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Biology

Professors Clotfelter‡, Goutte, Hood‡, Miller†, and Temeles; Associate Professors Purdy (Chair) and Trapani; Assistant Professors Edwards, Jeong*, Kim, Ragkousi and Wu Orr; Senior Lecturer Levin†; Visiting Assistant Professor Roche; Visiting Lecturer Heussler; Lab Coordinators Bowers 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 five categories:

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.

Five advanced courses in Biology, three of which must involve laboratories and one of which must be at the 400 level. Note that Special Topics courses 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.

Among the five advanced courses, one course in each of the two core areas of biology should be completed by the end of the junior year:

i) Cellular/Molecular Core: either Cell Structure and Function (BIOL 291) or Molecular Genetics (BIOL 370/371).
ii) Organismal Core: either Ecology (BIOL 230) or Evolutionary Biology (BIOL 320/321).

Two courses in chemistry (CHEM 151 or 155 and CHEM 161 or 165 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 151/155 and/or CHEM 161/165 are requisites for several Biology courses. Students are therefore encouraged to take CHEM 151/155 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 117/124 is recommended for BIOL 351 and BIOL 400.

One course to fulfill the STEM diversity, equity, and inclusion requirement. For the Class of 2023 and beyond, Biology Majors must take one science-oriented course that is approved by the department and that focuses on structural and systemic issues of diversity, equity, and inclusion.

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 usually 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; students placing out of BIOL 191 must take both courses from the Cellular/Molecular Core. 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). Placing out will not reduce the number of courses required for the major.

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 2022-23.†On leave fall semester 2022-23.‡On leave spring semester 2022-2023.

150, 250 Being Human in STEM

(Offered as BIOL-150, CHEM-250) This is an interactive course that combines academic inquiry and community engagement to investigate identity, inequality and representation within Science Technology Engineering and Mathematics (STEM) fields--at Amherst and beyond. We begin the course by grounding our understanding of the STEM experience at Amherst in national and global contexts. We will survey the interdisciplinary literature on the ways in which identity - race, gender, class, ability, sexuality- and geographic context shape STEM persistence and belonging. We will bring this literature into conversation with our own Amherst experiences. These challenging conversations require vulnerability, openness and the ability to tolerate discomfort. We will work from day one to build a brave space whose foundation is trust, accountability and growth. Students will design group projects that apply themes from the literature and our seminar discussions to develop resources and engage the STEM community, whether at the college, local, or national level. Course work includes critical reading and discussion, reflective writing, and collaborative work culminating in community engagement proposals which students will share with the campus and the broader public.

Open to sophomores, juniors and seniors. This course will be taught in two sections

Offered Spring Semester: Professors Jaswal and Trapani.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2020, January 2021, January 2022, Spring 2022, Spring 2024

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 Hood and Temeles; Lab Coordinator Kristensen.

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

2022-23: Offered in Fall 2022
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

181L Adaptation & Organism Lab

Lab 

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2022, Fall 2023

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 or CHEM 165. Limited to 96 students. Fall semester: Professors Goutte and Edwards, Lab Coordinator Bowers. Spring semester: Professors Purdy and Ragkousi, Lab Coordinator Bowers. 

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

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Spring 2019, Fall 2022, Fall 2023, Spring 2024

191L Molecules, Genes, & Cells Lab

Lab

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2022, Fall 2023, Spring 2024

210, 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 110 or equivalent. Limited to 40 students. Spring Semester. Professor Temeles.

2022-23: Offered in Spring 2023
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 2024

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 emphasis 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. Fall and Spring semester. Professors Kim and Roche.

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2019, Fall 2020, Fall 2021, Fall 2022, Fall 2023

221 Developmental Biology with Lab

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 2022. Professor Ragkousi.

 

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2014, Spring 2016, Fall 2022, Spring 2024

241 Genetic Analysis with lab

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 16 students. Not open to first-year students. Spring Semester. Professor Goutte.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2012, Spring 2013, Spring 2014, Spring 2015, Spring 2017, Spring 2018, Spring 2022

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. Fall 2022. Professor Purdy.

