Biological Sciences

Faculty of the Department of Molecular and Cellular Biology

Douglas A. Melton, Professor of Molecular and Cellular Biology (Chair of MCB)
Howard C. Berg, Professor of Molecular and Cellular Biology and Professor of Physics
Daniel Branton, Higgins Research Professor of Biology
John S. Chant, Associate Professor of Biochemistry and Molecular Biology
Lauraine A. Dalton, Preceptor in Molecular and Cellular Biology
James E. Davis, Senior Lecturer on Chemistry and Chemical Biology and on Molecular and Cellular Biology
John E. Dowling, Harvard College Professor and the Maria Moors Cabot Professor of Natural Sciences
Catherine Dulac, Assistant Professor of Molecular and Cellular Biology
Brian D. Dynlacht, Associate Professor of Molecular and Cellular Biology
Raymond L. Erikson, American Cancer Society Professor of Cellular and Developmental Biology
William Techumseh Fitch, Lecturer on Biology
William D. Fixsen, Senior Lecturer on Molecular and Cellular Biology
William M. Gelbart, Professor of Molecular and Cellular Biology (Head Tutor, Biology)
Walter Gilbert, Carl M. Loeb University Professor (on leave spring term)
Guido Guidotti, Higgins Professor of Biochemistry
Stephen C. Harrison, Higgins Professor of Biochemistry
J. Woodland Hastings, Paul C. Mangelsdorf Professor of Natural Sciences (on leave spring term)
Craig P. Hunter, Assistant Professor of Molecular and Cellular Biology
Nancy Kleckner, Professor of Biochemistry and Molecular Biology
Jeremy R. Knowles, Amory Houghton Professor of Chemistry and Biochemistry and Dean of the Faculty of Arts and Sciences
Samuel M. Kunes, John L. Loeb Associate Professor of the Natural Sciences
Richard M. Losick, Maria Moors Cabot Professor of Biology
Robert Lue, Senior Preceptor in Molecular and Cellulaar Biology
Tom Maniatis, Thomas H. Lee Professor of Molecular and Cellular Biology
Andrew P. McMahon, Professor of Molecular and Cellular Biology
Markus Meister, Jeff C. Tarr Professor of Molecular and Cellular Biology
Matthew S. Meselson, Thomas Dudley Cabot Professor of the Natural Sciences
James L. Michel, Assistant Professor of Medicine (Medical School) (Head Tutor, Biochemical Sciences)
Venkatesh N. Murthy, Assistant Professor of Molecular and Cellular Biology (Medical School)
Robert E. Pruitt, Associate Professor of Biology (on leave fall term)
Elizabeth J. Robertson, Professor of Molecular and Cellular Biology
Jack L. Strominger, Higgins Professor of Biochemistry
James C. Wang, Mallinckrodt Professor of Biochemistry and Molecular Biology
Don C. Wiley, John L. Loeb Professor of Biochemistry and Biophysics, Tutor in Biochemical Sciences (on leave fall term)

Associate Member of the Department of Molecular and Cellular Biology

Stuart L. Schreiber, Morris Loeb Professor of Chemistry and Chemical Biology (on leave 1999-00)

Faculty of the Department of Organismic and Evolutionary Biology

Daniel L. Hartl, Professor of Biology (Chair of OEB)
Peter S. Ashton, Charles Bullard Professor of Forestry
David A. Baum, Associate Professor of Biology
Fakhri A. Bazzaz, Mallinckrodt Professor of Biology
Andrew A. Biewener, Professor of Biology
Kenneth J. Boss, Professor of Biology
Colleen M. Cavanaugh, Professor of Biology
A. W. Crompton, Fisher Professor of Natural History (on leave fall term)
Michael J. Donoghue, Professor of Biology
Brian D. Farrell, John L. Loeb Associate Professor of the Natural Sciences, Associate Curator in Entomology
David R. Foster, Senior Lecturer on Biology
David A. Haig, Associate Professor of Biology
James Hanken, Professor of Biology
N. Michele Holbrook, Thomas D. Cabot Associate Professor of Organismic and Evolutionary Biology
Farish A. Jenkins, Jr., Alexander Agassiz Professor of Zoology in the Museum of Comparative Zoology and Professor of Biology, and Professor of Anatomy in the Harvard-MIT Division of Health Sciences and Technology (Medical School)
Andrew H. Knoll, Professor of Biology
George Varick Lauder, Professor of Biology
Richard C. Lewontin, Alexander Agassiz Professor of Zoology in the Museum of Comparative Zoology and Professor of Biology
Karel F. Liem, Henry Bryant Bigelow Professor of Ichthyology
Charles R. Marshall, Professor of Biology and of Geology
James J. McCarthy, Alexander Agassiz Professor of Biological Oceanography in the Museum of Comparative Zoology
Stephen R. Palumbi, Professor of Biology
Donald H. Pfister, Asa Gray Professor of Systematic Botany (on leave fall term)
Naomi E. Pierce, Sidney A. and John H. Hessel Professor of Biology (on leave fall term)
Kerry L. Shaw, Associate Professor of Biology
Otto T. Solbrig, Bussey Professor of Biology
P. Barry Tomlinson, Edward C. Jeffrey Professor of Biology
John R. Wakeley, Assistant Professor of Biology
Robert M. Woollacott, Professor of Biology (on leave spring term)

Other Faculty Offering Instruction in Biological Sciences

William H. Bossert, David B. Arnold, Jr. Professor of Science
George M. Church, Professor of Genetics (Medical School)
Charles A. Czeisler, Associate Professor of Medicine (Medical School)
John M. Doyle, John L. Loeb Associate Professor of the Natural Sciences
Timothy F. Flannery, Visiting Professor of Australian Studies
Melvin J. Glimcher, Harriet M. Peabody Professor of Orthopedic Surgery (Medical School)
Stephen J. Gould, Alexander Agassiz Professor of Zoology in the Museum of Comparative Zoology and Professor of Geology
Arthur L. Lage, Associate Professor of Veterinary Medicine (Medical School)
Ralph Mitchell, Gordon McKay Professor of Applied Biology (on leave spring term)
Hidde Ploegh, Mallinkrodt Professor of Immunopathology and Professor of Pathology (Medical School)

The courses designated Biological Sciences are an introductory sequence in the Biochemical Sciences and Biology concentrations. They are jointly organized by the departments of Molecular and Cellular Biology and Organismic and Evolutionary Biology. The courses designated Molecular and Cellular Biology are the responsibility of that department. The courses designated Biology are the responsibility of the Department of Organismic and Evolutionary Biology. The Head Tutor for the Biochemical Sciences concentration is James L. Michel. The Head Tutor for the Biology concentration is William M. Gelbart.

Biological Sciences

Primarily for Undergraduates

Biological Sciences 1. Introductory Genetics, Molecular, Cellular, and Developmental Biology
Catalog Number: 1812
William M. Gelbart, William D. Fixsen, and Robert Lue
Half course (spring term). M., W., F., at 12, and three hours of laboratory each week. EXAM GROUP: 5
Principles of biology, with special reference to cellular and molecular mechanisms, genetics, and development. Emphasis on cells as evolved systems for the capture and transformation of energy and the processing of information.
Note: Lectures and weekly laboratory/discussion sessions.
Prerequisite: Chemistry 5, Chemistry 10, or permission of instructor.

Biological Sciences 2. Organismic and Evolutionary Biology
Catalog Number: 1922
P. Barry Tomlinson, Andrew A. Biewener, and Brian D. Farrell
Half course (fall term). M., W., F., at 12, and three hours of laboratory each week. EXAM GROUP: 5
An evolutionary and integrative functional approach to plant and animal biology, emphasizing organism, populations, and their environment. Topics to be covered include basic morphology and physiology, population and community dynamics, ecology, principles of population genetics, and evolutionary theory.
Note: Knowledge of introductory molecular biology is strongly recommended (Biological Sciences 1 or equivalent).

Biological Sciences 10. Introductory Molecular Biology
Catalog Number: 1938
Richard M. Losick
Half course (fall term). M., W., F., at 10, and a one-hour meeting each week; approximately four afternoon laboratory sessions to be arranged over the course of the semester. EXAM GROUP: 3
The first semester of a year-long integrated introduction to the basic principles of biochemistry, molecular and cell biology. Topics covered: the biochemistry and molecular biology of nucleic acids; the Central Dogma; DNA, RNA, and protein synthesis; mutation and repair; recombination and transposition; the genetic code; the turning on and off of genes; RNA, ribozymes, and splicing; development.
Note: Chemistry 17 may be taken concurrently.
Prerequisite: Chemistry 17 or 20.

