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Amherst College Courses

Amherst College Courses

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Biology

Professors Clotfelter, Goutte (Chair), Hood‡, Miller†, Poccia†, Temeles, and Williamson*; Associate Professor Trapani; Assistant Professors Edwards, Jeong*, Purdy, and Ragkousi; Senior Lecturer Levin†; Visiting Assistant Professors Henderson-Stull, Lee, and Pater; Visiting Lecturer Goodwin; Lab Coordinators Emerson and Kristensen.

The Biology curriculum is designed to meet the needs of students preparing for postgraduate work in biology or allied careers, as well as to provide the insights of biology to other students whose area of specialization lies outside biology.

Courses for Non-Major Students. BIOL-104, 106, 108, 110 and 114 each focuses on a particular topic within biology and is specifically intended for students who do not major in biology. These courses will not count toward the Biology major and do not meet the admission requirements for medical school. The two semesters of introductory biology (BIOL-181 and 191) may also be taken by non-majors who wish a broad introduction to the life sciences.

Major Program. The Biology major consists of four categories:

  1. Two introductory biology courses (BIOL-181 and 191) taken in either order. BIOL-181 is offered in the fall semester. BIOL-191 is offered in both the fall and spring semesters. Please note that BIOL-191 requires completion of, or concurrent enrollment in, CHEM-161.
  2. Five advanced courses in Biology, three of which much involve laboratories and one of which must be at the 400 level. Additional stipulations for the advanced courses depend upon Class year (see below). Note that Special Topics and courses numbered below BIOL-181 do not fulfill this requirement and at least three of the five advanced courses must be completed at Amherst College, unless approved in advance by the Department.
    a) For the Class of 2022 and beyond, one advanced course in each of two core areas of biology is required by the end of the junior year:

    i) Cellular/Molecular Core:
    Cell Structure and Function (BIOL-291) or Molecular Genetics (BIOL-370/371).

    ii) Organismal Core:
    Ecology (BIOL-230) or Evolutionary Biology (BIOL-320/321).

    b) For the Class of 2021 and earlier, one course in each of three subject bins is required, two of which must be completed by the end of the junior year. These subject bins include:

    i) Molecular and cellular mechanisms of life processes: Cell Structure and Function (BIOL-291), Molecular Neurobiology (BIOL-301), Biochemical Principles (BIOL-330), Biochemistry (BIOL-331), Molecular Genetics (BIOL-370), Molecular Genetics with Lab (BIOL-371), and Biophysics (BIOL-400)

    ii) Integrative processes that show the relationship between molecular mechanisms and macroscopic phenomena: Introduction to Field Biology (BIOL-201), Plant Form, Function, and Diversity (BIOL-213), Developmental Biology (BIOL-221), Genetic Analysis of Biological Processes (BIOL-241), Form and Function (BIOL-264), Microbiology (BIOL-271), Microbiology with Lab (BIOL-271), Neurophysiology (BIOL-350) and Neurophysiology with Lab (BIOL-351), Immunology (BIOL-372), Genome Biology (BIOL-380) and Genome Biology with Lab (BIOL-381);

    iii) Evolutionary explanations of biological phenomena: Plants as Models in Organismal Biology (BIOL-211), Ecology (BIOL-230), Animal Behavior (BIOL-280), Animal Behavior with Lab (BIOL-281), Evolutionary Biology (BIOL-320) and Evolutionary Biology with Lab (BIOL-321).
  1. Two courses in chemistry (CHEM151 or 155 and CHEM161 or above) and two courses in mathematics, statistics, and/or physics at the level of MATH-111, STATS-111, and PHYS-116 or above. Students with Advanced Placement scores may satisfy the requirement by taking upper-level courses. CHEM-151 and/or CHEM-161 are requisites for several Biology courses. Students are therefore encouraged to take CHEM151/155 in the fall of their first year, particularly students whose planned courses emphasize cellular and molecular mechanisms. Students preparing for graduate study in life sciences should consider taking CHEM-221 and 231, PHYS-117, and a course in statistics in addition to the minimum requirements for the Biology major. Note that CHEM-221 and 231 are requisites for BIOL-331, and that prior completion of PHYS-117 or 124 is recommended for BIOL-351 and BIOL-400.
  2. Senior Biology Major Requirements: in their senior year, Biology majors must complete two comprehensive requirements: attendance at Departmental seminars (Mondays 4:00-5:00 p.m.) and a Senior Comprehensive Examination administered by the Department in the fall semester. All Biology majors and prospective Biology majors are also encouraged to attend the Biology seminars, which are open to the public.

Students with a Biology AP score of 5 who wish to place out of either Introductory Biology course must first seek permission from the Biology Department. Students placing out of BIOL-181 must substitute a course from the Organismal Core (2.a.ii) or Evolutionary explanations subject bin (2.b.iii); students placing out of BIOL-191 must substitute a course from the Cellular/Molecular Core (2.a.1) or Molecular and cellular mechanisms subject bin (2.b.i), according to Class year. Students placing out of BIOL-181 or BIOL-191, or in exceptional cases both, must take a minimum of four semesters of laboratory work (one intro lab and three upper-level labs or four upper-level labs).

Please note that courses taken as pass/fail cannot be counted toward the major. This includes biology courses as well as ancillary courses (i.e., chemistry, physics, and math). Also, a Biology course cannot be counted towards the Biology major if it is also being used to meet the requirements of another major, unless the course in question is required by both majors.

