Biology
Ecology and Evolutionary Biology courses
Molecular, Cellular, and Developmental Biology courses
Ecology and Evolutionary Biology department home page
Molecular, Cellular, and Developmental Biology department home page
Directors of undergraduate studies: Leo Buss (Ecology and Evolutionary Biology), 101 OML, 432-3837, maureen.cunningham@yale.edu; Douglas Kankel (Molecular, Cellular, and Developmental Biology), 754 KBT, 432-3839, catherine.blackmon@yale.edu
FACULTY OF THE DEPARTMENT OF ECOLOGY AND EVOLUTIONARY BIOLOGY
Professors
Leo Buss, Michael Donoghue, *Jacques Gauthier, *Vivian Irish, *Kenneth Kidd, Jeffrey Powell, Richard Prum, *Oswald Schmitz, *David Skelly, Stephen Stearns, *J. Rimas Vaišnys, Günter Wagner
Associate Professor
Paul Turner
Assistant Professors
Suzanne Alonzo, Antonia Monteiro, Thomas Near, David Post, Melinda Smith, Jeffrey Townsend
Lecturers
Gisella Caccone, Dianella Howarth, Marta Martínez Wells
FACULTY OF THE DEPARTMENT OF MOLECULAR, CELLULAR, AND DEVELOPMENTAL BIOLOGY
Professors
Sidney Altman, *Kim Bottomly, Ronald Breaker, John Carlson, *Lynn Cooley, Stephen Dellaporta, Xing-Wang Deng, Paul Forscher, *Sankar Ghosh, *Mark Hochstrasser, Vivian Irish, Douglas Kankel, *Michael Kashgarian, Haig Keshishian, *Perry Miller, Mark Mooseker, *Jon Morrow, Timothy Nelson, L. Nicholas Ornston, Thomas Pollard, Shirleen Roeder, Joel Rosenbaum, *Alanna Schepartz, Michael Snyder, Robert Wyman
Associate Professors
Craig Crews, Savithramma Dinesh-Kumar, Christine Jacobs-Wagner, Frank Slack, *Hugh Taylor, Weimin Zhong
Assistant Professors
Thierry Emonet, Martín García-Castro, Scott Holley, Elke Stein, David Wells
Lecturers
Carol Bascom-Slack, Iain Dawson, Mary Klein, Kenneth Nelson, *Aruna Pawashe, Barry Piekos, William Segraves, Joseph Wolenski
*A joint appointment with primary affiliation in another department or school.
The major in Biology is offered jointly by the Departments of Ecology and Evolutionary Biology (EEB) and Molecular, Cellular, and Developmental Biology (MCDB), providing students with opportunities to take courses in both departments. There are two principal areas of concentration in the major and two directors of undergraduate studies.
The science of biology is extremely broad, ranging across the domains of molecules, cells, organisms, and ecosystems. Moreover, biology explores questions of evolutionary history and the processes of evolutionary change as well as the mechanisms by which cells, organisms, and ecosystems function. Students majoring in Biology receive a thorough yet varied liberal education and preparation for professional careers in a diverse array of fields. Practical applications of biology include the development of pharmaceuticals, the practice of medicine, and the scientific bases for understanding the history and complexity of the environment and the need for its protection.
The major in Biology offers two areas of concentration. Area of concentration I, ecology and evolutionary biology, is intended for students wishing to concentrate in the basic sciences underlying ecological, evolutionary, organismal, computational, conservation, and environmental biology. Area of concentration II, molecular, cellular, and developmental biology, offers programs for students wishing to concentrate on molecular biology and genetics and their applications to problems in cell biology, development, neurobiology, and certain aspects of computational biology. Interdisciplinary opportunities are available within Area II in the biotechnology and neurobiology tracks.
Students who are primarily interested in plant science are invited to consult with an appropriate faculty member.
The Biology major offers opportunities for independent research in both laboratory and field work. With approval, research can be conducted under the supervision of faculty members in any Yale department. Some programs for study abroad are available to Biology majors and are especially appropriate for those in Area I; approved programs can fulfill some of the requirements for the major. Interested students should consult the director of undergraduate studies and the Office of International Education and Fellowship Programs.
Students majoring in Biology must take all courses in the major for a letter grade. College seminars do not count toward the requirements of the major. The Biology major should not be taken as one of two majors with Molecular Biophysics and Biochemistry.