Biological Sciences 11. Basic Principles of Biochemistry and Cell Biology
Catalog Number: 8673
Guido Guidotti
Half course (spring term). Lectures M., W., F., at 10, and a one-hour meeting during which problem sets are solved; one laboratory session each week. EXAM GROUP: 3
An introduction to the structure and function of cells and the major experimental tools used in modern cell biology. Emphasis is placed on the biochemistry and organization of proteins, macromolecules, and organelles so that an understanding of intracellular and cell-to-cell regulation can be achieved. Topics include structure and function of proteins, biochemical catalysis, energy conversion, the maintenance of cellular compartments, and transmembrane and cell-cell signaling.
Prerequisite: Chemistry 27 or equivalent, but Chemistry 27 may be taken concurrently.

Biological Sciences 14. Genetics
Catalog Number: 6434
Daniel L. Hartl
Half course (spring term). Tu., Th., 8:30–10. EXAM GROUP: 12, 13
Provides a clear, comprehensive, and rigorous introduction to genetics. Maintains a balance between genetics as a body of knowledge pertaining to genetic transmission, function, and mutation (the principles) and genetics as an experimental approach to understanding complex biological mechanisms (genetic analysis). Principal goals: To understand the basic processes of gene transmission, mutation, regulation, and evolution. To formulate genetic hypotheses, deduce their consequences, and test predictions against observed data. To use the principles of probability and statistics in solving quantitative problems in genetics. To appreciate the social and historical context in which genetics has developed and is continuing to develop. To become familiar with genetic resources and information available through the Internet.

Biological Sciences 25 (formerly MCB 25 and Psychology 12). Behavioral Neuroscience
Catalog Number: 6052
John E. Dowling and Mark G. Baxter
Half course (spring term). Lectures, Tu., Th., 10–11:30, and a 90-minute section meeting to be arranged. EXAM GROUP: 12, 13
An introduction to the organization and function of the nervous system and its role in behavior. Topics include the cell biology of neurons, electrical and biochemical signaling by neurons, mechanisms of sensation and perception, control of movement, learning and memory, language, motivation and emotion. Surveys research on mental illness, neurological diseases, and computational models of brain function.
Note: Introductory Biology recommended.

Molecular and Cellular Biology

Primarily for Undergraduates

*Biochemical Sciences 91r. Introduction to Research
Catalog Number: 6083
James L. Michel (Medical School) and members of the Faculty
Half course (fall term; repeated spring term). Hours to be arranged.
Laboratory research in topics related to the Biochemical Sciences concentration under the direction of, or approved by, members of the Board of Tutors in Biochemical Sciences. A paper must be submitted by the end of the reading period both to the laboratory sponsor and to the Biochemical Sciences Tutorial Office for review by the Course Director and members of the Board of Tutors.
Note: Open only to Biochemical Sciences concentrators; written permission of both the tutor and laboratory sponsor must be submitted to the Biochemical Sciences Tutorial Office prior to enrolling in the course. This introductory research course is intended to prepare students for Biochemical Sciences 99, and may be repeated only with prior permission of the Chairman of the Board of Tutors. (Forms are available in the Tutorial Office.)

*Biochemical Sciences 99. Laboratory Research for Honors Thesis
Catalog Number: 6670
James L. Michel (Medical School) and members of the Faculty
Full course (indivisible). Hours to be arranged.
For honors candidates writing a thesis in Biochemical Sciences. Indivisible for students enrolling in the fall term. Students intending to enroll in the fall are required to submit a written proposal to the Head Tutor by the end of the preceding spring term. Students may enter the course at midyear only with the permission of the Head Tutor. (Students intending to enroll in the spring must submit a written proposal to the Biochemical Sciences Tutorial Office by the beginning of the fall term.) The thesis proposal must be approved by the Chairman of the Board of Tutors prior to enrolling in Biochemical Sciences 99.

MCB 16. Developmental Biology
Catalog Number: 0749
Andrew P. McMahon
Half course (fall term). M., W., 8:30–10. EXAM GROUP: 1
A comprehensive lecture course in developmental biology. The principles and mechanisms of animal development are emphasized and illustrated using several animal models. Emphasis placed on experimental approaches at the molecular, genetic, and cellular levels of organization. For example, we will discuss how the egg gives rise to an adult with a functioning skeleton, brain and other organs, how sex is determined and how these mechanisms are conserved from insects to humans.
Prerequisite: Biological Sciences 1. Biological Sciences 10 or 11 suggested but not required.

MCB 61. Physical Biochemistry
Catalog Number: 0190
James E. Davis
Half course (spring term). Tu., Th., 11:30–1, and one section per week. EXAM GROUP: 13, 14
An introduction to solution thermodynamics and kinetics as applied to molecular interactions in biochemistry. Includes an introduction to statistical thermodynamics, as well as treatment of multiple equilibria, ions in solution, interaction among small molecules and macromolecules, transport properties of macromolecules, enzyme catalyzed reactions, and spectroscopy.
Prerequisite: Mathematics 1a and 1b (Mathematics 21a recommended), Physics 11, Chemistry 7 or 10.

For Undergraduates and Graduates

Attention is called to the following course: Chemistry 170. Chemical Biology
MCB 112. Structure and Function of Proteins and Nucleic Acids
Catalog Number: 6380
Stephen C. Harrison
Half course (fall term). Tu., Th., 5–7 p.m. EXAM GROUP: 18
Principles of protein and nucleic acid structure and assembly. This course is especially intended for first year graduate students in MCB and Biophysics, but it is also appropriate for suitably prepared undergraduates.
Prerequisite: Biological Sciences 10 and 11 (or equivalent) and MCB 61, or equivalent preparation in physical chemistry.

*MCB 113. Principles of Genetic Analysis
Catalog Number: 0875 Enrollment: Limited to 25
Nancy Kleckner and members of the Faculty
Half course (spring term). Tu., Th., 2–4. EXAM GROUP: 16, 17
Advanced treatment of principles and methods of genetic analysis. Classical and molecular approaches are discussed as applied to a range of organisms from bacteria to man.
Prerequisite: Biological Sciences 10, 11, and 14 (or equivalents).

MCB 116. Experimental Embryology
Catalog Number: 1207 Enrollment: Limited to 24
Elizabeth J. Robertson
Half course (fall term). M., 2–6. EXAM GROUP: 7, 8, 9
An introduction to basic problems in developmental biology by direct experimentation. Both classical and modern molecular manipulations of developing embryos are performed to study cell specification, differentiation, organ formation, and embryonic induction. Various aspects of pattern formation are analyzed including the establishment of polarity and body axes, making use of frogs, chicken, mice and fish.
Note: Laboratory course primarily for advanced undergraduates and graduate students.
Prerequisite: Biological Sciences 10 or 11 or MCB 16.

MCB 117. Experimental Neuroscience
Catalog Number: 3175 Enrollment: Limited to 16
Markus Meister, John E. Dowling, Samuel M. Kunes, and Ken Nakayama
Half course (fall term). Tu., Th., 2–5. EXAM GROUP: 16, 17, 18
An inquiry-based approach to neuroscience that uses state of the art technology to study the development and function of the nervous system. Topics include neural development, growth cone behavior, properties of voltage gated channels, systems neurobiology, and psychophysics. Experimental approaches include immunohistochemistry, molecular genetics, time lapse video-enhanced microscopy, gene expression of channels, whole cell voltage clamp, and extracellular recording.
Note: Primarily for upper-level undergraduate and graduate students; background in biochemistry or physics suggested.
Prerequisite: Biological Sciences 25.

MCB 119. Experimental Genetics
Catalog Number: 4472 Enrollment: Limited to 15
John S. Chant
Half course (spring term). W., 12–3, F., at 3. EXAM GROUP: 5, 6, 7
The primary objective is to illustrate the principles and techniques of genetics with the most tractable eukaryotic genetic organism, budding yeast. Because yeast cells are used so widely in biology as the organism of choice or as an organism for working with heterologous proteins, the course also serves as an introduction for those who wish to use yeast in the future. Each week, a genetic experiment is performed. For each experiment, there is some modest background reading from research literature and a short lecture at the beginning of the laboratory period.

MCB 121. Animal Genetic Systems
Catalog Number: 9084 Enrollment: Limited to 20
Craig P. Hunter
Half course (spring term). M., W., 4–6. EXAM GROUP: 9
This course will focus on the development and use of model genetic systems (C. elegans, Drosophila, zebrafish, mouse, and human) to study animal development, physiology, and evolution. Introductory lectures will precede discussions and student presentations of the current and classical literature.
Prerequisite: Biological Sciences 10, 11, and 14 (or equivalents).