Departmental Honors Program. Honors work in Biology is an opportunity to do original laboratory or field research and to write a thesis based on this research. The topic of thesis research is chosen in consultation with a member of the Biology Department who agrees to supervise the Honors work. Honors candidates take three courses of thesis research in their senior year (BIOL-498 in the fall and BIOL-499 in the spring), usually with the double course in the spring (designated as BIOL-499D) in addition to the other requirements for the major, except that Honors candidates may take four rather than five advanced Biology courses, subject to the laboratory and subject area constraints. 

Courses for Premedical Students. Students not majoring in Biology may fulfill the two-course minimum premedical requirement in Biology by taking two laboratory courses numbered 181 or above in Biology. Students interested in health professions should consult a member of the Health Professions Committee regarding specific requirements and visit the Amherst Health Professions webpage.

*On leave 2018-19.
†On leave fall semester 2018-19.
‡On leave spring semester 2018-19.

104 Food, Fiber, and Pharmaceuticals

It is perhaps impossible to experience a day without plants. From the air we breathe, the bed we sleep in, the soap we wash with and clothes we put on, to the foods we consume and the medicines we take, we are very much dependent upon plants and their products. Through a combination of lecture, discussion, and observation, we will explore how, why, and when plants became vital to people and their societies. Several economically important plant groups will be studied, including those that provide food and beverages, medicines and narcotics, spices, perfumes, fuels, and fiber. What are the characteristics of these groups enabling their exploitation, and what is the history of these associations? How and when were plants domesticated and what are the consequences of large-scale agriculture? What impacts do human population growth and habitat destruction have on the ways that people interact with plants now and in the future?  Finally, we will explore the role of technology in efforts to both improve and synthesize plant products. Three classroom hours per week. Two local field trips.

Limited to 26 students. This course is for non-majors. Students majoring in Biology will be admitted only with consent of the instructor. Omitted 2018-19. Senior Lecturer Levin.

2023-24: Not offered
Other years: Offered in Spring 2013, Fall 2013, Fall 2015, Fall 2016, Fall 2017, Spring 2020

114 The Evolution of Human Nature

(Offered as BIOL 114 and ANTH 114) After consideration of the relevant principles of animal behavior, genetics, and population biology, it will be shown that extensions of the theory of natural selection—kin selection, reciprocal altruism, parent-offspring conflict, sexual selection, and parental manipulation of sex ratios—provide unifying explanations for the many kinds of social interactions found in nature, from those between groups, between individuals within groups and between genes within individuals. The emphasis throughout will be on the special physical, social and psychological adaptations that humans have evolved, including the instincts to create language and culture, conflict and cooperation within and between the sexes, moral emotions, the mating system and family, kinship and inheritance, reciprocity and exchange, cooking, long distance running, homicide, socioeconomic hierarchies, warfare, patriarchy, religions and religious beliefs, deceit and self-deception, systems of laws and justice and the production, performance and appreciation of art. Along the way, we will consider how misrepresentations of evolutionary theory have been used to support political and social ideologies and, more recently, to attack evolutionary theory itself as scientifically flawed and morally corrupt.

Limited to 30 students. Omitted 2018-19. Professor Emeritus Zimmerman.

2023-24: Not offered
Other years: Offered in Spring 2008, Spring 2010, Spring 2011, Spring 2016, Spring 2017, Spring 2018

213 Plant Form, Function, and Diversity

Plants are one of the most successful and abundant groups of organisms on earth, comprising the majority of terrestrial biomass, being integral to ecosystem structure, and providing humans with food, shelter, and materials. Yet plants face extraordinary challenges given their relative immobility and, in response, have evolved a surprising suite of behaviors and morphologies. Plants can tolerate extreme abiotic environments, manipulate animals for services (protection, pollination, dispersal), and travel to the far corners of the planet. In this course, we will explore the range of plant diversity, form and function. This will include an introduction to basic principles of plant function including physical processes occurring in plants, water relations in whole plants and plant tissues, cell physiology and biochemistry, and growth and development. In the laboratory, students will explore plant diversity and anatomy, as well as conduct genetic, biochemical, and physiological studies. The course includes three hours of lecture and three hours of laboratory per week.

Requisite: BIOL 181. Recommended: BIOL 191. Limited to 16 students. Not open to first-year students. Fall semester. Professor Pater.

2023-24: Not offered
Other years: Offered in Fall 2018

214 Neurobiology

(Offered as BIOL 214 and NEUR 214) An introduction to the structure and function of the nervous system, this course explores the basic functions of neurons and synapses as well as neural mechanisms of sensation at molecular, cellular, circuit and system levels. Basic topics in neurobiology and neurophysiology will be covered with emphases on neuroscience history and understanding how neuroscientists approach the study of the nervous system. Three class hours per week.

Requisite: BIOL 191. Limited to 45 students. Omitted 2018-19. Professor Trapani.

Other years: Offered in Fall 2019, Fall 2020, Fall 2021, Fall 2022, Spring 2023, Fall 2023, Spring 2025

250 “According to Science…” ; Scientific Communication and Literacy in Biology

We’ve all seen the headlines claiming that scientists have proven everything from coffee consumption increasing your lifespan, sugar being as addictive as cocaine, and Alzheimer’s being caused by lack of sleep. In this course we will review media coverage of scientific findings and contrast that coverage with the primary literature upon which the articles or documentaries were based. Students will have the opportunity to lead discussion and present independent literature research in both oral and written format. Students will be encouraged to further improve their science communications skills by publishing their findings to a class blog. 

Requisite: Two Amherst College introductory level courses in the fields of Biology, Chemistry, or Physics. Limited to 20 students. Not open to first-year students. This course does not count toward the Biology major. Spring semester. Professor Pater.