2022-23: Offered in Fall 2022
Other years: Offered in Spring 2018, Spring 2019, Fall 2022, Spring 2024

275H Evolutionary Ecology Laboratory

This laboratory explores how underlying evolutionary changes in the morphology, physiology or behavior of organisms is associated with ecological interactions within or between species.  By employing artificial selection, students will determine whether genetic variation associated with herbivore defense exists.  This experiment will be coupled with ecological investigations to explore the potential of herbivory to drive evolutionary change.  Further, using the model system Daphnia, students will have the opportunity to engage in a self-directed, in-depth investigation of selection on morphological and life history traits across varying environmental conditions.  Throughout the course, students will engage with the scientific literature, and develop their skills in identifying relevant and important questions, experimental design and hypothesis formulation, data analysis and scientific communication.  Three hours of laboratory each week.

Requisite: Prior completion of or concurrent registration in either BIOL-230 or BIOL-320. Limited to 16 students. Professors Miller and Temeles and Lab Instructor Kristensen. Spring semester.

2022-23: Offered in Spring 2023

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. Four classroom hours per week.

Requisite: BIOL 181. Limited to 20 students. Fall semester. Professor Clotfelter.

2022-23: Offered in Fall 2022
Other years: Offered in Spring 2015, Fall 2015, Fall 2016, Fall 2017, Spring 2019, Fall 2019, Fall 2022, Fall 2023

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.

In Fall 2022, the laboratory will be structured as a course-based undergraduate research experience (CURE).  The focus will be a semester-long, field-based project on bird behavior in the Wildlife Sanctuary.  Students will be outdoors in the Sanctuary under almost all weather conditions.  Students can register for either laboratory section, both of which will be held at low enrollments to facilitate field research.

Requisite: BIOL 181. Limited to 8 students per lab section. Fall semester. Professor Clotfelter.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2011, Fall 2013, Fall 2022, Fall 2023

290, 290H, 390, 490, 490H Special Topics

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

Fall and spring semesters. The Department.

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2011, Spring 2012, Fall 2012, Spring 2013, Fall 2013, Spring 2014, Fall 2014, Spring 2015, Fall 2015, Spring 2016, Fall 2016, Spring 2017, Fall 2017, Spring 2018, Fall 2018, Spring 2019, Fall 2019, Spring 2020, Fall 2020, Spring 2021, Fall 2021, Spring 2022, Fall 2022, Fall 2023, Spring 2024

291 Cell Structure and Function with Lab

Offered as BIOL 291 and BCBP 291) An analysis of the structure and function of eukaryotic cells. 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. There will be three classroom hours consisting of both lectures and problem-solving sessions, and three hours of laboratory per week. The course will be taught in-person.

Requisite: BIOL 191, CHEM151/155, and CHEM161/165. Limited to 16 students. Spring semester. Professor Edwards.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2012, Spring 2013, Spring 2014, Spring 2015, Spring 2016, Spring 2017, Spring 2018, Spring 2019, Fall 2023

301 Molecular Neurobiology with Lab

(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 and synapse development as well as neuronal 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 supplement the lectures and also emphasize current molecular techniques utilized in the study of the nervous system. Four classroom hours and three hours of laboratory per week.

While enrolled at the college, students may only take either BIOL/NEUR-301 or BIOL/NEUR-303.

Requisite: Bio 191. Bio/Neur 214 strongly suggested.

Limited to 18 students. Spring semester. Professor Roche.

2022-23: Not offered
Other years: Offered in Fall 2011, Fall 2014, Fall 2015, Fall 2016, Spring 2020, Spring 2021, Spring 2022, Fall 2023

303 Molecular Neurobiology: CURE

(Offered as BIOL 303 and NEUR 303) 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. This course is a CURE, a Course-based Undergraduate Research Experience. Students will engage in scientific practice, make novel discoveries, participate in iterative work and collaborate with peers. The goal is to give students an authentic research experience in which they will acquire necessary skills for being a scientist. Four classroom hours and three hours of laboratory per week.