*MCB 122. Regulation of Cell Proliferation
Catalog Number: 1403 Enrollment: Limited to 20
Brian D. Dynlacht and Raymond L. Erikson
Half course (fall term). M., W., 4–6. EXAM GROUP: 9
The molecular and biochemical events occurring during the cell cycle are considered. Topics include growth factors and their receptors, second messengers, protein phosphorylation, gene expression, oncogenes, and tumor suppressor genes. The course consists of lectures and student presentations from the current literature. An optional laboratory section will be offered to students with limited laboratory experience interested in the techniques considered in the lectures.
Prerequisite: Biological Sciences 10 or equivalent.

[MCB 125. Introduction to Biophysics]
Catalog Number: 5593
Howard C. Berg and John M. Doyle
Half course (spring term). Hours to be arranged.
An introduction to physical processes that underlie the behavior of cells and organisms and the methods used for their study. Elements of probability and statistics, the random walk, and diffusion. Physics of fluid flow, mechanical design, locomotion, and flight. Biological effects of radiation.
Note: Expected to be given in 2000–01. Physics 1b, 11b or 15b and Biological Sciences 10 or 11 are very helpful but not required.
Prerequisite: Elementary calculus and one-half course in physics.

MCB 129. Molecular Genetics of Neuronal Development
Catalog Number: 8956 Enrollment: Limited to 25
Samuel M. Kunes
Half course (fall term). Tu., Th., 10:30–12. EXAM GROUP: 12, 13
Topics include the control of neuronal differentiation and cellular identity, cell birth and cell death, axon guidance and the mechanisms of synaptic specificity. Emphasis placed on genetic approaches to understanding the development of the nervous system.

MCB 138. Function of Neural Systems
Catalog Number: 1153 Enrollment: Limited to 25
Markus Meister
Half course (spring term). W., F., 10–11:30. EXAM GROUP: 3, 4
Introduction to the known functional principles of large neural circuits responsible for the processing of sensory input, learning and memory, and control of movement. Covers the physiological mechanisms underlying collective neural function and the methods useful in their analysis.
Prerequisite: Biological Sciences 25 and one half course in physics or permission of instructor.

MCB 141. Molecular and Developmental Neurobiology
Catalog Number: 5205
Catherine Dulac
Half course (spring term). Tu., Th., 8:30–10. EXAM GROUP: 10, 11
Molecular basis of sensory perception and formation of related neuronal networks during vertebrate development. Topics will include: mechanisms of sensory discrimination at the level of receptor molecules and receptor cells; coding of sensory information by the brain; establishment of appropriate connections in the developing brain. Molecular and genetic approaches to memory and behavior will be discussed.
Prerequisite: Biological Sciences 10 and 25.

MCB 142 (formerly MCB 242). Chromosomes
Catalog Number: 7948 Enrollment: Limited to 20
Matthew S. Meselson
Half course (spring term). W., 2–4. EXAM GROUP: 7, 8
Selected aspects of the structure, replication, segregation, recombination and function of chromosomes. Lectures, student presentations and critical discussion of the scientific literature.

*MCB 155. Molecular Mechanisms of Gene Control
Catalog Number: 6230
Tom Maniatis
Half course (fall term). Tu., Th., at 4pm, plus two-hour section times to be arranged. EXAM GROUP: 15
An advanced course on the control of gene regulation. Topics include mechanisms of gene regulation at the level of transcription, chromatin structure, DNA rearrangement, DNA methylation, RNA processing, mRNA localization, and protein systhesis and degradation. The course is taught through weekly lectures and readings from the current literature. Topics covered in lectures and the reading assignments are discussed in sections. Students are required to critically evaluate and discuss recent papers in sections. Term paper and two exams.
Prerequisite: Biological Sciences 10 and 11 (or equivalent), and permission of instructor.

MCB 169. Molecular and Cellular Immunology
Catalog Number: 2518
Hidde Ploegh (Medical School)
Half course (fall term). Tu., Th., 10–11:30, and a ninety minute discussion section per week to be arranged. EXAM GROUP: 12, 13
Basic elements of the immune system. Molecular biology of antigen recognition structures on B and T lymphocytes. Cellular and genetic basis of immunity. Regulation and development of the immune system.
Prerequisite: Biological Sciences 1 and 10. Genetics and Cell Biology strongly recommended.

MCB 176. Membrane Function
Catalog Number: 3186
Guido Guidotti
Half course (fall term). M., 2–4, and a two-hour session during which research papers are analyzed. EXAM GROUP: 7, 8
The structure of membrane proteins and their involvement in transport and signal transduction.
Prerequisite: Biological Sciences 10 and 11 or equivalent.

MCB 177. Macromolecular Assemblies in Genetic Processes
Catalog Number: 3102 Enrollment: Limited to 30
James C. Wang
Half course (spring term). M., 7–9 p.m. EXAM GROUP: 9
Selected topics on complex assemblies in replication, repair, recombination and transcription will be discussed. Emphasis is on the structural basis of an assembly and the macromolecular interactions underpinning its function.
Prerequisite: Biological Sciences 10 and 11 or equivalent, or permission of the instructor. Background in macromolecular structures and functions encouraged.

MCB 186. Circadian Biology: From Cellular Oscillators to Sleep Regulation
Catalog Number: 2854
J. Woodland Hastings and Charles A. Czeisler (Medical School)
Half course (fall term). W., 2-5, and a one hour discussion section per week to be arranged. EXAM GROUP: 7, 8, 9
Properties, mechanisms and functional roles of circadian (daily) rhythms in organisms ranging from unicells to mammals. Cellular and molecular components, regulation of gene expression and physiological functions, genetic and biochemical analyses of circadian rhythms, and neurobiology of the mammalian circadian pacemaker. Mathematics and modeling of oscillatory systems and applications to circadian rhythms. Experimental studies of human rhythms, including the sleep-wake cycle and hormone rhythms, with applications to sleep disorders.
Prerequisite: Biological Sciences 1 and 2; Biological Sciences 25 desirable.

Cross-listed Courses

Biophysics 101. Genomics and Computational Biology

Primarily for Graduates

MCB 212. Topics in Biophysics
Catalog Number: 3351
Howard C. Berg
Half course (spring term). F., 2:30–4. EXAM GROUP: 7, 8
Motility and sensory transduction: Chemotaxis in bacteria; flagellar motility; prokaryotic and eukaryotic motor molecules. A term paper and seminar will be required.

[*MCB 225. Topics in Neurobiology]
Catalog Number: 1149 Enrollment: Limited to 15.
John E. Dowling
Half course (fall term). Hours to be arranged.
An in-depth review of one or a few topics of current interest to the participants. Student seminars, occasional guest lecturers, and discussions.
Note: Expected to be given in 2000–01. Open to graduate students and undergraduates with special interests and training.
Prerequisite: Biological Sciences 25 or equivalent.

MCB 268. Molecular Immunology: Seminar
Catalog Number: 2196 Enrollment: Limited to 20.
Jack L. Strominger
Half course (spring term). M., Th., 4–6. EXAM GROUP: 9, 18
One session each week is a lecture on the topics. At the second session, four papers are read from the current literature; each is presented by a student in 20-30 minutes. Reading of the papers, seminar presentations, and class participation are the only work of the course.
Prerequisite: MCB 169 or permission of instructor required for undergraduates, but not essential for graduate students.

[MCB 289. Photobiology]
Catalog Number: 7087 Enrollment: Limited to 10.
Half course (spring term). Hours to be arranged.
Interaction of light with biological systems and the relevant principles of photochemistry and photophysics. Topics include vision, photosynthesis, bioluminescence, photoinactivation and mutation, photomovement and morphogenesis, photoperiodism, and photomedicine.
Note: Expected to be given in 2000–01.
Prerequisite: Biological Sciences 1 and Biological Sciences 11 or equivalent, and an introductory course in physics.