2023-24: Not offered
Other years: Offered in Spring 2020, January 2021, January 2022, Spring 2022

264 Form and Function

Functional morphology is the study of relationships between anatomical structures and the ecology and behavior of organisms. For example, how does a bird’s wing produce flight? Can we predict differences in bird flight habits based on wing shape or feather structure? How do wings of flying and flightless birds differ? The course begins by focusing on the importance of animal body size and metabolism before moving on to various forms of locomotion, and then to morphological adaptations for prey capture, predator avoidance, and reproduction. We will also discuss how animal morphology inspires human innovation. The course includes a combination of lecture, discussion, and practical activities. Three hours per week.

Requisite: BIOL 181. Not open to first-year students. Limited to 24 students. Omitted 2018-19. Professor Clotfelter.

Other years: Offered in Spring 2017, Spring 2020, Spring 2022, Spring 2025

270 Microbiology

Microbes inhabit the world's oceans, deserts, lakes, soils, and atmosphere, and play a vital role in the Earth's biogeochemical cycles. As humans, we harbor a diverse microbial flora estimated to outnumber our own human cells. During this course, we will explore this microbial world by investigating the structure, physiology, genetics, and evolution of microorganisms with a focus on bacteria, but including discussions of archaea, viruses, and microbial eukaryotes. The goal of the course is to gain an understanding of the unique properties of microbes that enable their persistence and diversification. We will also pay special attention to microbial interactions with eukaryotic organisms, by studying both host and microbe contributions to virulence, mutualism, and symbiotic relationships. Three hours of lecture and one hour of discussion per week.

Requisite: BIOL 191. Recommended: BIOL 181. Limited to 15 students. Not open to first-year students. Spring semester. Professor Purdy.

Other years: Offered in Spring 2018, Spring 2019, Fall 2022

271 Microbiology with Lab

Microbes inhabit the world's oceans, deserts, lakes, soils, and atmosphere, and play a vital role in the Earth's biogeochemical cycles. As humans, we harbor a diverse microbial flora estimated to outnumber our own human cells. During this course, we will explore this microbial world by investigating the structure, physiology, genetics, and evolution of microorganisms with a focus on bacteria, but including discussions of archaea, viruses, and microbial eukaryotes. The goal of the course is to gain an understanding of the unique properties of microbes that enable their persistence and diversification. We will also pay special attention to microbial interactions with eukaryotic organisms by studying both host and microbe contributions to virulence, mutualism, and symbiotic relationships. Laboratory exercises will include explorations of microbial functions and diversity in a variety of contexts using both classical and molecular approaches. Three hours of lecture, three hours of laboratory and one hour of discussion per week.

Requisite: BIOL 191. Recommended: BIOL 181. Limited to 15 students. Not open to first-year students. Spring semester. Professor Purdy.

2023-24: Not offered
Other years: Offered in Spring 2018, Spring 2019

281 Animal Behavior with Lab

Shaped by millions of years of evolution, animals have evolved myriad abilities to respond to their environment, their potential predators and prey, and members of their own species. This course examines animal behavior from both a mechanistic and a functional perspective. Drawing upon examples from a diverse range of taxa, and using articles from the primary scientific literature, we will discuss topics such as behavioral endocrinology, sexual selection and mating systems, animal communication, and kinship and cooperation. Four classroom hours and three laboratory hours per week; the laboratory projects will require additional time outside of class.

Requisite: BIOL 181. Limited to 15 students per lab section. Spring semester. Visiting Lecturer Goodwin and Lab Coordinator Kristensen.

Other years: Offered in Fall 2011, Fall 2013, Fall 2022, Fall 2023

330 Biochemical Principles of Life at the Molecular Level

(See CHEM 330)

350 Neurophysiology

(Offered as BIOL 350 and NEUR 350)  This course will provide a deeper understanding of the physiological properties of the nervous system. We will address the mechanisms underlying electrical activity in neurons, as well as examine the physiology of synapses; the transduction and integration of sensory information; the function of nerve circuits; the trophic and plastic properties of neurons; and the relationship between neuronal activity and behavior. Laboratories will apply electrophysiological methods to examine neuronal activity and will include experimental design as well as analysis and presentation of collected data. Throughout the course, we will focus on past and current neurophysiology research and how it contributes to the field of neuroscience. Three classroom lecture hours, plus a fourth discussion hour to be used for group work, paper presentations, and review sessions.

Requisites: BIOL 191 and CHEM 151; PHYS 117 or 124 is recommended. Limited to 12 students. Open to juniors and seniors. Fall semester. Professor Trapani.

2023-24: Not offered
Other years: Offered in Fall 2007, Fall 2008, Spring 2010, Fall 2012, Fall 2013, Fall 2016, Fall 2017, Fall 2018, Fall 2019, Fall 2020, Fall 2021

370 Molecular Genetics

(Offered as BIOL 370 and BCBP 370) A study of the molecular mechanisms underlying the transmission and expression of genes. DNA replication and recombination, RNA synthesis and processing, and protein synthesis and modification will be examined. Both prokaryotic and eukaryotic systems will be analyzed, with an emphasis upon the regulation of gene expression. Application of modern molecular methods to biomedical and agricultural problems will also be considered. Four classroom hours per week.

Requisite: BIOL 191 or equivalent. Limited to 15 students. Open to juniors and seniors.  Omitted 2018-19.

 

2023-24: Not offered
Other years: Offered in Fall 2008, Fall 2010, Fall 2011, Fall 2013, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Fall 2019, Spring 2020, Fall 2020, Fall 2021

371 Molecular Genetics W/Lab

(Offered as BIOL 371 and BCBP 371) A study of the molecular mechanisms underlying the transmission and expression of genes. DNA replication and recombination, RNA synthesis and processing, and protein synthesis and modification will be examined. Both prokaryotic and eukaryotic systems will be analyzed, with an emphasis upon the regulation of gene expression. Application of modern molecular methods to biomedical and agricultural problems will also be considered. The laboratory component will focus upon recombinant DNA methodology. Four classroom hours and four hours of laboratory per week.