Prerequisites. The basic science courses required of all Biology majors are MCDB 120a, E&EB 122a or b, and either MCDB 121La or E&EB 123Lb; CHEM 113, 114, or 118a taken with their appropriate laboratories as well as two terms of organic chemistry with laboratories (CHEM 125, 126L satisfies both chemistry requirements); two terms of PHYS 150a, 151b or higher; and one term of MATH 115a or b or above, but not MATH 190a. Note that the prerequisites fulfill most of the usual premedical science requirements. Students in Area I may, with permission of the director of undergraduate studies, substitute three course credits in geochemistry and geophysics, engineering, or mathematics for organic chemistry.
Placement. Students who have scored 710 or higher on the SAT subject matter Biology M test (formerly SAT II), or who have scored 5 on the Advanced Placement test in biology, may be exempt from taking MCDB 120a and its associated laboratory. Students scoring 5 on the Advanced Placement biology test may also be exempt from E&EB 122b and its associated laboratory, but beginning with the Class of 2010, students are no longer permitted to place out of E&EB 122a or b and 123Lb using the SAT subject matter test M or E. Students with equivalent scores on one of the corresponding chemistry tests may also be exempt from taking MCDB 120a, but should first discuss their preparation in biology with the director of undergraduate studies in MCDB.
Acceleration credit awarded in chemistry, mathematics, and physics, or completion of advanced courses in those departments, is accepted instead of the relevant prerequisites for the Biology major. Students who already have mathematics preparation equivalent to MATH 115a or b or higher are encouraged to take additional mathematics, such as MATH 120a or b, 222a or b, or 225a or b.
Placement in chemistry courses is arranged by the Chemistry department. Because the required chemistry courses are prerequisite to a number of biology courses, students are strongly urged to take general and organic chemistry in the freshman and sophomore years. Students who place out of general chemistry should take organic chemistry during their freshman year. Finishing the prerequisites early allows for a more adventurous program in later years.
Requirements of the major. Beyond the prerequisites, the B.A. degree requires seven lecture courses or seminars, two laboratories, and the senior requirement (E&EB 470a or b, MCDB 470a or b, E&EB 475a or b, MCDB 475a or b, or the senior essay); the B.S. degree requires seven lecture courses or seminars, two laboratories, and the senior requirement (two terms of E&EB 475a or b or MCDB 475a or b or 485). The intensive B.S. degree requires, in addition to the prerequisites, seven lecture courses or seminars, two laboratories, and the senior requirement (E&EB 495 or MCDB 495).
Core requirements for Area I, ecology and evolutionary biology, Class of 2010 and subsequent classes. Area I majors in the Class of 2010 and subsequent classes are required to take MCDB 202a, E&EB 210a or the equivalent, E&EB 220a, and 225b.
Electives for Area I, ecology and evolutionary biology, Class of 2010 and subsequent classes. In addition to the four core courses, students in the Class of 2010 and subsequent classes must take three electives, one of which must be a course in organismal diversity chosen from E&EB 246b, 250a, <255b>, 257b, 264a, 272b, or MCDB 290b. Students who wish to take electives from other departments should obtain the approval of the director of undergraduate studies in EEB.
Core requirements for Area I, ecology and evolutionary biology, Class of 2009 and previous classes. Students in the Class of 2009 and previous classes may fulfill the same core requirements as the Class of 2010, or they may take the core courses E&EB 160a, MCDB 202a, and E&EB 210a.
Electives for Area I, ecology and evolutionary biology, Class of 2009 and previous classes. Members of the Class of 2009 and previous classes must take four electives, including at least one course from each of the following three subject areas:
Group A, ecology: E&EB 220a, 230a, 326a, 360b, 365a, 370a
Group B, evolution: E&EB 225b, 316La, G&G 250a
Group C, organisms: E&EB 246b, 250a, <255b>, 257b, 264a, 272b, MCDB 290b
In Groups A and B, the course listed first is the recommended first course in that subject. One additional EEB or MCDB course numbered 140 or higher is also required. Students who wish to draw on electives in other departments should obtain the approval of the director of undergraduate studies in EEB.
Laboratory requirement. For all classes, two laboratories beyond the prerequisites are required for the major.