Requisite: BIOL 214 or BIOL 260. Limited to 18 students. Spring semester. Professor Kim.

2022-23: Offered in Spring 2023

311 Small Proteins in Antibiotic Resistance with Lab

This lab course will provide students a hands-on mentored research experience in protein biochemistry and molecular microbiology. Through structured primary literature analysis and studying historic examples, students will learn how to develop scientific questions, structure a convincing scientific argument, design experiments to address those questions, and critically analyze data and present findings to a scientific community. In addition to participation in classroom workshops and discussions, students will work in teams to design and execute a short research project in an area of active faculty research investigating a small protein involved in antibiotic resistance in the bacterial model organism Escherichia coli. Students will present the results of their work in a short research talk and in a poster session. At the end of this course, students will be able to apply the fundamental skills used to conduct scientific research.

Professor Wu Orr

2022-23: Offered in Spring 2023

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; BIOL 191 recommended. Limited to 30 students. Not open to first-year students. Spring semester. Professor Miller

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2024

330 Biochemical Principles of Life at the Molecular Level

(Offered as CHEM 330 and BIOL 330) What are the molecular underpinnings of processes central to life? We will explore the chemical and structural properties of biological molecules and learn the logic used by the cell to build complex structures from a few basic raw materials. Some of these complex structures have evolved to catalyze chemical reactions with an enormous degree of selectivity and specificity, and we seek to discover these enzymatic strategies. We will consider the detailed balance sheet that shows how living things harvest energy from their environment to fuel metabolic processes and to reproduce and grow. Examples of the exquisite control that permits a cell to be responsive and adapt its responses based on input from the environment will be considered. We will also consider some of the means by which cells respond to change and to stress. A student may not receive credit for both CHEM/BIOL 330 and BCBP/BIOL/CHEM 331. Note: BCBP/BIOL/CHEM 331 is a requirement for the biochemistry track of the BCBP major, so prospective BCBP majors should not enroll in CHEM/BIOL 330 if they are considering the biochemistry track of the major.

Requisite: BIOL 191 and CHEM  221. Limited to 40 students with 20 students per discussion section.

Spring Semester: Professor Bishop.

2022-23: Offered in Spring 2023
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Spring 2020, Spring 2021, Spring 2022, Spring 2024

331 Biochemistry with Lab

(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 CHEM/BIOL 330 and BCBP/BIOL/CHEM 331.

BCBP/BIOL/CHEM 331 is a requirement for the biochemistry track of the BCBP major, so prospective BCBP majors should not enroll in CHEM/BIOL 330 if they are considering the biochemistry track of the major.

Requisite: CHEM 231 and BIOL 191; or consent of the instructor. (CHEM 231 may be taken either as a prerequisite or as a co-requisite.) Limited to 30 students. Fall semester. Professors Jaswal and Wu Orr. 

2022-23: Offered in Fall 2022
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

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. 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.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2012, Fall 2013, Fall 2016, Fall 2022

371 Molecular Genetics with 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 30 students. Not open to first-year students. Omitted 2022-23. Visiting Professor Heussler.

2022-23: Not offered
Other years: Offered in Fall 2017, Fall 2018, Fall 2019, Spring 2020, Fall 2020, Fall 2021, Spring 2024

391 Quantitative Imaging with Lab: From Molecules to Cells and Beyond

(Offered as BIOL 391, BCBP 391, and NEUR-391) Fluorescence imaging offers a window to understanding the structure and function of living cells at sub-cellular resolution. This laboratory and lecture course will focus on advanced quantitative fluorescence microscopy techniques used for imaging a range of biological model systems and functions ranging from single molecules to cells to tissues. Students will learn the fundamentals of optics and microscopy and gain hands-on experience with classical and emerging imaging technology including: confocal microscopy, deconvolution and super-resolution methods, and live-cell imaging. Fluorescent probes and genetically encoded biosensors will be discussed and used to study dynamic cellular processes. In addition, the class will explore a range of image processing and analysis workflows. Lectures dealing with the theory, mechanics, and application of fluorescent imaging methods will be intermingled with extended laboratory sessions in our new Amherst Biological Imaging Center.

