Biological Sciences

Faculty of the Department of Molecular and Cellular Biology

Douglas A. Melton, Thomas Dudley Cabot Professor of the Natural Sciences (Chair of MCB)
Howard C. Berg, Professor of Molecular and Cellular Biology and Professor of Physics (on leave 2000-01)
Daniel Branton, Higgins Research Professor of Biology
John S. Chant, Associate Professor of Biochemistry and Molecular Biology
James E. Davis, Senior Lecturer on Chemistry and Chemical Biology and Molecular and Cellular Biology
John E. Dowling, Harvard College Professor and Llura Gund Professor of Neurosciences
Catherine Dulac, 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 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 2001-2002)
Guido Guidotti, Higgins Professor of Biochemistry (on leave spring term)
Stephen C. Harrison, Higgins Professor of Biochemistry and Molecular Biology (Acting Head Tutor, Biochemical Sciences)
J. Woodland Hastings, Paul C. Mangelsdorf Professor of Natural Sciences
Craig P. Hunter, Assistant Professor of Molecular and Cellular Biology, Associate Professor of Molecular and Cellular Biology
Nancy Kleckner, Herchel Smith Professor of Molecular Biology
Jeremy R. Knowles, Amory Houghton Professor of Chemistry and Biochemistry and Dean of the Faculty of Arts and Sciences
Samuel M. Kunes, Professor of Molecular and Cellular Biology (on leave fall term)
Richard M. Losick, Harvard College Professor and Maria Moors Cabot Professor of Biology
Robert A. Lue, Senior Lecturer on Molecular and Cellular Biology, Director of Undergraduate Studies in the Biological Sciences
Tom Maniatis, Thomas H. Lee Professor of Molecular and Cellular Biology
Andrew P. McMahon, Frank B. Baird, Jr. Professor of Science (on leave fall term)
Markus Meister, Jeff C. Tarr Professor of Molecular and Cellular Biology
Matthew Meselson, Thomas Dudley Cabot Professor of the Natural Sciences
Andrew W. Murray, Professor of Molecular and Cellular Biology
Venkatesh N. Murthy, Assistant Professor of Molecular and Cellular Biology, Tutor in Biochemical Sciences
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 (on leave spring term)
Don C. Wiley, John L. Loeb Professor of Biochemistry and Biophysics, Tutor in Biochemical Sciences (on leave 2000-01)

Associate Members of the Department of Molecular and Cellular Biology

Elena M. Kramer, Assistant Professor of Biology
Hidde Ploegh, Mallinckrodt Professor of Immunopathology and Professor of Pathology (Medical School)
Stuart L. Schreiber, Morris Loeb Professor of Chemistry and Chemical Biology
Gregory L. Verdine, Professor of Chemistry, Associate of Pforzheimer House

Faculty of the Department of Organismic and Evolutionary Biology

Daniel L. Hartl, Higgins Professor of Biology (Chair of OEB)
David A. Baum, Associate Professor of Organismic and Evolutionary Biology
Fakhri A. Bazzaz, Mallinckrodt Professor of Biology
Andrew A. Biewener, Charles P. Lyman Professor of Biology, Charles P Lyman Professor of Biology
Kenneth J. Boss, Professor of Biology
Colleen M. Cavanaugh, 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
Gonzalo Giribet, Assistant Professor of Biology and Assistant Curator of Invertebrates
David A. Haig, Associate Professor of Biology
James Hanken, Alexander Agassiz Professor of Zoology in the Museum of Comparative Zoology (on leave fall term)
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, Fisher Professor of Natural History
Elena M. Kramer, Assistant Professor of Biology
George V. Lauder, Professor of Biology and Alexander Agassiz Professor of Zoology in the Museum of Comparative Zoology
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, Master of Pforzheimer House, Director of the Museum of Comparative Zoology (on leave fall term)
Stephen R. Palumbi, Professor of Biology (on leave fall term)
Donald H. Pfister, Asa Gray Professor of Systematic Botany
Naomi E. Pierce, Sidney A. and John H. Hessel Professor of Biology
Otto T. Solbrig, Bussey Professor of Biology, Acting Director of the David Rockefeller Center for Latin American Studies
P. Barry Tomlinson, Edward C. Jeffrey Professor of Biology, Emeritus
John R. Wakeley, Assistant Professor of Biology
Robert M. Woollacott, Professor of Biology