Core requirements for Area II, molecular, cellular, and developmental biology. Biology majors in Area II are required to take MCDB 202a, 300a or its equivalent, and either 205b or 210a. (MCDB 210a is not a core course for the biotechnology track.) Equivalents for MCDB 300a are defined as either (a) both MB&B 300a and 301b or (b) MB&B 300a only, if the student took MCDB 120a or 200b prior to 300a or has the permission of the director of undergraduate studies in MCDB.
Electives for Area II, molecular, cellular, and developmental biology. In addition to the prerequisites and core requirements, the standard degree program requires four electives, including three from MCDB (numbered 150 or above), EEB (numbered 140 or above), or MB&B (numbered 200 or above), and one upper-level MCDB elective numbered 350 or higher.
Two laboratories at the 200 level or higher are also required for the standard program, one of which can be selected from courses in EEB or MB&B.
The neurobiology track requires MCDB 320a and three courses chosen from BENG 410a, CPSC 475a, MCDB 240b, 310a, 315b, 330a, 410a, 415b, 425a, 440b, 460b, PSYC 176a, 270b, 320a, and either MCDB 215a or PSYC 200a. (Students should note that PSYC 110a or b is a prerequisite for many psychology courses but does not substitute as an elective in the neurobiology track.) Two laboratories chosen from MCDB courses are also required as electives. Students interested in the neurobiology track should consult an adviser for the track.
Neurobiology track advisers:
P. Forscher, 222 KBT (432-6344)
H. Keshishian, 640A KBT (432-3478)
E. Stein, 232 KBT (432-8402)
D. Wells, 226 KBT (432-3481)
R. Wyman [F], 610A KBT (432-3475)
W. Zhong, 616 KBT (432-9233)
The biotechnology track requires MCDB 370b and three courses chosen from any MCDB course numbered 150 or above, MB&B 405a, 420a, 421b, 443b, BENG 351a, 352b, 410a, 457b, 464b, CENG 210a, 411a, 412b, CPSC 437a, 445b, 470a, or 475a. Two laboratories are required: either two from MCDB (including at least one from MCDB 341La or Lb to MCDB 345Lb) or BENG 355L or CENG 412b. Students interested in the biotechnology track should consult an adviser for the track.
Biotechnology track advisers:
R. Breaker, 506 KBT (432-9389)
X. W. Deng, 352B OML (432-8908)
K. Nelson, 725 KBT (432-5013)
M. Snyder, 926 KBT (432-6139)
J. Wolenski, 330 KBT (432-6912)
Many of the courses in other departments have prerequisites that can be substituted for an upper-level elective with permission of the MCDB director of undergraduate studies.
For all tracks in Area II, if both MCDB 205b and 210a are taken, one counts as a core course and the other as an elective. If both MB&B 300a and 301b are taken, one counts as a core course (in place of MCDB 300a) and one as an elective. Two laboratory courses from MCDB 342La, 343La, 344Lb, and 345Lb can be used together as one elective credit. If used as an elective, these laboratories cannot also fulfill the laboratory requirement. A relevant intermediate or advanced course from another department in science, mathematics, or statistics may be accepted as an elective with permission of the director of undergraduate studies. College seminars cannot be substituted for electives. Students should be aware that many of the courses listed above from other departments have prerequisites.
Senior requirement. In addition to the course requirements described above, all students must satisfy a senior requirement undertaken during the senior year. A booklet listing the requirements of each track and degree is available in the office of the director of undergraduate studies (101 OML for Area I, 754 KBT for Area II). All students must fill out a checklist of requirements and go over it with the undergraduate registrar, Maureen Cunningham (Area I) or Catherine Blackmon (Area II), by the spring term of their junior year. For the B.A. degree the requirement can be met in any one of three ways: by submitting a senior essay of fifteen to twenty pages evaluating current research in a field of biology; by successful completion of one term of tutorial work (E&EB 470a or b or MCDB 470a or b); or by successful completion of one term of individual research (E&EB 475a or b or MCDB 475a or b).
A senior choosing to fulfill the requirement with a senior essay must consult with a faculty adviser on the scope and literature of the topic and submit the adviser's written approval to the appropriate director of undergraduate studies no later than the course selection period of the term in which the paper is due. The senior essay may be related to the subject matter of a course, but the essay is a separate departmental requirement in addition to any work done in a course and does not count toward the grade in any course. The senior essay must be completed and submitted to the office of the director of undergraduate studies by the last day of classes. Students electing this option should obtain an approval form from the office of the director of undergraduate studies.