Requisite: BIOL 191 or equivalent. Limited to 15 students. Open to juniors and seniors. Fall semester. Visiting Assistant Professor Lee.

Other years: Offered in Fall 2017, Fall 2018, Fall 2019, Spring 2020, Fall 2020, Fall 2021, Fall 2024, Spring 2025

376 Epigenetics

Genes can undergo heritable modifications that alter gene expression without altering genetic information. Such modifications are known as epigenetic modifications, and are heritable through cell division (mitotic inheritance) or from parents to progeny (meiotic inheritance). Epigenetic modifications can be influenced by multiple biological and environmental factors, thus providing an added layer to the relationship between genotype and phenotype. Understanding how this layer of regulation works is becoming an important aspect of molecular genetics. In this course, we will cover basic concepts in epigenetics and discuss various epigenetic processes such as DNA methylation, chromatin remodeling, gene imprinting, post-translational histone modification, epigenomics, environmental epigenetics and the relationship between epigenetic modification and human health. Three hours of lecture per week plus one fourth hour per week.

Requisite: BIOL 191. Limited to 30 students. Spring semester. Visiting Assistant Professor Lee.

2023-24: Not offered
Other years: Offered in Spring 2019

400 Molecular and Cellular Biophysics

(See PHYS 400)

430 Seminar in Behavioral Ecology

This course will explore the relationship between an animal's behavior and its social and ecological context. The topic for 2018 will be the evolution of sexual dimorphism in animals. Sexual dimorphism is widespread in animals, yet its causes remain controversial and have generated much debate. In this seminar, we will examine a variety of sexual dimorphisms in different groups of animals and consider hypotheses for how these sexual dimorphisms may have evolved. We will then consider how these hypotheses are tested in an attempt to identify the best approaches to studying the evolution of sexual dimorphism. Then we will look at evidence that either supports or refutes various hypothesized mechanisms for the evolution of sexual dimorphisms in different animal groups. Finally, we will consider whether some mechanisms for the evolution of sexual dimorphism are more convincing among certain kinds of organisms than others. Three hours per week.

Requisite: One or more courses from BIOL 181, BIOL 230, BIOL 280, BIOL 281, BIOL 320, BIOL 321 or consent of the instructor. Not open to first-year students or to students who have taken the seminar in previous years. Limited to 15 students. Fall semester. Professor Temeles.

2023-24: Not offered
Other years: Offered in Spring 2010, Spring 2012, Fall 2013, Fall 2018, Fall 2019

454 Seminar in Tropical Biology

Most biodiversity on our planet can be found in tropical latitudes. Tropical rainforests, for example, which account for less than 10% of the Earth’s surface, may contain 50-75% of all plant and animal species. In this course we will examine some of the myriad biotic interactions that occur in the tropics using an ecological, evolutionary, and behavioral approach. The course is divided into two main components: a two-week interterm field course and an advanced seminar. During the field course students will be immersed in the tropical forests of Costa Rica, with a focus on organismal identification and comparisons between three sites (lowland Caribbean rainforest, cloud forest, and coastal dry forest). While in Costa Rica, we will utilize the expertise of local specialists to learn more about taxonomic groups that are particularly significant in the tropics, such as bats, frogs, and epiphytic plants. Back on campus, students will analyze tree diversity data collected in the field, as well as focus on a specific topic within tropical biology, around which students will design experimental research that will be presented in an NSF-style grant proposal. Three hours per week.

Requisite: BIOL 181. Recommended: one or more of the following courses: BIOL 230/ENST-210, BIOL 201, BIOL 280/281, BIOL 320/321. Not open to first-year students. Limited to 12 students. Admission with consent of the instructor. Spring semester. Professor Clotfelter and Senior Lecturer Levin.

457 Seminar in Developmental Genetics

This seminar will focus on understanding germ cells. Although germ cells do not contribute to the form or function of an individual, they have the important role of providing the continuity of life between generations. In many animals, they are among the first cells to be differentiated from others during embryonic development. Elaborate mechanisms ensure that the genetic information in these reproductive cells is protected and packaged in unique ways to be used in the next generation. We will explore primary literature readings that probe the specialized development and genetic regulation of germ cells in model systems such as mice, flies, and nematodes. Classic developmental genetic approaches as well as more modern molecular genetic methods will be discussed as ways of understanding these extraordinary cells. Students will use oral and written formats to present paper reviews and analyses. Three classroom hours a week.   

Prerequisite: One of the following: BIOL 221, BIOL 241, BIOL 291, BIOL 370, BIOL 371 or consent of the instructor. Limited to 15 students. Fall Semester. Professor Goutte.

2023-24: Not offered
Other years: Offered in Fall 2018, Spring 2020

470

Seminar in Biology of Metals: A Molecular/Cellular Perspective 

Metals are required for the function of about one third of all proteins and are involved in vital biological processes such as photosynthesis, respiration, gene regulation, DNA replication and repair, signal transduction, and antioxidant defense. However, essential metals are potentially toxic due to the same properties that make them indispensable. To cope with such a paradox, metals must be tightly regulated.

This advanced seminar will focus on the molecular and cellular biology of metals. Topics of discussion will include metal homeostasis strategies (e.g. import/export, chelation, subcellular compartmentalization), metal cofactors of biochemical processes, inherited metal metabolism disorders, and genetics of hyperaccumulators. We will also discuss prospects of manipulating metal homeostasis to aid human health and environmental sustainability. The course will consist of discussions of primary literature and student presentations. Assignments will include written reviews of literature.