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, Professor of Physics (on leave spring term)
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
Hidde Ploegh, Mallinckrodt 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 Acting 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 50. Genetics and Genomics
Catalog Number: 9370
Daniel L. Hartl
Half course (spring term). M., W., F., at 12 and three hours of laboratory/discussion each week. EXAM GROUP: 5
For 2000-2001 academic year, this course is only offered during the spring term. Analysis of genes and genomes with emphasis on function, transmission, mutation, and evolution, with examples from animals, plants, bacteria, and fungi. Discusses classical and current methods of gene and genome analysis, including genetic, molecular, quantitative, and bioinformatic approaches. For current Biology and Biochemical Sciences concentrators, this course may be taken in lieu of Biological Sciences 1. Please refer to the respective concentration notes for additional information on the new course sequence.
Note: Lectures and weekly laboratory/discussion section. For 2000-2001 academic year, this course is only offered during the spring term.

Biological Sciences 51 (formerly Biological Sciences 2). Integrative Biology of Organisms
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 integrative and functional approach to plant and animal biology in an evolutionary context, emphasizing common attributes of whole organisms and their solutions to problems imposed by the physical environment. Topics to be covered include development and organization of body plans, gas exchange, transport and excretion, information processing, support and locomotion, and the acquisition of energy sources.
Note: Knowledge of introductory molecular, cellular biology, and genetics is recommended.

Biological Sciences 52 (formerly 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
An integrated introduction to the basic principles of molecular 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. For current Biology and Biochemical Sciences concentrators, this course may be taken in lieu of Biological Sciences 10. Please refer to the respective concentration notes for additional information on the new course sequence.
Note: Chemistry 17 may be taken concurrently.
Prerequisite: Chemistry 17 or 20.

Biological Sciences 53. Diversity and History of Life
Catalog Number: 3342
David A. Baum, David A. Haig, and John R. Wakeley
Half course (spring term). Tu., Th., 10–11:30 and a weekly three-hour lab. EXAM GROUP: 12, 13
A study of the process of organic evolution and its result: the structural, functional, and genetic diversity of organisms. Emphasis on recent advances in understanding phylogenetic relationships among the primary groups of organisms, major events in the history of life, and the fundamental concepts and methods of evolutionary biology.
Note: Replaces Biology 17 and 20.
Prerequisite: BS 1 or 50, BS 2 or 51, or permission of instructor.

Biological Sciences 54. Introductory Cell Biology
Catalog Number: 0801
Robert A. Lue, Raymond L. Erikson, and Douglas A. Melton
Half course (spring term). 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 integrated introduction to the structure, function, and interactions of cells, with an emphasis on their molecular composition and dynamics. The course will emphasize experiments and hypotheses that have led to our current understanding of the cell. The discussion sections will focus on problem-solving and evaluation of data. The laboratory exercises will provide exposure to several techniques commonly used in cell biology. For current Biology and Biochemical Sciences concentrators, this course may be taken in lieu of Biological Sciences 11. Please refer to the respective concentration notes for additional information on the new course sequence.
Prerequisite: Chemistry 17 or 20.

Biological Sciences 56. Structure, Function, and Physical Chemistry of Macromolecules
Catalog Number: 5424
Stephen C. Harrison and Don C. Wiley
Half course (spring term). M., W., F., at 9 and a weekly section to be arranged. EXAM GROUP: 2
An introduction to macromolecular structure that integrates the basic principles of equilibrium thermodynamics, chemical kinetics, and molecular dynamics. Particular attention is paid to the forces and energetics of single molecules and how they determine the properties of ensembles of these molecules. Specific examples of such phenomena will be drawn from biochemistry and cell biology.
Prerequisite: BS 10 or 52 and BS 11 or 54 are recommended but not required; Chemistry 10 or equivalent; Chemistry 27 or Chemistry 30; Physics 11 (may be taken concurrently); Math 21a; Math 21b is recommended but not required.

Biological Sciences 80 (formerly Biological Sciences 25). 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
Stephen C. Harrison 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 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 ordinarily be repeated no more than once.

*Biochemical Sciences 99. Laboratory Research for Honors Thesis
Catalog Number: 6670
Stephen C. Harrison 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. Students may enter the course at midyear only with the permission of the Head Tutor. The thesis proposal must be approved by the Head Tutor prior to enrolling in Biochemical Sciences 99.