Graduate Courses of Reading and Research

*MCB 300. Introduction to Research
Catalog Number: 4816
Douglas A. Melton 7232 and James C. Wang 4870

*MCB 304. Biochemistry of Cell Cycle Regulation
Catalog Number: 2980
Brian D. Dynlacht 1940

*MCB 306. Biophysics and Physiology of Neurons
Catalog Number: 1695
Venkatesh N. Murthy (Medical School) 2424

*MCB 310. Molecular Embryology of the Mouse
Catalog Number: 8031
Elizabeth J. Robertson 3313

*MCB 312. Military and Arms Control Applications of Biology and Chemistry
Catalog Number: 2063
Matthew S. Meselson 1319

*MCB 314. Molecular Genetics of Cell-Cell Interactions in Plants
Catalog Number: 5564
Robert E. Pruitt 3376 (on leave fall term) (on leave fall term)

*MCB 322. Genetics and Development
Catalog Number: 7290
Craig P. Hunter 2803

*MCB 326. Biochemical Virology
Catalog Number: 0243
Raymond L. Erikson 7506

*MCB 344. Molecular and Developmental Neurobiology
Catalog Number: 2292
Catherine Dulac 2801

*MCB 351. Structural Molecular Biology
Catalog Number: 1977
Stephen C. Harrison 3597

*MCB 352. Structural Molecular Biology
Catalog Number: 2099
Don C. Wiley 3598 (on leave fall term)

*MCB 359. Molecular Genetics
Catalog Number: 6278
Nancy Kleckner 4697

*MCB 360. Proteins Involved in DNA Transactions
Catalog Number: 0407
James C. Wang 4870

*MCB 364. Vertebrate Development
Catalog Number: 1396
Andrew P. McMahon 3312

*MCB 365. Neurobiology
Catalog Number: 8349
John E. Dowling 3545

*MCB 369. Structure, Composition, and Function of Biological Membranes
Catalog Number: 4636
Daniel Branton 4139

*MCB 373. Cellular Biochemistry and Physiology
Catalog Number: 8053
J. Woodland Hastings 1311 (on leave spring term)

*MCB 374. Developmental Neurobiology
Catalog Number: 6167
Samuel M. Kunes 3486

*MCB 375. Molecular Biology of Cell Division
Catalog Number: 4991
John S. Chant 3485

*MCB 376. Molecular Biology
Catalog Number: 4159
Walter Gilbert 1306 (on leave spring term)

*MCB 377. Genetics and Development
Catalog Number: 5598
William M. Gelbart 4774

*MCB 378. Motile Behavior of Bacteria
Catalog Number: 5729
Howard C. Berg 1377

*MCB 381. Microbial Development
Catalog Number: 4994
Richard M. Losick 3561

*MCB 382. Molecular Immunology
Catalog Number: 5515
Jack L. Strominger 1193

*MCB 386. Molecular Evolution
Catalog Number: 0763
Matthew S. Meselson 1319

*MCB 390. Function of Neuronal Circuits
Catalog Number: 8883
Markus Meister 3007

*MCB 391. Biochemistry
Catalog Number: 4888
Guido Guidotti 1203

*MCB 395. Eukaryotic Gene Regulation
Catalog Number: 7697
Tom Maniatis 7231

*MCB 399. Vertebrate Developmental Biology
Catalog Number: 7699
Douglas A. Melton 7232

Biology


Certain courses at the Medical School, offered by the FAS through the Division of Medical Sciences (see listings in this catalog), may be counted for concentration credit by individual petition to the Biology Undergraduate Committee prior to filing the study card. These include courses in Biological Chemistry and Molecular Pharmacology, Cell Biology, Genetics, Immunology, Microbiology, Neurobiology, Pathology, and Virology.

With the approval of the Biology Undergraduate Committee, summer courses taken at the Marine Biological Laboratory, Woods Hole, Massachusetts, may be counted toward degree requirements, but the total credit so obtained shall not exceed one course. Similarly, and with the same limitations, courses taken in Tropical Biology under the auspices of the Organization for Tropical Studies may be counted toward degree requirements.

Primarily for Undergraduates

Biology 10. Biology of Invertebrate Animals
Catalog Number: 7873
Kenneth J. Boss
Half course (fall term). Lectures M., W., F., at 11; laboratory to be arranged. EXAM GROUP: 4
Surveys the diversity of invertebrate animals, including their natural history and probable evolutionary relationships. The ecological and behavioral adaptations as well as the morphology and systematics of these organisms are explored, using common marine, freshwater, terrestrial, and parasitic species.
Note: Biological Sciences 2 recommended, or permission of instructor required.

Biology 17. Evolution
Catalog Number: 1799
Andrew J. Berry
Half course (spring term). Tu., 8:30–10. EXAM GROUP: 10, 11
A study of the course and processes of organic evolution. Topics include the history of ideas of evolution, population genetics, population ecology, speciation, the concept of adaptation, coevolution, evolutionary rates, evolutionary convergences, mass extinctions, and biogeography.
Note: Complements Biology 19.
Prerequisite: Science B-15 or Biological Sciences 2, and some familiarity with elementary mathematical manipulations.

Biology 19. Population Biology: Ecology
Catalog Number: 3365
Fakhri A. Bazzaz
Half course (spring term). M., W., F., at 10, weekly discussion section, and a minimum of three field trips on Saturday and/or Sunday. EXAM GROUP: 3
Relationships of organisms to their environment at the individual, population, and community level. Topics in pure and applied ecology including adaptations to physical environment, competition, concept of the niche, population dynamics, predator-prey interactions, herbivore effects, community ecology, ecosystem structure and stability, and resource management.
Note: Complements Biology 17.
Prerequisite: Mathematics 1 and Science B-15 or Biological Sciences 2 or equivalent.

Biology 20. Biological Diversity
Catalog Number: 0761
David A. Baum and Andrew H. Knoll
Half course (spring term). Tu., Th., 10–11:30; one afternoon of laboratory/discussion each week. EXAM GROUP: 12, 13
An overview of the structural and functional diversity of organisms, with emphasis on recent advances in understanding phylogenetic relationships among the primary branches of life and the evolution of diversity through geological time. Emphasis placed on the origin and diversification of major groups, the maintenance of diversity in natural ecosystems, and the conservation of biodiversity.
Prerequisite: Biological Sciences 2, or permission of instructor.

Biology 21. Structure and Physiology of the Vertebrates
Catalog Number: 0921
Andrew A. Biewener
Half course (fall term). Lectures, M., W., F., at 9, and five hours of laboratory each week. EXAM GROUP: 2
Introduction to the relationship between physiology, structure, and function of vertebrates. Lectures concentrate on selected organ systems (musculoskeletal, cardiovascular, respiratory, reproductive, excretory, and digestive) paying particular attention to how they have evolved and diversified within living groups in relationship to environmental problems. General principles of structure-function relationship are emphasized. Physiology and morphological dissection labs concentrate on higher vertebrates and are correlated with one another and with the lectures.
Prerequisite: Biological Sciences 1 and 2.

Biology 22. Animal Behavior
Catalog Number: 2539
Tecumsch Fitch
Half course (fall term). Lectures, Tu., Th., at 10; evening movie section to be arranged; ninety minutes of discussion section each week. EXAM GROUP: 12
A review of the behavior of animals under natural conditions, with emphasis on both mechanistic and evolutionary approaches. Topics include motivation and behavioral endocrinology; development; behavioral genetics and learning; signaling, deceit and animal consciousness; orientation, migration and biological rhythms: optimization and game theory; parental investment and mating systems; selfishness, altruism, and reciprocity; and sociality in vertebrates and invertebrates.
Prerequisite: Science B-15, B-29 or Biological Sciences 1, 2, or Biology 20 or equivalent, or permission of instructor.

Biology 24. Biology of Plants
Catalog Number: 1343
N. Michele Holbrook and P. Barry Tomlinson
Half course (spring term). Tu., Th., 11:30–1, one afternoon laboratory per week, plus occasional field trips. EXAM GROUP: 13, 14
Introduction to the structure, diversity, and physiology of plants with an emphasis on evolutionary relationships and adaptations to life on land. Topics include growth, resource acquisition, interactions with other organisms (i.e., fungi, bacteria, insects), reproduction, and survival in extreme environments. Laboratory sessions provide an overview of plant and diversity and an introduction to basic physiological processes.

Tutorials, Independent Research, and Thesis Work

See Biology 121a, Biology 121b, and Biology 200r for other independent research options. Students who participate in independent research laboratory courses (Bio 98r, 99ar, 99b, 121a, 121b, or 200r) are required to attend a safety session on matters concerned with safety procedures before or at the beginning of their laboratory work. Credit for these courses is contingent upon compliance with this requirement. Biology 95hf: Tutorial for Credit course descriptions and syllabi are available in the Biology Undergraduate Office and on Biology’s Web site at http://www.mcb.harvard.edu. Students should enter on the study card the “Catalog Number” of the Tutorial in which they wish to enroll.
*Biology 91r. Supervised Reading
Catalog Number: 2817
William M. Gelbart and members of the Department
Half course (fall term; repeated spring term). Hours to be arranged.
Supervised reading on topics not covered by regular courses of instruction. Normally work is directed by a Biology faculty member of the Department of Molecular and Cellular Biology or Organismic and Evolutionary Biology. For biology concentrators, work may be supervised by faculty in other departments or outside Cambridge, provided it is co-sponsored by a Biology faculty member. Nonconcentrators must conduct their reading project with a Biology faculty member. Students must submit a registration request to the Biology Undergraduate Office before course enrollment. A written report of approximately 10 pages describing the material covered during the term is due in duplicate in the Undergraduate Office on the date indicated on the registration form.
Note: May be taken only once for concentration credit. May not be taken with the director/sponsor of an independent research course. Registration forms will not be accepted after Study Cards are due.
Prerequisite: Four semesters of introductory biology.