Requisite: One of the following courses, BIOL 241, 251, 291, 330, 331, or consent of the instructor. Limited to 15 students. Omitted 2018-19. Professor Jeong.

2023-24: Not offered
Other years: Offered in Spring 2016, Spring 2022

Departmental Courses

110 Contagion

Infection by contagious microorganisms remains a leading cause of death in many parts of the world. This course will explore the biological mechanisms of infectious diseases, as well as the challenges associated with fighting their emergence and spread. We will focus on diseases of global health importance, such as HIV/AIDS, cholera, and tuberculosis, to discuss the strategies pathogens have evolved that ensure their successful transmission. In light of their ability to effectively outwit our own immune systems, we must devise new means to overcome these disease-causing microbes. Here, the challenges are legion. We will see that the answer lies not only with an understanding of biology to formulate treatments and prevention measures, but this knowledge must be integrated with awareness of complex societal issues to inform and implement solutions. Discussions will focus upon the many perspectives from which infectious diseases are encountered, drawing on resources from the literature on microbiology, ethics, and policy, as well as personal accounts and current news stories. Three hours of lecture and discussion per week. This course is for non-science majors and will not count toward the Biology major.

Limited to 40 students. Omitted 2018-19. Professor Purdy.

2023-24: Not offered
Other years: Offered in Spring 2013, Spring 2014, Fall 2017, Spring 2021

181 Adaptation and the Organism

An introduction to the evolution, ecology, and behavior of organisms and how these relate to the diversity of life. Following a discussion of the core components of evolutionary theory, we'll examine how evolutionary processes have shaped morphological, anatomical, physiological, and behavioral adaptations in organisms that solve many of life's problems, ranging from how to find or acquire food and avoid being eaten, to how to attract and locate mates, and how to optimize reproduction throughout a lifetime. We'll relate and compare characteristics of animals, plants, fungi, protists, and bacteria, examining how and why these organisms have arrived at various solutions to life's problems. Laboratory exercises will complement lectures and will involve field experiments on natural selection and laboratory studies of vertebrates, invertebrates and plants. Four classroom hours and three laboratory hours per week.

Fall semester. Professors Clotfelter and Hood; Lab Coordinators Kristensen and Goodwin.

Please note that the same textbook is used for both BIOL 181 and BIOL 191.

Other years: Offered in Spring 2012, Spring 2013, Spring 2014, Spring 2015, Spring 2016, Spring 2017, Spring 2018, Fall 2018, Fall 2021, Fall 2022, Fall 2023, Fall 2024

191 Molecules, Genes and Cells

An introduction to the molecular and cellular processes common to life with an emphasis on control of energy and information flow. Central themes include metabolism, macromolecular function, and the genetic basis of cellular function. We examine how membranes work to establish the internal composition of cells; how the structure of proteins including enzymes affects protein function; how energy is captured, stored and utilized by cells; and how cells communicate, move and divide. We explore inheritance patterns and underlying molecular mechanisms of genetics, the central dogma of information transfer from DNA replication to protein synthesis, and recombinant DNA methods and medical applications. Laboratories include genetic analyses, enzyme reaction kinetics, membrane transport, and genomic analysis. Two hours of lecture, two hours of team-based learning, and three laboratory hours per week.

Requisite: Prior completion of, or concurrent registration in, CHEM 161. Limited to 96 students. Fall and spring semesters. The Department.

Please note that the same textbook is used for both BIOL 181 and BIOL 191.

Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Spring 2019, Fall 2022, Spring 2023, Fall 2023, Fall 2024, Spring 2025

201 Introduction to Field Biology: Disease Ecology

Advances in organismal biology hinge upon an understanding of natural history and are enhanced by quantitative observation, hypothesis formation, and experimentation with systems that occur in nature. In this course, we will apply these principles specifically to the study of infectious diseases in natural populations. With a combination of lecture, discussion, and field-based activities, the course will focus on deriving important questions and the variety of approaches to address them. While covering the fundamentals of disease ecology, the applicability of the field-based approaches to other areas of organismal biology will be emphasized as a foundation for further studies. Three classroom hours and three laboratory/field work hours per week.

Requisite: BIOL 181. Limited to 16 students. Omitted 2018-19. Professor Hood.

Other years: Offered in Fall 2023, Fall 2024

221 Developmental Biology

How can a single cell, the fertilized egg, give rise to all the specialized cells of an adult? What gives rise to biological form? What is the molecular logic of the pathways that progressively refine cellular identities? How do cells "talk" to one another so as to coordinate their behaviors as embryos develop form and function? How can parts of an organism be regenerated with only the appropriate regions remade, structured identically to the missing ones? How does a stem cell differ from a non-stem cell? How can genetically identical organisms be cloned? This course will offer an integrative study of the development of animals, leading to the formulation of the principles of development, including an introduction to experimental embryology and developmental physiology, anatomy, genetics and "evo-devo." Laboratory work explores embryonic development and regeneration in amphibians, sea urchins, nematodes, flatworms, fruit flies, fish, and chickens. Three classroom hours plus three hours of laboratory per week.

Requisite: BIOL 191. Not open to first-year students. Limited to 16 students. Fall Semester. Professors Ragkousi.