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). Hours to be arranged.
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.
Note: Expected to be given in 2001–02.
Prerequisite: BS 10 or 52, BS 11 or 54 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). Hours to be arranged.
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.
Note: Expected to be given in 2001–02.
Prerequisite: BS 10 or 52, BS 11 or 54, and BS 14 or 50.

[MCB 114. Structure and Function of Membrane Proteins]
Catalog Number: 8244
Don C. Wiley
Half course (spring term). Hours to be arranged. EXAM GROUP: 18
An advanced course on the relationship between the atomic structure and the biological function of membrane proteins. Topics include: Both methods, such as: membrane protein crystallization; 2D electron crystallography; single particle image reconstruction; X-ray diffraction; and biological examples, such as: bacterial rhodopsin, photosynthetic and respiratory proteins, toxins, pores, and ion channels. Because relatively few membrane protein structures have been determined, a complete examination of current knowledge is possible. Students are required to evaluate and discuss scientific papers.
Note: Expected to be given in 2001–02.
Prerequisite: BS 10 or 52, BS 11 or 54 and MCB 61 or equivalent preparation in physical chemistry.

MCB 115. Cellular Basis of Neuronal Function
Catalog Number: 8703 Enrollment: Limited to 30.
Venkatesh N. Murthy
Half course (fall term). W., F., 10–11:30. EXAM GROUP: 3, 4
Processes involved in the function of neurons will be explored, with emphasis on biophysical and cell biological approaches. Topics include excitable membranes, intracellular membrane trafficking, cytoskeletal dynamics, synaptic transmission, dendritic integration, and synaptic plasticity.
Prerequisite: BS 25 or BS 80 or equivalent.

MCB 116. Experimental Embryology
Catalog Number: 1207 Enrollment: Limited to 24. 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: BS 10 or 52, or BS 11or 54, or MCB 16 or 118 or their equivalents.

MCB 117. Experimental Neuroscience
Catalog Number: 3175 Enrollment: Limited to 16.
Markus Meister, John E. Dowling, Catherine Dulac, Venkatesh N. Murthy, 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: BS 25 or 80.

[MCB 118 (formerly MCB 16). Developmental Biology]
Catalog Number: 0749
Andrew P. McMahon
Half course (fall term). Hours to be arranged.
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.
Note: Expected to be given in 2001–02.
Prerequisite: BS 1 or 50. BS 10 or 11 or 52 or 54 suggested but not required.

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 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: BS 10 or 52 or equivalent.

[MCB 129. Molecular Genetics of Neuronal Development]
Catalog Number: 8956 Enrollment: Limited to 25.
Samuel M. Kunes
Half course (fall term). Hours to be arranged.
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.
Note: Expected to be given in 2001–02.

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: BS 25 or 80 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: BS 10 or 52, and BS 25 or 80.

MCB 142 (formerly MCB 242). Chromosomes
Catalog Number: 7948 Enrollment: Limited to 25.
Matthew Meselson
Half course (spring term). W., 2–4. EXAM GROUP: 7, 8
Selected aspects of the structure, replication, segregation, recombination, and function of chromosomes. Current findings will be considered in a historical context. Lectures, student presentations and critical discussion of the scientific literature.

MCB 150. Developmental Genetics and Genomics
Catalog Number: 5703
Craig P. Hunter
Half course (spring term). M., W., 2–3:30. EXAM GROUP: 7, 8
An advanced course on the genetic control of plant and animal development. Topics include classical and molecular genetic analysis of developmental processes and mechanisms in nematodes, flies, fish, plants, mice, and man. The course will consist of lectures, student presentations, several written assignments, and an exam.
Prerequisite: BS 1or 50, BS 10 or 52, BS 11 or 54, their equivalents or permission of instructor.

*MCB 155. Molecular Mechanisms of Gene Control
Catalog Number: 6230
Tom Maniatis
Half course (fall term). Tu., Th., 4–5:30, plus two-hour section times to be arranged. EXAM GROUP: 18
An advanced course on the control of gene regulation. Topics include: mechanisms of gene regulation at the level of transcription, chromatin structure, DNA methylation, RNA processing, mRNA localization, and protein synthesis 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. Two exams.
Prerequisite: BS 10 or 52 and BS 11 or 54 (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: BS 1 or 50 and BS 10 or 52. Genetics and Cell Biology strongly recommended.