B.S. degree. The requirements for the B.S. degree are the same as for the B.A. degree except for the senior requirement, which differs in its greater emphasis on individual research. The senior requirement is two consecutive terms of E&EB 475a or b or MCDB 475a or b, at least one of which must be taken during the senior year, or MCDB 485. Students are expected to spend ten hours per week conducting individual research. Ordinarily, both terms of E&EB 475a or b or MCDB 475a or b are taken during the senior year, but it is possible for a student to begin work toward the senior requirement in the spring of the junior year, continue it over the summer, and complete it during the final year, an arrangement that may be particularly useful for students doing fieldwork. Yale College does not grant academic credit for summer research unless the student is enrolled in an independent research course in Yale Summer Session.
Intensive B.S. degree. Requirements for the intensive B.S. degree are the same as for the B.A. degree except that students fulfill the senior requirement by taking E&EB 495 or MCDB 495, Intensive Research, which earns four course credits. Seniors in the intensive major are expected to spend twenty hours per week conducting individual research.
It should be noted that the research courses E&EB 475a or b and 495 and MCDB 475a or b, 485, and 495 exist primarily to fulfill the senior requirement. Some students may wish to take E&EB 475a or b or MCDB 475a or b earlier in their course of study. This contributes to the thirty-six course credits required for graduation, but does not substitute for any of the other requirements of the major including the senior requirement. Students may take up to three credits in E&EB 475a or b and MCDB 475a or b during their undergraduate career.
Laboratory preparation for research. Students concentrating in Area II or with an interest in molecular evolution and who are planning to undertake research are advised to take one or more of the following laboratories first: MCDB 341La or Lb, either MCDB 342La or E&EB 316La, MCDB 343La, 344Lb, or 345Lb.
Combined B.S./M.S. degree program. Exceptionally able and well-prepared students may accelerate their professional education by completing a course of study leading to the simultaneous award of the B.S. and M.S. degrees after eight terms of enrollment. Completion in fewer than eight terms is not allowed. The requirements are as follows:
1. Candidates must satisfy the Yale College requirements for the B.S. degree. In addition to the three or four core requirements (depending on track) specified for the standard major, the three or four electives must be graduate-level courses designated "G" in the printed YCPS. One of these is a graduate seminar selected with the approval of the director of undergraduate studies. Students must earn a grade of A or A– in two graduate-level courses and a grade of B– or higher in the rest.
2. Six credits outside the major must be taken in the last two years, and at least two undergraduate courses in the last two terms.
3. In addition to the courses specified above, students must complete two graduate research courses for six course credits: (a) MCDB 585b or E&EB 585b, a two-credit course typically taken in the second term of the junior year. At the start of the course, each student forms a committee comprised of the adviser and two faculty members that meets to discuss the research project. Two of the members of this committee must be members of either the MCDB or EEB faculty, as appropriate to the thesis topic. At the end of the course, the student completes a detailed prospectus describing the thesis project and the work completed to date. The committee evaluates an oral and written presentation of the prospectus and whether the student may continue in the combined program; (b) MCDB 595/E&EB 595, a four-credit, yearlong course that is similar to MCDB 495 and E&EB 495 and is taken during the senior year. During the course, the student gives an oral presentation describing the work. At the end of the course, the student is expected to present his or her work to the department in the form of a poster presentation. In addition, the student is expected to give an oral thesis defense, followed by a comprehensive examination of the thesis conducted by the thesis committee. Upon successful completion of this examination, as well as all other requirements, the student is awarded the combined B.S./M.S. degree.
Students must also satisfy the requirements of Yale College for the simultaneous award of the bachelor's and master's degrees, including the following:
1. Students must apply in writing to the appropriate director of undergraduate studies and obtain departmental approval no later than the beginning of the second term of their junior year. Applications must be submitted by November 1 in the preceding term. Students must have the approval of both the director of undergraduate studies and the director of graduate studies to receive graduate credit for the graduate courses they select.
2. Students must have two-thirds A or A– grades in all of their courses and two-thirds A or A– grades in Biology courses, including prerequisites, in order to be admitted to the program.
Advising. Freshmen considering a major in Biology are invited to consult with one of the two directors of undergraduate study and/or a faculty member in EEB or MCDB who is a fellow of their residential college. Once an area of concentration is chosen, students should find a faculty adviser in the appropriate department. Students in the neurobiology or biotechnology track should consult an adviser for their track (listed above). Students in EEB should consult one of the advisers assigned to their class. The course schedules of all Biology majors (including sophomores intending to major in Biology) must be signed by a faculty member in one of the two participating departments; the signature of the director of undergraduate studies is not required. Students whose regular adviser is on leave can consult the director of undergraduate studies to arrange for an alternate.