Other years: Offered in Fall 2014, Spring 2016, Fall 2022, Fall 2024

230 Ecology

(Offered as BIOL 230 and ENST 210) A study of the relationships of plants and animals (including humans) to each other and to their environment. We'll start by considering the decisions an individual makes in its daily life concerning its use of resources, such as what to eat and where to live, and whether to defend such resources. We'll then move on to populations of individuals, and investigate species population growth, limits to population growth, and why some species are so successful as to become pests whereas others are on the road to extinction. The next level will address communities, and how interactions among populations, such as competition, predation, parasitism, and mutualism, affect the organization and diversity of species within communities. The final stage of the course will focus on ecosystems, and the effects of humans and other organisms on population, community, and global stability. Three hours of lecture per week.

Requisite: BIOL 181 or ENST 120 or equivalent. Limited to 50 students. Fall and spring semesters. Professor Temeles.

Other years: Offered in Spring 2012, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Spring 2019, Spring 2020, Fall 2020, Fall 2021, Spring 2023, Fall 2024

241 Genetic Analysis of Biological Processes

In this course we will explore genetic analysis as a means of probing the mysteries of the molecular world. Scientists often turn to the study of genes and mutations when trying to decipher the molecular mechanisms that underlie such diverse processes as the making of an embryo, the response of cells to their environment, or the defect in a heritable disease. All of the reading in the course will be from the primary literature, where students will engage with data from genetic experiments that shed light on the workings of a signal transduction pathway. Students will learn from these examples how to use genetic analysis to formulate models that explain the molecular function of a gene product. In the laboratory students will apply these approaches to their own semester-long project, taking responsibility for experimental design and execution as well as data interpretation and analysis. Three hours of lecture and four hours of laboratory per week; the laboratory projects will require time outside of class hours.

Requisite: BIOL 191. Limited to 24 students. Not open to first-year students. Omitted 2018-19. Professor Goutte.

2023-24: Not offered
Other years: Offered in Spring 2012, Spring 2013, Spring 2014, Spring 2015, Spring 2017, Spring 2018, Spring 2022, Spring 2023, Spring 2025

260 Animal Physiology

This course will examine the function of tissues, organs, and organ systems, with an emphasis on the relationship between structure and function. Building outward from the level of the cell, we will study bodily processes including respiration, circulation, digestion and excretion. In addition, the course will address how different organisms regulate these complex processes and how ion and fluid balance is maintained. We will also study the nervous system in the context of sensory systems, focusing on how external stimuli are transformed into meaningful neuronal signals and processed by the brain. Weekly discussions will include readings from primary literature. Four classroom hours per week.

Requisite: BIOL 191 and either BIOL 181 or NEUR 226. Omitted 2018-19. Professor Trapani.

Other years: Offered in Fall 2011, Spring 2013, Spring 2017, Spring 2021, Fall 2021, Spring 2025

280 Animal Behavior

Shaped by millions of years of evolution, animals have evolved myriad abilities to respond to their environment, their potential predators and prey, and members of their own species. This course examines animal behavior from both a mechanistic and a functional perspective. Drawing upon examples from a diverse range of taxa, and using articles from the primary scientific literature, we will discuss topics such as behavioral endocrinology, sexual selection and mating systems, animal communication, and kinship and cooperation. FourThree classroom hours.

Requisite: BIOL 181. Limited to 20 students. Spring semester. Visiting Lecturer Goodwin.

Other years: Offered in Spring 2015, Fall 2015, Fall 2016, Fall 2017, Spring 2019, Fall 2019, Fall 2022, Fall 2023, Fall 2024

291 Cell Structure and Function

(Offered as BIOL 291 and BCBP 291) An analysis of the structure and function of cells in plants, animals, and bacteria. Topics to be discussed include the cell surface and membranes, cytoskeletal elements and motility, cytoplasmic organelles and bioenergetics, the interphase nucleus and chromosomes, mitosis, meiosis, and cell cycle regulation. Four classroom hours and three hours of laboratory per week.

Requisite: BIOL 191. Limited to 28 students. Spring semester. Professors Edwards and Poccia.

Other years: Offered in Spring 2012, Spring 2013, Spring 2014, Spring 2015, Spring 2016, Spring 2017, Spring 2018, Spring 2019, Spring 2023, Fall 2023, Fall 2024

301 Molecular Neurobiology

(Offered as BIOL 301 and NEUR 301)  An analysis of the molecules and molecular mechanisms underlying nervous system function, development, and disease. We will explore the proteins that contribute to the unique structure and function of neurons, including an in-depth analysis of synaptic communication and the molecular processes that modify synapses.  We will also study the molecular mechanisms that control brain development, from neurogenesis, neurite growth, and synaptogenesis to cell death and degeneration. In addition to analyzing neural function, throughout the course we will also study nervous system dysfunction resulting when such molecular mechanisms fail, leading to neurodevelopmental and neurodegenerative disease. Readings from primary literature will emphasize current molecular techniques utilized in the study of the nervous system. Four classroom hours and three hours of laboratory per week.

Requisite: BIOL 191 and CHEM 161. Not open to first-year students. Admission with consent of the instructor.  Limited to 24 students. Omitted 2018-19. Members of the Department.

Other years: Offered in Fall 2011, Fall 2014, Fall 2015, Fall 2016, Spring 2020, Spring 2021, Spring 2022, Fall 2023, Fall 2024

320 Evolutionary Biology

Evolution is a powerful and central theme that unifies the life sciences. In this course, emphasis is placed on microevolutionary mechanisms of change, and their connection to large-scale macroevolutionary patterns and diversity. Through lectures and readings from the primary literature, we will study genetic drift and gene flow, natural selection and adaptation, molecular evolution, speciation, the evolution of sex and sexual selection, life history evolution, and inference and interpretation of evolutionary relationships. Three hours of lecture and one hour of discussion each week.