[MCB 176. Membrane Structure and Function]
Catalog Number: 3186
Guido Guidotti
Half course (fall term). M., W., 2–4. EXAM GROUP: 7, 8
The structure of membrane proteins and their involvement in transport and signal transduction: membrane structure; membrane protein synthesis and insertion in the bilayer; transporters and pumps; channels; electron transport and H+gradients; ATP synthase; 7 transmembrane segment receptors and G proteins; receptor tyrosine kineses and dimerization.
Note: Expected to be given in 2001–02.
Prerequisite: BS 10 or 52 and BS 11or equivalent.

[MCB 177. Macromolecular Assemblies in Genetic Processes]
Catalog Number: 3102 Enrollment: Limited to 30.
James C. Wang
Half course (spring term). Hours to be arranged.
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.
Note: Expected to be given in 2001–02.
Prerequisite: BS 10 or 52 and BS 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: BS 1 or 50 and BS 2 or 51; BS 25 or 80 desirable.

MCB 188. Genetics and Biochemistry of Chromosome Behavior
Catalog Number: 8561
Nancy Kleckner
Half course (spring term). Tu., Th., 2–4. EXAM GROUP: 16, 17
Chromosome morphogenesis in prokaryotic and eukaryotic organisms. Topics will include chromosome structure, interactions between chromosomes (sisters and homologs), DNA recombination and repair, topoisomerases, transposable elements and site-specific recombination, epigenetic inheritance. Genetic, cytological, and biochemical approaches will be integrated. Lecture, reading, and discussion of classical and current literature and consideration of future experimental directions.
Prerequisite: BS 10 or 52, BS 11 or 54, and BS 14 or 50.

*MCB 190. Bioinformatics: The Computational Analysis of Genes and Proteins
Catalog Number: 7805 Enrollment: Limited to 40.
Walter Gilbert and Christopher Peter Sears
Half course (spring term). Tu., Th., 2:30–4. EXAM GROUP: 16, 17
This course provides advanced students with an introduction to the theory and methods of computational molecular biology. The focus is on the use of bioinformatics to solve problems in biology. We will cover the practical application and theoretical basis of computational methods used to search for, classify, analyze and model DNA, RNA, and protein sequences. Topics covered include: accessing molecular databases, pattern database similarity searching, classification of sequence and structure, alignment of sequences, rapid similarity searching, phylogenies, pattern learning, and gene prediction algorithms. Emphasis is placed on practical methods, an understanding of the primary literature, and an assessment of analytical methods.
Note: The course includes a computer lab and an introduction to algorithm programming.
Prerequisite: BS10 and 11. Calculus 1 and Introductory Computer Science recommeded. Or permission of the Instructor.

Cross-listed Courses

Biophysics 101. Genomics and Computational Biology

Primarily for Graduates

MCB 200a. Introduction to Graduate Study in Molecular and Cellular Biology
Catalog Number: 7215 Enrollment: Enrollment is limited to MCB graduate students.
Members of the faculty
Half course (fall term). Tu., Th., 2–3:30. EXAM GROUP: 16, 17
The goal of the course is to introduce the methods and logic of modern biology as developed through reading and discussion of research papers in macromolecules, cell biology, and gene regulation. Students are expected to actively participate in critical evaluation and discussion. The course will be team-taught by faculty.

MCB 200b. Introduction to Graduate Study in Molecular and Cellular Biology
Catalog Number: 2655 Enrollment: Enrollment is limited to MCB graduate students.
William M. Gelbart and members of the faculty.
Half course (spring term). M., 10:30–12, W., 4–5:30. EXAM GROUP: 3, 4, 9
The goal of the course is to introduce the methods and logic of modern biology as developed through reading and discussion of research papers in genetics, developmental biology, and neuroscience. Students are expected to actively participate in critical evaluation and discussion. The course will be team taught by faculty.

[MCB 212. Topics in Biophysics]
Catalog Number: 3351
Howard C. Berg
Half course (spring term). Hours to be arranged.
Motility and sensory transduction; Chemotaxis in bacteria; flagellar motility; prokaryotic and eukaryotic motor molecules. A term paper and seminar will be required.
Note: Expected to be given in 2001–02.

