Department of Ecology and Evolutionary Biology

  • Chair

    Lars O. Hedin

  • Departmental Representative

    Daniel I. Rubenstein

  • Director of Graduate Studies

    Bryan T. Grenfell

  • Professor

    Joshua M. Akey, also Lewis-Sigler Institute for Integrative Genomics

    Andrew P. Dobson

    Bryan T. Grenfell, also Woodrow Wilson School

    Lars O. Hedin, also Princeton Environmental Institute

    Simon A. Levin

    Stephen W. Pacala

    Daniel I. Rubenstein

    David S. Wilcove, also Woodrow Wilson School, Princeton Environmental Institute

     

  • Associate Professor

    Andrea L. Graham

    Robert M. Pringle

    Corina E. Tarnita

  • Assistant Professor

    Julien Ayroles, also Lewis-Sigler Institute for Integrative Genomics

    Carolyn S. McBride, also Princeton Neuroscience Institute

    C. Jessica E. Metcalf, also Woodrow Wilson School

    Christina P. Riehl

    Mary C. Stoddard

    Bridgett M. vonHoldt

  • Associated Faculty

    Asif A. Ghazanfar, Psychology, Princeton Neuroscience Institute

    Bess B. Ward, Geosciences, Princeton Environmental Institute

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.

Information and Department Plan of Study

Required core courses:

All EEB concentrators must take EEB 309 and EEB 321 in the fall of their junior year or earlier.

Departmental areas of study:

Students must successfully complete a minimum of eight upper-level departmental courses at least six of which must be EEB courses. The first semester of organic chemistry and biochemistry (MOL 345) can each be counted as a departmental course (the second semester of organic chemistry will not count as a departmental). Other MOL courses might count as an EEB departmental with prior approval of the departmental representative.

Students can choose their departmental courses from six areas of study as listed below, but must ensure that their courses represent at least 4 different areas of study (the EEB 309 and 321 core courses count towards Evolution and Genomics and Ecology and Environment, respectively).  Students can find sample sequences for each area of study on the EEB website.

(i) Ecology and Environment: EEB 321; EEB 308; *EEB 346; *EEB 338; EEB 417.

(ii) Evolution and Genomics:  EEB 309; EEB 326; EEB 409;

(iii) Behavior and Organismal Biology:  EEB 311; EEB 313; EEB 314; EEB 329; EEB 403; *EEB 404; *EEB 406.

(iv) Disease Ecology:  EEB 304; EEB 327; *EEB 328; EEB 351.

(v) Conservation Biology:  EEB 308; *EEB 380; EEB 417.

(vi) Mathematical and Computational Biology:  EEB 324; EEB 325;

* offered as part of semester abroad program (described below)

Laboratory requirement

EEB students must complete an upper-level laboratory course. In most cases, this will be satisfied by the core course EEB 321.

No course taken P/D/F can be counted as a departmental or to satisfy requirements of the major. 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 – these can be counted as departmentals, but the grades do not transfer.

Integrated Biology Sequence: Students who have completed the two-year sequence need to meet with the EEB departmental representative to discuss placement in upper-level EEB courses.

Pre-medical students:

Students interested in medical school should consider taking EEB 314 Comparative Physiology, which will count towards one of the four required areas of study. In addition, medical schools require two terms of physics; physics can be delayed to senior year if necessary. The Health Profession advisers recommend that students wanting to study abroad, especially in EEB’s field semesters, should do so. They recommend contacting them early to formulate a plan of study that allows students to fulfill the pre-medical prerequisites and allows them to study abroad during spring of the junior year.

Advanced Placement

Advanced placement will be granted to students who received a 5 on the Biology AP exam. Nevertheless, all students planning on majoring in EEB must take EEB 211 freshman or sophomore year.

Prerequisites

The Department of Ecology and Evolutionary Biology requires the following prerequisites, which should normally be completed by the end of sophomore year.

  • Two terms of introductory biology: EEB 211 and either EEB/MOL 214 or EEB/MOL 215);
  • Mastery of calculus to the level of MAT 103 or above, 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);
  • A statistics course. SML 201 is preferred, but the following courses will fulfill the requirement: ECO 202, ORF 245, POL 345, PSY 251, or WWS 332. It is best to fulfill the statistics requirement before junior year, but it must be fulfilled by the end of the fall term senior year.

 

Prerequisite courses may not be taken using the P/D/F grading option.

Early Concentration

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.

Independent Work

Junior independent work:  Early in the fall semester, students will attend periodic meetings on Tuesday evenings that will include presentations by EEB faculty members on their research and on opportunities for senior independent work. After the faculty presentations, which will take place early in the semester, students will identify those faculty with whom they are interested in working. If the faculty member agrees to take on a particular student, together they will develop a plan of study.

During the second half of the fall term, students participate in a tutorial in which they work closely with a faculty member to address a problem and write a first junior paper. In the spring semester, students carry out a program of independent research with their faculty adviser, which can include empirical or theoretical work. Students will summarize this research project in a second junior paper, which can be structured either as a research project, a literature review, or a research proposal.  The second junior paper is due in early May.

Senior Independent Work. During the fall of the junior year each student selects a senior thesis adviser (see above). 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.