Requisite: BIOL 181 and BIOL 191. This course is offered for those students not taking the associated laboratory (BIOL 321) and the combined enrollment for these courses will be 30 students. Not open to first-year students. Spring semester. Professor Miller.

Other years: Offered in Spring 2023, Spring 2025

321 Evolutionary Biology with Lab

Evolution is a powerful and central theme that unifies the life sciences. In this course, emphasis is placed on microevolutionary mechanisms of change, and their connection to large-scale macroevolutionary patterns and diversity. Through lectures and readings from the primary literature, we will study genetic drift and gene flow, natural selection and adaptation, molecular evolution, speciation, the evolution of sex and sexual selection, life history evolution, and inference and interpretation of evolutionary relationships. The laboratory investigates evolutionary processes using computer simulations, artificial selection experiments, and a semester-long project that characterizes phenotypic breeding relationships among individuals and integrates these results with analyses of molecular sequence variation for genes contributing to mating recognition. Three hours of lecture, one hour of discussion and four hours of laboratory work each week.

Requisite: BIOL 181 and BIOL 191. Limited to 16 students. Not open to first-year students. Spring semester. Professor Miller.

Other years: Offered in Fall 2011, Fall 2012, Spring 2016, Spring 2017, Spring 2019

331 Biochemistry

(Offered as BIOL 331, BCBP 331, and CHEM 331) Structure and function of biologically important molecules and their role(s) in life processes. Protein conformation, enzymatic mechanisms and selected metabolic pathways will be analyzed. Additional topics may include: nucleic acid conformation, DNA/protein interactions, signal transduction and transport phenomena. Four classroom hours and four hours of laboratory work per week. Offered jointly by the Departments of Biology and Chemistry. A student may not receive credit for both BCBP/BIOL/CHEM 331 and CHEM 330. 

Requisite: CHEM 221 and BIOL 191; or consent of the instructor. CHEM 231 is a co-requisite. Limited to 30 students. Spring semester. The Department. 

Other years: Offered in Spring 2012, Spring 2013, Spring 2014, Spring 2015, Spring 2016, Spring 2017, Spring 2018, Spring 2019, Fall 2019, Fall 2020, Fall 2021, Fall 2022, Fall 2023, Fall 2024

351 Neurophysiology with Lab

(Offered as BIOL 351 and NEUR 351) This laboratory course will provide a deeper understanding of the physiological properties of the nervous system. We will address the mechanisms underlying electrical activity in neurons, as well as examine the physiology of synapses; the transduction and integration of sensory information; the function of nerve circuits; the trophic and plastic properties of neurons; and the relationship between neuronal activity and behavior. Laboratories will apply electrophysiological methods to examine neuronal activity and will include experimental design as well as analysis and presentation of collected data. Throughout the course, we will focus on past and current neurophysiology research and how it contributes to the field of neuroscience. Lecture meetings will be combined with BIOL 350 students for three classroom hours plus a fourth hour to be used for group work, paper presentations, and review sessions. Three hours of laboratory work per week.

Requisites: BIOL 191 and CHEM 151; PHYS 117 or 124 is recommended. Limited to one lab section with 18 students. Open to juniors and seniors. Fall semester. Professor Trapani.

2023-24: Not offered
Other years: Offered in Fall 2012, Fall 2013, Fall 2016, Fall 2022

372 Immunology

The immune response is a consequence of the developmentally programmed or antigen-triggered interaction of a complex network of interacting cell types. These interactions are controlled by regulatory molecules and often result in the production of highly specific cellular or molecular effectors. This course will present the principles underlying the immune response and introduce the methodology of immunological research. Examples from clinical immunology will illustrate the recent impact of immunotherapies on the treatment of disease. In addition to lectures, tutorials will be used to provide a detailed introduction to the research literature of immunology. The class will meet twice per week and will also include periodic individual tutorials by appointment.

Requisite: BIOL 191 and two of the following or their equivalents: 241, 251, 271, 291, 310, 330, 331, 351, 380, 381,400, 420, or consent of the instructor. Limited to 15 students. Omitted 2018-19. Emeritus Professor Goldsby.

2023-24: Not offered
Other years: Offered in Fall 2011, Spring 2014, Fall 2017, Fall 2019

380 Genome Biology

A study of the architecture and interactions of genetic systems. Advances in genomics are providing insights into a variety of important issues, from the structural limits of DNA-based inheritance to the discovery of novel infectious and genetic diseases. We will address how heritable information is organized in different groups of organisms. We will also cover a major challenge of this emerging field—the application of vast amounts of genetic data to understanding genomic integrity and regulation. We will critically assess the genome as a "cooperative assemblage of genetic elements" and conclude by discussing the consequences of genomic structure for shaping species traits and long-term evolutionary potential. Three hours of lecture per week. 

Requisite: BIOL 181 and 191. This course is designed as an overflow class for those who cannot take BIOL 381 and the combined enrollment for these courses will be 30 students. Omitted 2018-19. Professor Hood.

Other years: Offered in Spring 2012, Spring 2013, Spring 2014, Spring 2017, Spring 2018, Spring 2020, Spring 2021, Spring 2022, Spring 2025

381 Genome Biology with Lab

A study of the architecture and interactions of genetic systems. Advances in genomics are providing insights into a variety of important issues, from the structural limits of DNA-based inheritance to the discovery of novel infectious and genetic diseases. We will address how heritable information is organized in different groups of organisms. We will also cover a major challenge of this emerging field—the application of vast amounts of genetic data to understanding genomic integrity and regulation. We will critically assess the genome as a "cooperative assemblage of genetic elements" and conclude by discussing the consequences of genomic structure for shaping species traits and long-term evolutionary potential. Three hours of lecture, and three hours of laboratory per week. Lab activities will require work outside of the scheduled meeting times.