Area I, ecology and evolutionary biology.
Class of 2008:
J. Powell, 170 ESC (432-3887)
M. M. Wells, 401 OML (432-6294)Class of 2009:
D. Post, 426B OML (432-3005)
R. Prum [Sp], 164 ESC (432-9423)
J. Townsend [F], 226 OML (432-4646)Class of 2010:
A. Monteiro, 326A OML (432-3109)
T. Near, 370A ESC (432-3002)Class of 2011:
L. Buss, 326B OML (432-3837)
Area II, molecular, cellular, and developmental biology. Any member of the MCDB department can serve as a faculty adviser to majors. College faculty advisers available to freshmen:
| BK | J. Wolenski, M. García-Castro | MC | X. W. Deng, H. Keshishian, K. Nelson, T. Pollard, F. Slack |
|
| BR | S. Dinesh-Kumar | PC | J. Carlson, C. Crews | |
| CC | M. Mooseker, R. Wyman [F] | SY | C. Jacobs-Wagner, S. Roeder | |
| DC | P. Forscher, V. Irish [Sp], L. N. Ornston, W. Zhong |
SM | S. Dellaporta,
D. Kankel, J. Rosenbaum, M. Snyder |
|
| TD | S. Holley | ES | E. Stein, D. Wells | |
| JE | T. Nelson, R. Breaker | TC | S. Altman [F] | |
REQUIREMENTS OF THE MAJOR
Prerequisites: MCDB 120a, E&EB 122a or b; either MCDB 121La or E&EB 123Lb; CHEM 113, 114, or 118a, with labs, and 2 terms of organic chem with labs (CHEM 125, 126L satisfies both reqs); PHYS 150a, 151b, or above; MATH 115a or b or higher (except MATH 190a); all courses taken for letter grades
Number of courses: B.A.—7 courses and 2 labs beyond prereqs taken for letter grades, totaling 8 or 9 course credits, incl senior req; B.S.—7 courses and 2 labs taken for letter grades, totaling 10 course credits, incl senior req; Intensive B.S.—7 courses and 2 labs taken for letter grades, totaling 12 course credits, incl senior req
Specific courses required: Area I: Ecology and evolutionary biology, Class of 2010 and later—E&EB 210a or equivalent, E&EB 220a, 225b, MCDB 202a; Area I: Ecology and evolutionary biology, Class of 2009 and earlier—E&EB 160a, MCDB 202a, E&EB 210a; Area II: Molecular, cellular, and developmental biology—Standard track—MCDB 202a, 300a, and either 205b or 210a; Neurobiology track—MCDB 202a, 300a, 320a, and either 205b or 210a; Biotechnology track—MCDB 202a, 205b, 300a, and 370b
Distribution of courses: Area I: Ecology and evolutionary biology, Class of 2010 and later—3 electives, 1 in organismal diversity; Area I: Ecology and evolutionary biology, Class of 2009 and earlier—4 electives, incl 1 from each of 3 elective groups; Area II: Molecular, cellular, and developmental biology—Standard track—3 electives from EEB numbered 140 or above, MCDB numbered 150 or above, or MB&B numbered 200 or above, and 1 addtl MCDB course numbered 350 or above; Neurobiology and biotechnology tracks—3 electives as specified
Substitution permitted: Area I—3 course credits in geochemistry and geophysics, engineering, or math for organic chem, with DUS permission; Area II—1 course relevant to biology in another dept, with DUS permission, except for college sems; higher-level courses for lower-level courses, with approval of adviser or DUS
Senior requirement: B.A.—MCDB 470a or b, or E&EB 470a or b, or MCDB 475a or b, or E&EB 475a or b, taken in senior year, or senior essay; B.S.—2 terms of MCDB 475a or b or E&EB 475a or b, at least 1 in senior year, or MCDB 485; Intensive B.S.—MCDB 495 or E&EB 495
Because the length of laboratory sessions depends on the particular experiment, only the starting times of some laboratory courses are given in the course listings. Students should allow several hours for each laboratory. For the Class of 2008, courses in EEB and MCDB count toward the natural science requirement unless otherwise indicated.