Senior Departmental Examination

 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.

Study Abroad

Special Features of the Plan of Study. EEB offers two tropical field study programs: a semester in Panama, and a semester in Kenya. Four courses, built around experiential learning, are taken in sequence at each location typically during the spring of junior year. Please note that seniors, who are well along with the thesis are encouraged to consider fulfilling their last two courses spring of senior year by attending the first half of the semester in Kenya or Panama. Seniors who choose to do this would return to campus at spring break to complete the senior thesis. Details of these programs can be found on the department's website.

Princeton’s 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, parasitology, and anthropology (focusing on Pre-Columbian peoples and their land-use practices).

2. Kenya. This four-course program on Tropical Biology and Sustainability, 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, other appropriate faculty from Princeton University, and faculty from Columbia University; Columbia students participate in the program. The courses delve into the ecology of savannas, conservation in Africa, the natural history of mammals, tropical agriculture, engineering and field hydrology and paleoecology.

3. 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.

Additional Information

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 211 Life on Earth: Chaos and Clockwork of Biological Design (also
MOL 211
) Fall STL
An examination of how life evolved and how organisms function. Design--'intelligent' and otherwise--will provide a unifying theme. Why do some microbes produce slime and others do not? Why are males brightly colored in some species, but in others females are the showy sex? Why do humans have knees that fail whereas horses and zebras do not? These and other 'why is it so' questions related to the origin and history of life, genetic code, biochemistry, physiology, morphology and body plans, sex and reproduction, cooperation, and ecosystems will be explored. This course is required of all EEB majors and fulfills a requirement for medical school. J. Akey, A. Dobson
EEB 214 Introduction to Cellular and Molecular Biology (See MOL 214)
EEB 215 Quantitative Principles in Cell and Molecular Biology (See MOL 215)
EEB 255B Life in the Universe (See GEO 255B)
EEB 304 Disease Ecology, Economics, and Policy (See ENV 304)
EEB 306 Human Evolution (See ANT 206B)
EEB 308 Conservation Biology Fall STN Students will use ecological principles and policy analysis to examine conflicts between human activities such as farming, forestry, and infrastructure development, and the conservation of species and ecosystem services. Two lectures, one preceptorial. D. Wilcove
EEB 309 Evolutionary Biology Fall All life on Earth has evolved and continues to evolve. This course will explore evolution at both the molecular and organismal level. We will examine the features that are universal to all life and that document its descent from a common ancestor that lived over 3 billion years ago. Topics include the origin of life, the evidence for natural selection, methods for reconstructing evolutionary history using DNA, population genetics, genome evolution, speciation, extinction, and human origins. This course will provide you with the basic tools to understand how evolution works and can produce the incredible diversity of life on our planet. B. vonHoldt
EEB 311A Animal Behavior Fall STN An examination of the mechanisms and evolution of the behavior of humans 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 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, one three-hour laboratory. Prerequisite: 210 or 211. Staff
EEB 315 Human Adaptation (See ANT 215)
EEB 321 Ecology: Species Interactions, Biodiversity and Society Fall STL How do wild organisms interact with each other, their physical environments, and human societies? Lectures will examine a series of fundamental topics in ecology -- herbivory, predation, competition, mutualism, species invasions, biogeographic patterns, extinction, climate change, and conservation, among others--through the lens of case studies drawn from all over the world. Readings will provide background information necessary to contextualize these case studies and clarify the linkages between them. Precepts and fieldwork will explore the process of translating observations and data into an understanding of how the natural world works. R. Pringle
EEB 324 Theoretical Ecology Spring 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 325 Mathematical Modeling in Biology and Medicine Not offered this year How can mathematical modeling help to illuminate biological processes? This course examines major topics in biology through the lens of mathematics, focusing on the role of models in scientific discovery. Students will learn how to build and analyze models using a variety of mathematical tools. Particular emphasis will be placed on evolutionary game theory. Specific topics will include: the evolution of cooperation and of social behavior from bacteria to humans; the evolution of multicellularity; the somatic evolution of cancer; virus dynamics (within host and within populations); and multispecies interactions and the evolution of mutualisms. C. Tarnita
EEB 327 Immune Systems: From Molecules to Populations (also
MOL 327
/
GHP 327
) Fall STN
Why is there immunological polymorphism in animal populations? Why do immune systems work as they do? This course examines the theories of host-parasite coevolution, including optimal host resource allocation to immune defense in light of parasite counter-strategies, and assesses the empirical evidence by which these theories are tested. Students look at the evolutionary ecology of mechanisms used by immune systems to recognize and kill parasites, finding similarities across animal taxa. Finally, students will map immune mechanisms onto host phylogenies to understand the order in which different mechanisms arose over evolutionary time. A. Graham
EEB 328 Ecology and Epidemiology of Parasites and Infectious Diseases (also
GHP 328
) Spring STL
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. Limited to students in the Tropical Ecology Program in Panama. A. Dobson
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. Staff
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. Staff
EEB 350 Vertebrate Tropical Ecology Not offered this year 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 351 Epidemiology: An Ecological and Evolutionary Perspective (See GHP 351)
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: 211 and 321. D. Rubenstein
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)