Requisite: BIOL 181 and 191. Limited to 18 students. Omitted 2018-19. Professor Hood.

2023-24: Not offered
Other years: Offered in Spring 2012, Spring 2014, Spring 2018, Spring 2020, Spring 2021, Spring 2022, Spring 2025

404 Seminar in Biochemistry

(Offered as BIOL 404 and BCBP 404) The topic of this advanced seminar will be cholesterol. It has been said that more Nobel prizes have been awarded for the study of cholesterol than any other biological topic, yet it is astonishing how much we have learned only in the last few years, and how much we still don't understand. The topics in this course will include biosynthesis, transport, regulation, physiology, and biophysics of cholesterol. In many cases, these subjects illuminate or are illuminated by cholesterol-related diseases, so the biochemical bases for high cholesterol medications and for a genetic propensity for getting heart disease from eating broccoli are likely to come up. The course will be based on the scientific literature, and will include writing and presentation assignments.

Requisite: BIOL 191 and 291 or 331 or equivalent. Limited to 18 students. Omitted 2018-19. Professor Williamson. 

2023-24: Not offered
Other years: Offered in Spring 2011, Fall 2016

424 Seminar in Evolution: Sex and Sexual Reproduction

The origin and maintenance of sexual reproduction stands as one of the great mysteries of evolutionary biology. This seminar will explore the nature of sex and sexual reproduction across organisms, consider hypotheses for its origin and maintenance, and study its diverse consequences in populations. Readings will incorporate articles from the primary literature and topics for consideration include the molecular machinery and origin of meiosis, variation in sex determination mechanisms (including the evolution of sex chromosomes), sex ratio evolution, mating system variation, sexual conflicts, and the evolutionary ecology of sex differences. Three hours per week.

Requisite: BIOL 181, BIOL 191, and one upper-level course in Biology. Limited to 16 students. Omitted 2018-19. Professor Miller.

Other years: Offered in Spring 2023, Fall 2023, Spring 2025

434 Seminar in Ecology: Plant-animal Interactions

Most animals on Earth obtain their energy from green plants, and, thus, it is not surprising that interactions between plants and animals have played a prominent role in our current understanding of how ecological processes such as predation, parasitism, and mutualism shape evolutionary patterns in plants and animals. The main topics that we will discuss in this seminar include pollination, fruit and seed dispersal, deception, herbivory, and phytocarnivory, considering both ecological and evolutionary perspectives. We will also examine the biodiversity consequences of the loss of these associations via human-induced environmental change. Class readings emphasize the relevant primary literature. Students will have the opportunity to lead discussion and present independent literature research in both oral and written format. Three classroom hours per week.

Requisite: One of the following Biology courses: BIOL 201, 211, 230, 280/1, 320/1, or consent of the instructor. Limited to 15 students. Not open to first-year students. Spring semester. Professor Temeles.

2023-24: Not offered
Other years: Offered in Fall 2011, Spring 2013, Fall 2014, Spring 2016, Spring 2019, Spring 2020, Spring 2021, Fall 2022, Fall 2024

440 Seminar in Conservation Biology

(Offered as BIOL 440 and ENST 441) Conservation biology is a highly interdisciplinary field, requiring careful consideration of biological, economic, and sociological issues. Solutions to biodiversity conservation and environmental challenges are even more complex. Yet, conservation is a topic of timely importance in order to safeguard biological diversity. Utilizing articles from the primary literature, course topics will include invasive species, restoration, climate change, and biodiversity banking, as well as how to determine appropriate conservation priorities. Three classroom hours per week.

Requisite: BIOL 230/ENST 210 or consent of the instructor. Not open to first-year students. Limited to 14 students. Omitted 2018-19. Senior Lecturer Levin.

2023-24: Not offered
Other years: Offered in Spring 2012, Fall 2013, Fall 2015, Fall 2016, Spring 2018, Fall 2019, Fall 2020, Spring 2022, Spring 2023, Spring 2025

450 Seminar in Physiology: Classic Papers in Neurophysiology

(Offered as BIOL 450 and NEUR 450) Concentrating on reading and interpreting primary research, this course will focus on classic and soon-to-be classic neurophysiology papers. We will discuss the seminal experiments performed in the 1950s that led to our understanding of action potentials; experiments in the 1960s and 1970s that unlocked how synapses function; and more recent research that combines electrophysiology with optical methods and genetic techniques to investigate the role of many of the molecular components predicted by the work from the earlier decades. Assignments will include written reviews of literature as well as oral presentations.

Requisite: PHYS 117 or PHYS 124 and one of NEUR 226, BIOL 260, BIOL 351, or consent of the instructor. Limited to 18 students. Not open to first-year students. Spring semester. Professor Trapani.

2023-24: Not offered
Other years: Offered in Spring 2013, Spring 2017, Spring 2019, Spring 2022, Spring 2025

Senior Departmental Honors Courses

498, 499D Senior Departmental Honors

Honors students take three courses of thesis research, usually, but not always, with the double course load in the spring. The work consists of seminar programs, individual research projects, and preparation of a thesis on the research project.

Open to seniors. Spring semester. The Department.

Other years: Offered in Spring 2012, Spring 2013, Spring 2014, Spring 2015, Spring 2016, Spring 2017, Spring 2018, Spring 2019, Spring 2020, Spring 2021, Spring 2022, Spring 2023, Spring 2025

Special Topics Courses

290, 290H, 490, 490H Special Topics

Independent reading or research course. Full course. Does not normally count toward the major.

Fall and spring semesters. The Department.

Other years: Offered in Fall 2022, Fall 2023, Fall 2024, Spring 2025