*MCB 250. Producing a Phenotype: Modern Genetics & Genomics
Catalog Number: 7006 Enrollment: Limited to 20.
William M. Gelbart
Half course (spring term). Tu., Th., 1–2:30. EXAM GROUP: 15, 16
A course consisting of a combination of lectures, student seminars, and computational investigations exploring the current interface between the genetic and genomic analysis of DNA sequences and gene products. The goal is to understand how these different approaches may be synthesized to understand biological processes.
Prerequisite: Courses in basic genetics and molecular biology, and permission of instructor.

MCB 268. Molecular Immunology: Seminar
Catalog Number: 2196 Enrollment: Limited to 20. 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. Limited to 10.
J. Woodland Hastings
Half course (spring term). W., 3–5. EXAM GROUP: 8, 9
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.
Prerequisite: BS 1 or 50 and BS 11 or 54 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 (on leave spring term)

*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 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 Meselson 1319

*MCB 314. Molecular Genetics of Cell-Cell Interactions in Plants
Catalog Number: 5564
Robert E. Pruitt 3376 (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 2000-01)

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

*MCB 360. Proteins Involved in DNA Transactions
Catalog Number: 0407
James C. Wang 4870 (on leave spring term)

*MCB 364. Vertebrate Development
Catalog Number: 1396
Andrew P. McMahon 3312 (on leave fall term)

*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

*MCB 374. Developmental Neurobiology
Catalog Number: 6167
Samuel M. Kunes 3486 (on leave fall term)

*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 2001-2002)

*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 (on leave 2000-01)

*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 Meselson 1319

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

*MCB 391. Biochemistry
Catalog Number: 4888
Guido Guidotti 1203 (on leave spring term)

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

*MCB 396. Regulation of Mitosis
Catalog Number: 5706
Andrew W. Murray 3765

*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: BS 2 or 51 recommended, or permission of instructor required.

Biology 19. Population Biology: Ecology
Catalog Number: 3365
William H. Bossert
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.
Prerequisite: Mathematics 1 and Science B-15 or BS 2 or 51 or equivalent.

Biology 21. Structure and Physiology of the Vertebrates
Catalog Number: 0921
George V. Lauder and 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 vertebrate groups in relationship to environmental factors. 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: BS 1 or 50 and 2 or 51.

Biology 22. Animal Behavior
Catalog Number: 2539
Naomi E. Pierce
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 BS 1, 2, 50, 51, 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., 10–11:30, one afternoon laboratory per week, plus occasional field trips. EXAM GROUP: 12, 13
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 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. Conceptual Issues in Evolutionary Biology
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 and Human Performance
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 Biotechnological 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. Immune Invasion: What Can We Learn from Sneaky Pathogens
Catalog Number: 4021
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hff. Oncogenes and the 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 95hfh. Size: It Really Does Matter
Catalog Number: 4969
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfi. Neurodegenerative Diseases and Neuropsychiatric Disorders
Catalog Number: 7067
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfj. Plant Sexual Reproduction: Evolutionary and Molecular Aspects
Catalog Number: 6361
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfl. Problem Solving in Neuroscience: An Interactive Case-Based Online Network (ICON)
Catalog Number: 7072
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfm. Games Parasites Play: The Cellular and Molecular Biology of Host/Parasite Interactions
Catalog Number: 7766
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfn. The Genetics of Cancer in the Postgenomic Era
Catalog Number: 8189
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfo. The Guinness Book of Plants: Extreme Plant Physiology
Catalog Number: 8456
William M. Gelbart and members of the Department
Half course (throughout the year). Hours to be arranged.

*Biology 95hfp. Biodiversity: Science, Policy, and Law
Catalog Number: 8846
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 102. Evolution and Biology of the Seed
Catalog Number: 7580
P. Barry Tomlinson
Half course (spring term). First class meeting 2/1 at 12 pm Herbarium Seminar Room. 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.
Prerequisite: BS 2 or 51 recommended.

[Biology 104. Plants and Human Affairs]
Catalog Number: 5281
Otto T. Solbrig
Half course (spring term). Hours to be arranged.
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.
Note: Expected to be given in 2001–02.
Prerequisite: Introductory Biology (Biological Sciences 51) or equivalent.

Biology 105. Environmental Consequences of Land Use Changes in Latin America
Catalog Number: 1226
Otto T. Solbrig
Half course (spring term). Tu., Th., 10–11:30. EXAM GROUP: 12, 13
Identifies and introduces key issues in land use changes and land management practices in Latin America. Three categories of land use are discussed: as ecosystems, as producers of raw materials, and as space. Agriculture, pastoral production, forestry, energy extraction and production, water supply, and land use for urbanization are considered as main areas of conflict and as factors that have influenced resource use. Lectures introduce case studies of actual situations in the forest-cropland interface. Students are expected to write a paper on a topic relating to class material.

