 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; both terms count as EEB departmentals. 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 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 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. The program also includes courses on coral reefs, vertebrate tropical biology, and anthropology (focusing on Pre-Columbian peoples and their land-use practices). Prerequisite: EEB 321 or an equivalent introductory ecology course.
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 the ecology of savannas, conservation in Africa, the natural history of mammals, global technology and engineering and field hydrology. 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. Prerequisite: EEB 211.
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 in 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.
EEB 210 Evolutionary Ecology (also MOL 210) Not offered this year STL
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 STL
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
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 GSS 301) Spring STN
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. Staff
EEB 302 Advanced Analysis of Environmental Systems (see ENV 302)
EEB 304 Disease Ecology, Economics, and Policy (see ENV 304)
EEB 305 Field Ecohydrology (see CEE 307)
EEB 306 Human Evolution (see ANT 206)
EEB 308 Conservation Biology Spring
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 other topics, such as the origin of life and 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. Two lectures, one preceptorial. Prerequisite: 210 or 211. L. Landweber
EEB 311 Animal Behavior Spring STN
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. Two 90-minute lectures, one preceptorial. J. Gould
EEB 312 Marine Biology (also ENV 312) STL
An intensive four-week course during June 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 Not offered this year
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, one three-hour laboratory. Prerequisite: 210 or 211. Staff
EEB 315 Human Adaptation (see ANT 215)
EEB 320 Molecular Evolutionary Genetics (also MOL 330) Not offered this year
How and where did life evolve? This course will discuss the evolution of the molecules that sustain life (DNA, RNA and proteins) at both the micro and macro evolutionary levels. We will explore the role of these molecules in the origin and continued evolution of life. Topics include the origin of eukaryotes and organelles, comparative genomics, population genetics, and human evolution. Two lectures, one preceptorial. 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. A. Dobson
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 (also WWS 490) 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. A. Dobson, B. Grenfell
EEB 332 Pre-Columbian Peoples of Tropical America and Their Environments (also LAS 350) Spring SA
An intensive course on the pre-European history of Amerind cultures and their environments in the New World tropics. Topics 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. Daily lectures, field trips, and laboratory experiences and incorporates methods and problems in field archaeology, paleoethnobotany and paleoecology, and archaeozoology. Limited to students in the Tropical Ecology Program in Panama. This course does not count as an EEB departmental. Prerequisite: EEB 321. Staff
EEB 336 The Diversity of Brains (see PSY 336)
EEB 338 Tropical Biology (also LAS 351) Spring STL
This intensive field course, 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; tropical climate and hydrology; biotic interactions; and contemporary and historical factors in shaping tropical landscapes, with emphasis on the Isthmian Landbridge and subsequent floral and faunal interactions. Two hours of lecture/discussion, six hours of laboratory, and two hours of data analysis daily. Limited to students in the Tropical Ecology Program in Panama. Prerequisite: 321. Y. Basset
EEB 346 Biology of Coral Reefs Spring STL
This intensive field 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. Two hours of lecture/discussion, six hours of laboratory, and two hours of data analysis daily. Snorkeling in open ocean and walking in wild terrain is common. Limited to students in the Tropical Ecology Program in Panama. Prerequisite: 321. S. Pacala
EEB 350 Vertebrate Tropical Ecology Spring
This intensive 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. Two hours of lecture/discussion, six hours of laboratory, and two hours of data analysis daily. Limited to students in the Tropical Ecology Program in Panama. Prerequisite: 321. Staff
EEB 355 Introduction to Statistics for Biology (also MOL 355) Spring QR
An applied introduction to probability and statistical methods in biology. Topics include data visualization, parameter estimation, frequentist hypothesis testing, Bayesian hypothesis testing, useful statistical tests, linear models, and an introduction to data-driven modeling. Analyses will use an appropriate statistical computer package. J. Storey, P. Andolfatto
EEB 404 Natural History of Mammals Spring STL
Students examine how mammals interact with diverse and potentially conflicting features of their environment in order to understand the concepts, methods, and material of comparative natural history. Perspectives include morphology, identification, evolution, ecology, behavior, habitat, and conservation. Original observations and experiments culminate in class, group, and individual research projects. This intensive field course entails two hours of lecture/discussion, six hours of laboratory, and two hours of data analysis daily. Limited to students in the Tropical Ecology Program in Kenya. Prerequisite: 210 or 211. D. Rubenstein
EEB 414 Genetics of Human Populations (also MOL 414) Spring
This 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, P. Andolfatto
EEB 417A Ecosystems and Global Change (also ENV 417A) Not offered this year STN
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. Two 90-minute classes. L. Hedin
EEB 417B Ecosystems and Global Change (also ENV 417B) Not offered this year STL
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. 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