*Biology 95hfa. Genetic Screens: Dissecting Complex Biological Systems
Catalog Number: 2052
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfb. The Science of Exercise: Cardiopulmonary and Muscular Adaptations
Catalog Number: 2607
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfc. Cell Signaling In the Immune System. Apoptosis and Immune Diseases as Targets for Biotechnology Research
Catalog Number: 2935
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfd. Gene Therapy
Catalog Number: 3437
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfe. Molecular Virology: Playing by a Different Set of Rules?
Catalog Number: 4021
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hff. Oncogenes and Molecular Initiation of Cancer
Catalog Number: 5745
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfg. Conservation Biology
Catalog Number: 4576
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfh. Disordered Immunity: The Basis for Autoimmunity
Catalog Number: 4969
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfi. Sensing Sound: The Psychophysics and Neurophysiology of Hearing
Catalog Number: 7067
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfj. Vertebrate Brain Development: Mechanisms and Diseases
Catalog Number: 6361
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfk. From Disease Genes to Crop Improvment: Molecular Analysis of Polygenic Traits
Catalog Number: 7431
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfl. Problem Solving in Neuroscience: An On-line E-case Environment and Student-Centered Interactive Learning
Catalog Number: 7072
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfm. Evolutionary Perspectives on Infectious Disease
Catalog Number: 7766
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfn. Biology and Evolution of the Dinosauria
Catalog Number: 8189
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfo. Games Parasites Play: The Cellular and Molecular Biology of Host/Parasite Interactions
Catalog Number: 8456
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfp. Sticky-Tricky-Hairy-Scary Plants: Ecology and Evolution of Plant-Animal Interactions
Catalog Number: 8846
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfq. Molecular Mechanisms of Cancer
Catalog Number: 8555
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfs. Signaling Mechanisms in Neuronal Cell Biology
Catalog Number: 9282
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hft. The Guinness Book of Plants: Extreme Plant Physiology
Catalog Number: 5169
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfv. Biodiversity: Science, Policy, and Law
Catalog Number: 8959
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfw. Ecology, Dectection, and Treatment: Disease in the Developing World
Catalog Number: 1197
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfx. Muscles, Metabolism, and Movement
Catalog Number: 8716
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 98r. Introduction to Research
Catalog Number: 8616
William M. Gelbart and members of the Department
Half course (fall term; repeated spring term). Hours to be arranged.
Introduction to independent research. Recommended for students (ordinarily in the junior year) with no previous independent laboratory or library research experience. For Biology concentrators, ordinarily work is directed by a member of the Biological Sciences departments (MCB: Molecular and Cellular Biology, and OEB: Organismic and Evolutionary Biology) or affiliates. Other research sponsors must be approved by petition to the Biology Undergraduate Office. Non-concentrators must conduct their project with a faculty member in MCB or OEB. All students must submit registration materials for Biology 98r at the time of enrollment. A written report of approximately 5 pages is due in duplicate in the Biology Undergraduate Office on the date indicated on the registration form.
Note: To be graded SAT/UNS only. Laboratory safety session required.

*Biology 99ar. Research for Honors Thesis
Catalog Number: 8037
William M. Gelbart and members of the Department
Half course (fall term; repeated spring term). Hours to be arranged.
Ordinarily taken by students planning on writing an honors thesis. Recommended as the research course to be taken in the first semester of the student’s senior year. For Biology concentrators, work may be directed by a member of the Biological Sciences departments (MCB: Molecular and Cellular Biology, and OEB: Organismic and Evolutionary Biology) or affiliates. Other research sponsors must be approved by the Biology Undergraduate Committee and require an MCB or OEB co-sponsor. Non-concentrators must conduct their project with a faculty member in MCB or OEB. All students must submit registration materials for Biology 99ar at the time of enrollment. A written report of approximately 10 pages is due in duplicate in the Biology Undergraduate Office on the date indicated on the registration form. All students enrolling in this course must submit a short thesis proposal prior to the first day of classes of the fall semester. The thesis proposal form is available in the Biology Undergraduate Office.
Note: May be repeated only with special permission of the Biology Undergraduate Committee. Laboratory safety session required.

*Biology 99b. Honors Thesis
Catalog Number: 7264
William M. Gelbart and members of the Department
Half course (fall term; repeated spring term). Hours to be arranged.
Taken by students who are actively writing their honors thesis (ordinarily in the second semester of their senior year). For Biology concentrators, work may be directed by a member of the Biological Sciences departments (MCB: Molecular and Cellular Biology, and OEB: Organismic and Evolutionary Biology) or affiliates. Other research sponsors must be approved by the Biology Undergraduate Committee and require an MCB or OEB co-sponsor. Non-concentrators must conduct their project with a faculty member in MCB or OEB. All students must submit registration materials for Biology 99b at the time of enrollment. These registration materials are available in the Biology Undergraduate Office. Honors levels determined by the Biology Undergraduate Committee based upon readers’ evaluations.

For Undergraduates and Graduates

Biology S-105. Biodiversity of Tropical Plants (No Catalog Number — requires Harvard Summer School registration)
Catalog Number: 00-1
P. Barry Tomlinson (spring term)
Half course (summer term). Hours to be arranged.
An in-residence course centered in Miami, Florida, at Fairchild Tropical Garden. An intensive field and laboratory examination of tropical plant families with access to specialized tropical ecosystems available in South Florida. Attention given to economic plants of the tropics and specialized groups (e.g., palms and cycads) which are well represented in the area. Emphasis on growth and function as well as morphology, anatomy, and systematics.
Note: Students are admitted on the basis of previous experience and are examined on the basis of independent research projects.

[Biology 102. Evolution and Biology of the Seed]
Catalog Number: 7580
P. Barry Tomlinson
Half course (spring term). Hours to be arranged; plus additional laboratory work at the Arnold Arboretum and the Harvard Forest.
The evolution of the seed had major impact on the development of modern vegetation and hence of modern faunas, but can be traced back to the late Devonian. The evolutionary origins of the seed will be traced on the basis of paleobotanical evidence and comparative morphology of modern vascular plants. The biological conflict between the functions of the ovule (the seed precursor) at the time of pollination and the need for protection of the embryo will be discussed, together with the biological implications of seed dormancy.
Note: Expected to be given in 2000–01.
Prerequisite: Biological Sciences 2 recommended.

Biology 103. Plant Systematics
Catalog Number: 8704
Michael J. Donoghue
Half course (spring term). M., W., F., at 11. EXAM GROUP: 4
An introduction to the phylogeny, evolution, and diversification of the flowering plants, with emphasis on the major plant families of the world. Native and cultivated plants of eastern North America are studied in laboratory and field work.
Prerequisite: Biological Sciences 2 or permission of instructor.

Biology 104. Plants and Human Affairs
Catalog Number: 5281
Otto T. Solbrig
Half course (spring term). Lectures, M., W., F., at 10, and five three-hour laboratories. EXAM GROUP: 3
An introduction to the biology and use of plants of economic importance, including the role that plants have played in the development of civilization and in modern society. Topics to be covered are the structure, classification, economic importance, chemistry, and evolution and culture of the principal crops of the world, and cultural methods, including the use of biotechnology in plant breeding.
Prerequisite: Introductory Biology (Biological Sciences 2) or equivalent.

Biology 107. Evolution of Plant Life in Geologic Time
Catalog Number: 1318
Andrew H. Knoll
Half course (fall term). Lectures, M., W., F., at 9, and three hours of laboratory each week. EXAM GROUP: 2
Origin, evolution, dispersal, paleoecology, and geologic history of the major groups of the plant kingdom. Laboratory study of representative groups, living and fossil.
Prerequisite: Biological Sciences 2 or permission of instructor. Biology 20 recommended.