[Biology 107. Evolution of Plant Life in Geologic Time]
Catalog Number: 1318
Andrew H. Knoll
Half course (fall term). Hours to be arranged.
Origin, evolution, dispersal, paleoecology, and geologic history of the major groups of the plant kingdom. Laboratory study of representative groups, living and fossil.
Note: Expected to be given in 2001–02.
Prerequisite: BS 2 or 51 or permission of instructor.

[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 2001–02.
Prerequisite: BS 1 or 50 and 2 or 51 or equivalent.

Biology 114 (formerly Biology 213). Vertebrate Viviparity
Catalog Number: 4953
David A. Haig
Half course (fall 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). Hours to be arranged.
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, plankton demonstrations, and critical analyses of current literature.
Note: Expected to be given in 2001–02. For Biology and other natural science concentrators.
Prerequisite: BS 1 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). Hours to be arranged.
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.
Note: Expected to be given in 2001–02.
Prerequisite: Permission of the instructor.

Biology 121a. Advanced Structure and Physiology of the Vertebrates
Catalog Number: 4049
Andrew A. Biewener and George V. Lauder
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 and George V. Lauder
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 (fall term). Hours to be arranged. EXAM GROUP: 10, 11
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.
Prerequisite: BS 1 or 50 and 2 or 51, and BS 11 or equivalent, or permission of instructor.

*Biology 130. Patterns and Processes in Fish Diversity
Catalog Number: 4624 Enrollment: Limited to 10.
Karel F. Liem and George V. Lauder
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: BS 1or 50, and 2 or 51, 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 2001–02. BS 2 or 51 recommended, or permission of instructor required.

[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). Hours to be arranged.
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 2001–02.
Prerequisite: BS 2 or 51 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). Hours to be arranged.
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.
Note: Expected to be given in 2001–02.
Prerequisite: BS 2 or 51, Biology 21, or equivalent.

Biology 149. Plant Ecology
Catalog Number: 2670
Fakhri A. Bazzaz
Half course (fall term). Tu., Th., 11:30–1; plus a two-hour discussion or laboratory session each week and occasional field trips. EXAM GROUP: 13, 14
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.
Prerequisite: BS 53, Biology 17, 19 or permission of instructor.

[Biology 152. Population Genetics]
Catalog Number: 0903
John R. Wakeley
Half course (fall term). Hours to be arranged.
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.
Note: Expected to be given in 2001–02.
Prerequisite: BS 53, 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.
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).
Note: Expected to be given in 2001–02.
Prerequisite: Biology 19, BS 53 (formerly Biology 17), or equivalent; Mathematics 1a and 1b.

Biology 160. Forest Ecology
Catalog Number: 4369
David R. Foster
Half course (fall term). Tu., Th., at 10. EXAM GROUP: 12
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: Includes two full weekend field trips to the Harvard Forest.
Prerequisite: BS 2 or 51 or permission of instructor.

[Biology 168. Life Cycles of Plants and Fungi]
Catalog Number: 2750
David A. Haig
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 2001–02.

[Biology 174. Topics in Behavioral Biology]
Catalog Number: 5199 Enrollment: Limited to 20.
Naomi E. Pierce, David A. Haig, Marc D. Hauser, and Richard W. Wrangham
Half course (spring term). Hours to be arranged.
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.
Note: Expected to be given in 2001–02.
Prerequisite: Biology 22, Anthropology 170, BS 25 or 80 or Science B-29 or permission of instructor.

Biology 187. Current Advances in Metazoan Diveristy and Evolution
Catalog Number: 3220
Gonzalo Giribet
Half course (spring term). Tu., Th., 11:30–1 and occasional laboratory sessions and field trips. EXAM GROUP: 13, 14

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.
Note: Expected to be given in 2001–02.

[*Biology 209. Phylogenetic Analysis: Seminar]
Catalog Number: 2534
David A. Baum
Half course (spring term). Hours to be arranged.
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.
Note: Expected to be given in 2001–02. Background in probability and statistics highly recommended.

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.

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.
Prerequisite: Biology 120 or permission of instructor.