[Biology 113. Nonvascular Autotrophs and Fungi: A Survey]
Catalog Number: 3672
Donald H. Pfister
Half course (fall term). Hours to be arranged.
The biology and classification of algae, bryophytes, lichens, and fungi are considered. The goal is to introduce students to the diversity of the groups and the ecology, growth, and development of these organisms. Begins by reviewing previous classification schemes used to cover these groups. Life cycles, anatomy-morphology, physiology, and ecology of each group are covered through lectures, laboratory, and field trips. Each student is expected to undertake an independent project.
Note: Expected to be given in 2000–01.
Prerequisite: Biological Sciences 1 and 2 or equivalent.

Biology 114 (formerly Biology 213). Vertebrate Viviparity
Catalog Number: 4953
David A. Haig
Half course (spring term). Tu., Th., at 1. EXAM GROUP: 15
Viviparity has evolved many times in vertebrate phylogeny. The course will review the diversity of parental care in vertebrates and explore the selective forces that have favored the evolution of live-bearing. The evidence for intergenerational conflicts will be considered.

Biology 118. Biological Oceanography
Catalog Number: 7752
James J. McCarthy
Half course (spring term). Tu., Th., 10–11:30, and a two-hour discussion session each week. EXAM GROUP: 12, 13
The ocean as an ecological system. Emphasis on the ecology and physiology of the plankton. Considerable attention to processes and events that demonstrate the complexity of environmental-organismal interactions. Discussion sessions treat special topics such as sampling strategies, man’s impact on the marine environment, and critical analyses of current literature.
Note: For Biology and other natural science concentrators.
Prerequisite: Biological Sciences 1 and 2 and Chemistry 5 and 7 or Chemistry 10. Biology 19 is recommended.

Biology 120. Physiology of Plants
Catalog Number: 2554
N. Michele Holbrook
Half course (fall term). M., W., F., at 10. EXAM GROUP: 3
Introduction to the physiology, biochemistry, and development of plants. Topics include photosynthesis, energy balance, transport processes, growth, biomechanics, and reproduction. Emphasis on the physiological basis for structural adaptations of plants in relation to environmental constraints and on mechanisms leading to developmental and physiological integration at the whole-plant level. Laboratory sessions provide an introduction to basic measurement techniques in plant physiology.
Prerequisite: Biological Sciences 1 or permission of the instructor.

Biology 121a. Advanced Structure and Physiology of the Vertebrates
Catalog Number: 4049
Andrew A. Biewener, A. W. Crompton, and others
Half course (spring term). Hours to be arranged.
The purpose of this course is to introduce students to experimental techniques used in investigating the structure and physiology of vertebrates. Each instructor offers a list of research projects to be undertaken in his or her laboratory, and is prepared to supervise 5 students per instructor. The only formal class meetings are at the beginning (at which time the students outline their projects) and at the end when the project results are presented. An extensive commitment of time in the laboratory is required. The grade is based on the work completed, the oral presentation, and a short paper prepared as if it were to be submitted for publication in a professional journal.
Note: Laboratory safety session required.
Prerequisite: Biology 21 preferred, or permission of instructor.

*Biology 121b. Advanced Structure and Physiology of the Vertebrates
Catalog Number: 4670
Andrew A. Biewener
Half course (fall term). Hours to be arranged.
Students may extend the initial project undertaken in Biology 121a into a thesis research project.
Note: Laboratory safety session required.
Prerequisite: Biology 121a.

[Biology 123. Biology of Symbiosis]
Catalog Number: 0508
Colleen M. Cavanaugh
Half course (spring term). Hours to be arranged.
An examination of the major aspects of endosymbiosis with emphasis on mutualism, although some parasitic interactions are covered. Topics include origins of the eukaryotic cell, specificity and recognition of partners, distribution and diversity of associations, and coevolution of host and symbiont.
Note: Expected to be given in 2000–01.
Prerequisite: Biological Sciences 1 and 2, and Biological Sciences 11 or equivalent, or permission of instructor.

Biology 124. Genetics and the Origin of Species
Catalog Number: 7568
Kerry L. Shaw
Half course (fall term). Tu., Th., 2:30–4. EXAM GROUP: 16, 17
Topics including the genetic nature of species and the genetic mechanisms of speciation will be covered. Lectures on the genetic and phenotypic principles of evolution, the genetic structure of populations and the genetics of functional systems involved in the origins of biodiversity will provide the conceptual tools for discussions of the classic and modern primary literature. Course participation consists of half lecture, half discussion of the primary literature (both classic and modern).
Prerequisite: Biological Sciences 1; Biology 17, Biology 20 or Biology 22 or permission of the instructor. Some background in genetics will be helpful.

*Biology 130. Patterns and Processes in Fish Diversity
Catalog Number: 4624 Enrollment: Limited to 10
Karel F. Liem
Half course (spring term). M., W., F., at 1. EXAM GROUP: 6
As the largest and most diverse group of vertebrates, fishes inhabit virtually all aquatic environments including deep seas, intertidal zones, coral reefs, polar waters, the vast Amazonian basin, and great East Africa lakes. Some even exploit anoxic systems and temporary water bodies uninhabitable by other vertebrates. Moreover, a single fish species may occupy diverse environments through extraordinary long distance horizontal and vertical migratory strategies. To explore this unparalleled diversity, the course emphasizes bridging traditional academic boundaries with integrative analyses of the biology underlying rapid evolutionary radiations, stasis, and mass extinctions.
Prerequisite: Biological Sciences 1 and 2, or permission of instructor.

[Biology 133. Biology of Mollusks]
Catalog Number: 0751
Kenneth J. Boss
Half course (spring term). Hours to be arranged. Field trips and laboratories to be arranged.
Predominantly an organismic approach to the biology of the Phylum Mollusca; surveys the diversity of marine, terrestrial, and fresh-water mollusks, including special aspects of current research in systematics, ecology, zoogeography, and paleontology.
Note: Expected to be given in 2000–01. Biological Sciences 2 recommended, or permission of instructor required.

Biology 134. Behavioral Genetics
Catalog Number: 5230
Kerry L. Shaw
Half course (spring term). M., 3–5. EXAM GROUP: 8, 9
A seminar that focuses on the primary literature in the field of behavioral genetics, addressing our current understanding of behavior at the molecular, organismal, population and phylogenetic levels. Where possible, we will examine case studies of research systems for which we have insights at each level of analysis.
Prerequisite: Biological Sciences 1 and 2; BS 10 or some background in molecular biology desirable. Biology 20 and 22 also helpful.

[Biology 135. The Biology, Biochemistry, and Physiology of the Skeletal System of Vertebrates]
Catalog Number: 1278 Enrollment: Limited to 30
Melvin J. Glimcher (Medical School)
Half course (spring term). Tu., Th., 8:30–10. EXAM GROUP: 10, 11
Topics include development of bone and cartilage as organs and as tissues; mechanisms for remodeling (including fracture repair and tissue and organ grafting); biochemical composition, structure, and ultrastructural organization of the inorganic mineral phase and its molecular and structural relationships to the constituents of the organic matrix; the primary, secondary, and higher-ordered structures of collagen and noncollagenous proteins of bone; general principles underlying the molecular mechinisms and calcification of bone and other mineralized biological tissue; the structure, chemical composition, and function of the articular cartilage; and the biomechanics and biochemistry of joint function: and the pathophysiology of bone and cartilage to illuminate the basic biology, chemistry, and physiology of these tissues and organs. Selected related topics in molecular biology, and the molecular basis of genetic abnormalities of the skeleton.
Note: Expected to be given in 2000–01.
Prerequisite: Biological Sciences 2 and inorganic and organic chemistry, or permission of instructor.

Biology 139. Evolution of the Vertebrates
Catalog Number: 8562
A. W. Crompton and Farish A. Jenkins, Jr.
Half course (spring term). Tu., Th., at 8. EXAM GROUP: 10
Concentrates on the major steps in vertebrate evolution and does not attempt a synoptic review of fossil vertebrates. Emphasis on the anatomical and physiological changes that occurred in the transitional stages leading to fish, amphibians, reptiles, birds, and mammals as well as the geographic locations and environment in which these groups originated.
Prerequisite: Biological Sciences 2, Biology 21, or equivalent.

[Biology 143. Biology of the Fungi]
Catalog Number: 1308
Donald H. Pfister
Half course (fall term). Hours to be arranged.
Introduction to the morphology, classification, evolution, and diversity of the fungi, including both parasitic and saprophytic members. Attention to biological phenomena unique to fungi and their relationships to systems of classification. Students learn how to collect, isolate, and identify a variety of fungi.
Note: Expected to be given in 2000–01. At least one weekend field trip to be arranged.
Prerequisite: Science B-15, Biological Sciences 1 or 2, or equivalent.

[*Biology 147. Biogeography]
Catalog Number: 7330
Peter S. Ashton
Half course (spring term). Hours to be arranged.
Introduction to the geography of plants and animals. Distribution of species and genera, and development of floras and faunas interpreted in terms of population and species ecology, and the processes of migration, adaptation, and speciation.
Note: Expected to be given in 2001–02. Some familiarity with evolutionary theory and the major groups of plants and animals desirable.

[Biology 149. Plant Ecology]
Catalog Number: 2670
Fakhri A. Bazzaz
Half course (fall term). Hours to be arranged; plus a two-hour discussion or laboratory session each week and occasional field trips.
Introduction to concepts and principles of plant ecology, with emphasis on experimental approaches to understanding the biology of plant populations. Special attention is given to plant ecophysiological response to the physical and biological environment including energy, water, nutrients, environmental contaminants, pollinators, herbivores, predators and pathogens; competition and resource sharing; the niche and the level of ecological variation; community organization.
Note: Expected to be given in 2000–01.
Prerequisite: Biology 17, 19, or permission of instructor.

Biology 152. Population Genetics
Catalog Number: 0903
John R. Wakeley
Half course (fall term). Tu., Th., 10–11:30. EXAM GROUP: 12, 13
Introduction to population genetic theory. Covers deterministic and stochastic theory of gene frequencies, and coalescent theory of sample-based statistics. Emphasis on patterns of observed genetic variation within and between populations, and how these can serve as the basis for inference about mutation, genetic drift, migration, natural selection, and population structure.
Prerequisite: Biology 17, calculus, and knowledge of statistics and probability.

Biology 154. Theoretical Population and Community Ecology
Catalog Number: 0244
William H. Bossert
Half course (spring term). Hours to be arranged. Weekly discussion group to be arranged.
Theoretical treatment of population dynamics, evolutionary ecology, and community organization. Includes discussion of population growth, life history adaptations, competition theory, foraging strategies, coevolution, community models, diversity and stability of communities, comparison of ecosystems (ordination).
Prerequisite: Biology 17, 19, or equivalent; Mathematics 1a and 1b.

[Biology 160. Forest Ecology]
Catalog Number: 4369
David R. Foster
Half course (fall term). Hours to be arranged.
Plant community organization, structure, and dynamics focusing primarily on the forest vegetation of northeastern North America. Field studies emphasize the role of natural and anthropogenic disturbance in controlling the pattern of vegetation, methods of vegetation analysis, and historical methods of vegetation reconstruction.
Note: Expected to be given in 2000–01. Includes three full weekend field trips to the Harvard Forest.
Prerequisite: Biological Sciences 2 or permission of instructor.

[Biology 168. Life Cycles of Plants and Fungi]
Catalog Number: 2750
Half course (fall term). Hours to be arranged.
A study of the diversity and evolution of plant, fungal, and algal life cycles, with an emphasis on interactions between the generations. Topics will include embryology, sexual differentiation, mating types, parent-offspring and intragenomic conflict.
Note: Expected to be given in 2000–01.

Biology 174. Topics in Behavioral Biology: The Evolution of Cooperation
Catalog Number: 5199 Enrollment: Limited to 20
Naomi E. Pierce, David A. Haig, Marc D. Hauser, Kerry L. Shaw, and Richard W. Wrangham
Half course (spring term). M., 3–6. EXAM GROUP: 8, 9
Major issues in behavioral biology are examined in this course, with topics to change each year. The evolution of cooperation will be the focus of the 2000 spring term. The evolution of both inter- and intra-specific cooperation will be considered, including topics such as the evolution of complex social behavior, intra-cellular symbiosis, mutualism, reciprocal altruism, Prisoner’s Dilemma, and conflict resolution. The course will involve invited speakers and participation of professors across disciplines.
Prerequisite: Biology 22, Anthropolgy 170, Biological Sciences 25 or Science B-29 or permission of instructor.

Primarily for Graduates

Biology 200r. A.B./A.M. Laboratory Research
Catalog Number: 3696
William M. Gelbart and members of the Department
Half course (fall term; repeated spring term). Hours to be arranged.
Open only to Advanced Standing Biology concentrators doing research as part of a combined A.B./A.M degree. To be taken in both the first and second semesters of the student’s fourth year. Work may be directed by a member of the Biological Sciences departments (MCB - Molecular and Cellular Biology, and OEB - Organismic and Evolutionary Biology) or affiliates. Other research sponsors must be approved by the Biology Undergraduate Committee and require an MCB or OEB co-sponsor. All students must submit registration materials for Biology 200r at the time of enrollment. In the first semester, a written report of approximately 10 pages is due in duplicate in the Biology Undergraduate Office on the date indicated on the registration form. In the second semester, the honors thesis will be submitted on the date indicated on the registration form. All students enrolling in this course must submit a short thesis proposal prior to the first day of classes. The thesis proposal form is available in the Biology Undergraduate Office.
Note: Laboratory safety session required.

Biology 208. Issues in Paleobiology: Seminar
Catalog Number: 1344
Andrew H. Knoll, Stephen J. Gould, and Charles R. Marshall
Half course (spring term). Hours to be arranged.
A seminar focusing on current issues in paleobiology. Examines patterns of diversification, morphological change, and extinction, as well as detailed patterns of change across selected stratigraphic boundaries.

*Biology 209. Phylogenetic Analysis: Seminar
Catalog Number: 2534
David A. Baum
Half course (spring term). W., 1–3, Tu., 2–4. EXAM GROUP: 6, 7, 16, 17
Methods for constructing and interpreting phylogenies. What phylogenies are, and how they may be inferred from molecular or morphological data. Hands-on experience of relevant software and discussion of the underlying assumptions of the main approaches. Methods for evaluating statistical robustness of phylogenetic hypotheses. Methods for utilizing phylogenies in evolutionary and comparative studies.

[Biology 211r. Form, Function, and Evolution]
Catalog Number: 2056
Karel F. Liem and guest lecturers
Half course (fall term). Hours to be arranged.
Intrinsic and extrinsic determinants of form, considered from the evidence of neontological studies of vertebrates. Emphasis on current research approaches and theories. Consists of a mixture of seminars by faculty, guest lecturers, and students. Topics vary from year to year and are determined by the interests of the students.
Note: Expected to be given in 2000–01.

[Biology 212r. Advanced Topics in Plant Physiology]
Catalog Number: 2176
N. Michele Holbrook
Half course (fall term). Hours to be arranged.
A critical discussion of current research in plant physiology including measurement techniques, modeling, and experimental approaches.
Note: Expected to be given in 2000–01.
Prerequisite: Biology 120 or permission of instructor.

[Biology 214. Macroevolution of Interactions]
Catalog Number: 7040
Brian D. Farrell and Naomi E. Pierce
Half course (spring term). Hours to be arranged.
Focuses on current issues pertaining to the macroevolution of resource use and interspecific interactions, with emphases on insects, plants, and fungi. The goal will be to achieve a synthetic view of the evolution of ecological specialization and the evolving trophic web. Topics will include the timing, liability, and consequences of evolutionary change in traits affecting interactions (including mutualisms and antagonisms), and their implications for community structure and diversity. Readings will be of paleontological, phylogenetic, and neontological studies from primary literature and recent reviews.
Note: Expected to be given in 2000–01.
Prerequisite: Biology 17, 20, 22 or equivalent, or permission of instructor.

Biology 216r (formerly Biology 216). Topics in Plant Evolution
Catalog Number: 3081
Michael J. Donoghue
Half course (fall term). Hours to be arranged.
Critical discussion of current research in plant evolution; topics to change from year to year.
Prerequisite: Background in plant systematics and evolutionary biology; permission of the instructor.

Biology 221. Bacterial Diversity
Catalog Number: 1234
Colleen M. Cavanaugh
Half course (spring term). Tu., 11:30–1. EXAM GROUP: 13, 14
The remarkable diversity of prokaryotes is examined. Physiological, genetic, ecological, and evolutionary characteristics of bacterial groups are discussed, as well as the relation of phenotype to phylogeny.
Prerequisite: Biological Sciences 1 and 2, and Biological Sciences 11 or equivalent, or permission of instructor.

*Biology 234. Topics in Marine Biology
Catalog Number: 4637 Enrollment: Limited.
Robert M. Woollacott
Half course (fall term). W., 2–5. EXAM GROUP: 7, 8, 9
Reproduction of marine invertebrates.
Note: Weekly class meeting and several field trips and laboratories through course of term. Independent projects required.