Biology 214. Macroevolution of Interactions
Catalog Number: 7040 Enrollment: Limited to 15.
Brian D. Farrell and Naomi E. Pierce
Half course (spring term). W., 1–3. EXAM GROUP: 6, 7
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.
Prerequisite: BS 53 (formerly Biology 17 and 20), Biology 22 or equivalent, or permission of instructor.

[Biology 221. Bacterial Diversity]
Catalog Number: 1234
Colleen M. Cavanaugh
Half course (spring term). Hours to be arranged.
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.
Note: Expected to be given in 2001–02.
Prerequisite: BS 1 or 50 and 2 or 51, and BS 11 or equivalent, or permission of instructor.

Biology 224 (formerly Biology 143). Biology of the Fungi
Catalog Number: 1308
Donald H. Pfister
Half course (fall term). Hours to be arranged. EXAM GROUP: 12
This is an intensive course covering the morphology, classification, evolution, and diversity of the fungi, including both parasitic and saprophytic members. Attention is given to biological phenomena unique to fungi. Readings and discussion will draw from the primary literature. Students will apply a variety of techniques to study fungi in the field and in the laboratory.
Note: At least one weekend field trip to be arranged.
Prerequisite: Biology 113 or permission of instructor.

Biology 227. Molecular Approaches to Environmental Microbiology
Catalog Number: 4444
Colleen M. Cavanaugh
Half course (spring term). First Class meeting will be February 6th at 1pm, subsequent class meetings to bearranged.
Examination of the new understanding of microbial activities and biodiversity in the environment resulting from the application of cellular and molecular techniques. Critical review and discussion of advances in studies of bacterial diversity and community structure in aquatic and terrestrial environments, uncultivable bacteria, symbiotic associations, microenvironments, and genetic and metabolic capabilities of microorganisms in their natural habitats.
Note: First Class meets T., 2/6 at 1p in Biolabs Rm 2062.
Prerequisite: Earth and Planetary Sciences 30 and BS 11 or equivalent, or permission of instructor.

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

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. In even years, we will use the primary literature to explore a particular topic chosen by the course participants, including mating systems, population structure, genetic signatures of demographic history, etc. In odd years, the course will be a more basic exploration of the growing field of molecular ecology and be appropriate for advanced undergraduates.

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 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.
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). Fall: M., at 2; Spring: M., at 1. EXAM GROUP: Fall: 7; Spring: 6
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 (spring term). Tu., 2:30–4. EXAM GROUP: 16, 17
A seminar course that will discuss evolutionary aspects of genomic imprinting and the related theory of parent-offspring conflict.

Biology 255. Nature and Regulation of Marine Ecosystems
Catalog Number: 7753
James J. McCarthy and guest lecturers
Half course (fall term). Tu., 1:30–4. EXAM GROUP: 15, 16, 17
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.
Prerequisite: Biology 19 and 118.

Biology 261. Seminar in Evolution and Development
Catalog Number: 8451
James Hanken and David A. Baum
Half course (spring term). W., 1–3. EXAM GROUP: 6, 7
Graduate seminar in evolution and development. This seminar will evaluate contemporary problems and issues in the field of evolutionary developmental biology. Weekly meetings will include student presentations based on assigned readings, plus occasional guest speakers. Examples will be drawn from both plants and animals.

[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 2001–02.

*Biology 299r. Forest Practice and Research
Catalog Number: 6128
David R. Foster
Half course (fall term; repeated spring term). Fall: Tu., Th., at 10; Spring: W., at 5. EXAM GROUP: Fall: 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

*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 308. Evolution of Floral Developmental Mechanisms
Catalog Number: 5535
Elena M. Kramer 3791

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

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

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

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

*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

*Biology 334. Behavioral Ecology
Catalog Number: 8279
Naomi E. Pierce 2889

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

*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 (on leave fall term)

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

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

*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 389. Population Biology and Evolution
Catalog Number: 0680
Stephen R. Palumbi 2406 (on leave fall term)

*Biology 399 (formerly Biology 220). Topics in Organismic and Evolutionary Biology
Catalog Number: 0764
George V. Lauder 2375
Half course (spring term). Hours to be arranged.
Presents the research interests and experiences of scientists in organismic and evolutionary biology. Specific topics treated vary from year to year.
Note: Required of all first-year graduate students in Organismic and Evolutionary Biology.