[Biology 236. Species and Speciation]
Catalog Number: 5400
Kerry L. Shaw
Half course (fall term). Hours to be arranged.
Views on the nature of species are examined. Genetic and phenotypic models and mechanisms of the speciation process are examined and evaluated by case studies from a selection of the major groups of organisms.
Note: Expected to be given in 2000–01.
Prerequisite: Biological Sciences 14 and Biology 17 or 20, or permission of the instructor.

Biology 239r. Topics in Molecular Ecology and Evolution
Catalog Number: 8124
Stephen R. Palumbi
Half course (spring term). W., 3–5. EXAM GROUP: 17, 18
Focuses on the use of molecular genetic tools to illuminate current issues in ecology, population biology, and evolution. We will use the primary literature to explore a particular topic chosen by the course participants. Topics to choose among will include mating systems and mate choice, population structure, genetic signatures of demographic history etc.

Biology 245. Topics in Plants and Environments
Catalog Number: 1739
Fakhri A. Bazzaz
Half course (fall term; repeated spring term). Tu., 7–10 p.m. EXAM GROUP: 18
A discussion of recent advancements in research on plants in their environments. Presentations of research proposals. Discussions on design and execution of field and laboratory experiments and readings and discussions of important recent papers.
Prerequisite: Biology 149, equivalent, or permission of instructor.

[*Biology 250. Tropical Ecology]
Catalog Number: 5931
Peter S. Ashton
Half course (fall term). Hours to be arranged.
An introduction to the ecology of tropical communities with particular emphasis on adaptations of terrestrial plants. Fall lectures provide background for an optional research field trip to northern South America in late December or January. Students not participating in the field trip are asked to write a paper on a subject of their choice.
Note: Expected to be given in 2000–01.

*Biology 251. Introduction to Vertebrate Surgery
Catalog Number: 2075 Enrollment: Limited to 16
Arthur L. Lage (Medical School)
Half course (fall term). W., 1–5. EXAM GROUP: 6, 7, 8, 9
Teaches students the basic principles of aseptic surgery with emphasis on practicality. Students learn basic “open” surgery as well as newer high tech videoscopic minimally invasive technique, obtaining hands-on experience in scrubbing, gowning, and sterile technique while serving as anesthetist and surgeon. Course covers surgical applications for a wide variety of species including biped and quadruped mammals, ruminant and non-ruminant mammals, birds, amphibians, and reptiles; students are expected to integrate their previously learned knowledge of anatomy and physiology into the context of the whole, living animal.
Note: Intended for the student interested in the application of surgical technique in higher studies in biology and related disciplines.
Prerequisite: Comparative Anatomy, Biology 21, or equivalent course.

[Biology 252. Coalescent Theory]
Catalog Number: 0118
John R. Wakeley
Half course (fall term). Tu., Th., 10–11:30. EXAM GROUP: 12, 13
The mathematics and computation of ancestral inference in population genetics. Theory relates observable genetic data to factors of evolution such as mutation, genetic drift, migration, natural selection, and population structure.
Note: Expected to be given in 2000–01.
Prerequisite: Biology 152 or consent of instructor: calculus and statistics or probability.

Biology 253r. Seminar in Evolutionary Genetics
Catalog Number: 8104
John R. Wakeley
Half course (fall term; repeated spring term). Tu., at 4. EXAM GROUP: 18
Readings and discussion of primary literature in population and evolutionary genetics.

[Biology 254. Genomic Imprinting and Parent-Offspring Conflict]
Catalog Number: 9341
David A. Haig
Half course (fall term). Tu., at 2:30. EXAM GROUP: 16, 17
A seminar course that will discuss evolutionary aspects of genomic imprinting and the related theory of parent-offspring conflict.
Note: Expected to be given in 2001–02.

[Biology 255. Nature and Regulation of Marine Ecosystems ]
Catalog Number: 7753
James J. McCarthy and guest lecturers
Half course (fall term). Hours to be arranged.
A presentation of topics that are of current interest in marine ecosystems. Emphasis on identification and quantification of biological and environmental factors important in the regulation of community structure in the intertidal, deep benthic, and planktonic realms.
Note: Expected to be given in 2000–01.
Prerequisite: Biology 19 and 118.

[Biology 257r. Topics in Microbial Ecology]
Catalog Number: 0621
Ralph Mitchell
Half course (fall term). Hours to be arranged.
A critical discussion of current research in microbial ecology.
Note: Expected to be given in 2000–01. A term paper and a seminar are required.

[Biology 267. Topics in Symbiosis: Seminar]
Catalog Number: 3859
Colleen M. Cavanaugh
Half course (fall term). Hours to be arranged.
Focuses on current issues in symbiosis. Emphasis on prokaryote-eukaryote associations and comparative approaches to both mutualistic and parasitic interactions. Critical review and discussion of one or a few topics of interest to participants.
Note: Expected to be given in 2000–01.

*Biology 299r. Forest Practice and Research
Catalog Number: 6128
David R. Foster 7770
Half course (fall term; repeated spring term). Tu., Th., at 10. EXAM GROUP: 12
Field and laboratory research into the history, biology, ecology, culture, and economic problems of local, regional, and world forests. Individual research projects.
Note: Open to students who satisfy the staff that they have had adequate training. Seminars, conferences, field and laboratory work at the Harvard Forest, Petersham, Massachusetts.

Graduate Courses of Reading and Research

*Biology 303. Theoretical Population Genetics
Catalog Number: 4248
John R. Wakeley 5680

*Biology 304. Mycology
Catalog Number: 4702
Donald H. Pfister 4344 (on leave fall term)

*Biology 305. The Fundamental Interconnectedness of All Things
Catalog Number: 3647
David A. Haig 1629

*Biology 307. Biomechanics, Physiology and Musculoskeletal Biology
Catalog Number: 2831
Andrew A. Biewener 1446

*Biology 309. Plant Anatomy and Morphology
Catalog Number: 4772
P. Barry Tomlinson 3712

*Biology 311. Behavioral Evolution and Sexual Selection
Catalog Number: 2061
Kerry L. Shaw 1852

*Biology 316. Plant Population Biology
Catalog Number: 3863
Otto T. Solbrig 3197

*Biology 320. Biomechanics and Evolution of Vertebrates
Catalog Number: 8915
George Varick Lauder 2375

*Biology 321. Paleontology and Macroevolutionary Theory
Catalog Number: 5799
Stephen J. Gould 1707

*Biology 322. Functional Anatomy of the Vertebrates
Catalog Number: 6365
A. W. Crompton 1154 (on leave fall term)

*Biology 323. Advanced Vertebrate Anatomy
Catalog Number: 8188
Farish A. Jenkins, Jr. 3558

*Biology 324. Molecular Evolution
Catalog Number: 2356
Daniel L. Hartl 3278

*Biology 325. Marine Biology
Catalog Number: 4643
Robert M. Woollacott 4135 (on leave spring term)

*Biology 334. Behavioral Ecology
Catalog Number: 8279
Naomi E. Pierce 2889 (on leave fall term)

*Biology 335. Ichthyology and Functional Anatomy of Fishes
Catalog Number: 4640
Karel F. Liem 3843

*Biology 337. Plant Phylogeny
Catalog Number: 0762
Michael J. Donoghue 3277

*Biology 339. Whole-Plant Physiology
Catalog Number: 5214
N. Michele Holbrook 1220

*Biology 341. Coevolution
Catalog Number: 2998
Brian D. Farrell 1985

*Biology 343. Microbial Ecology and Symbiosis
Catalog Number: 1288
Colleen M. Cavanaugh 2538

*Biology 345. Biological Oceanography
Catalog Number: 4676
James J. McCarthy 4343

*Biology 348. Plant Ecology
Catalog Number: 2885
Fakhri A. Bazzaz 7926

*Biology 355. Evolutionary Developmental Biology
Catalog Number: 9192
James Hanken 2719 (fall term only)

*Biology 357. Population Biology and Mathematical Biology
Catalog Number: 5392
William H. Bossert 1049

*Biology 359. Paleobotany
Catalog Number: 0248
Andrew H. Knoll 7425

*Biology 361. Molecular Systematics and Development of Plants
Catalog Number: 1985
David A. Baum 1218

*Biology 387. Tropical Botany
Catalog Number: 6715
Peter S. Ashton 6086

*Biology 389. Population Biology and Evolution
Catalog Number: 0680
Stephen R. Palumbi 2406

*Biology 399 (formerly Biology 220). Topics in Organismic and Evolutionary Biology
Catalog Number: 0764
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Half course (spring term). .
Presents the research interests and experiences of members of the Department on integrative topics in the field. The specific topic and list of faculty participants varies from year to year.
Note: Required of all first-year graduate students in Organismic and Evolutionary Biology.