Pre requisites : BIOL 191 and PHYS 117 and one of BIOL/BCBP291, BIOL221 or BIOL/NEUR 301, or with instructor permission.

There will be two 80-minute lectures and a weekly laboratory. Limited to 12 students. Fall semester. Professors Edwards and Kim  

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2022, Spring 2024

400 Molecular and Cellular Biophysics

(Offered as PHYS 400, BIOL 400, BCBP 400, and CHEM 400) How do the physical laws that dominate our lives change at the small length and energy scales of individual molecules? What design principles break down at the sub-cellular level and what new chemistry and physics becomes important? We will answer these questions by looking at bio-molecules, cellular substructures, and control mechanisms that work effectively in the microscopic world. How can we understand both the static and dynamic shape of proteins using the laws of thermodynamics and kinetics? How has the basic understanding of the smallest molecular motor in the world, ATP synthase, changed our understanding of friction and torque? We will explore new technologies, such as atomic force and single molecule microscopy that have allowed research into these areas. This course will address topics in each of the three major divisions of Biophysics: bio-molecular structure, biophysical techniques, and biological mechanisms.

Requisite: CHEM 161/165, PHYS 116/123, PHYS 117/124, BIOL 191 or evidence of equivalent coverage in pre-collegiate courses. Spring semester. Professor Carter. 

2022-23: Offered in Spring 2023
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Spring 2015, Spring 2016, Spring 2017, Spring 2018, Spring 2019, Spring 2020, Spring 2021, Spring 2022, Spring 2024

424 Seminar in 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. Spring semester. Prof. Miller. 

2022-23: Offered in Spring 2023
Other years: Offered in Fall 2023

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. Fall 2022. Professor Temeles.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2011, Spring 2013, Fall 2014, Spring 2016, Spring 2019, Spring 2020, Spring 2021, Fall 2022

440, 441 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 close reading and discussion of articles from the primary literature, the course will explore key topics including overexploitation (including connections between the wildlife trade and emergent diseases such as COVID-19), habitat fragmentation, climate change, restoration, protected areas, payments for ecosystem services, as well as how to determine appropriate conservation priorities. Three classroom hours per week.

Requisite: BIOL 230/ENST 210 or BIOL 320, or consent of the instructor. Not open to first-year students. Limited to 14 students. Spring Semester. Senior Lecturer Levin.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2012, Fall 2013, Fall 2015, Fall 2016, Spring 2018, Fall 2019, Fall 2020, Spring 2022

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 BIOL-214, BIOL 260, BIOL 350/351, or consent of the instructor. Limited to 18 students. Not open to first-year students. Spring Semester. Professor Trapani.

2022-23: Not offered
Other years: Offered in Spring 2013, Spring 2017, Spring 2019, Spring 2022, Spring 2024

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 or ENST 200. 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. Omitted 2022-23. Professor Clotfelter and Senior Lecturer Levin.

2022-23: Not offered
Other years: Offered in Spring 2024

498, 499, 499D Senior Departmental Honors

Honors students take three courses of thesis research, usually, but not always, with the double course load in the spring (499D). The work consists of seminar programs, individual research projects, and preparation of a thesis on the research project.

Open to seniors. Fall semester. The Department.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Fall 2019, Fall 2020, Fall 2021, Fall 2022, Fall 2023

Related Courses

AMST-296 Black Women and Reproductive Justice in the African Diaspora (Course not offered this year.)
ANTH-245 Medical Anthropology (Course not offered this year.)
ECON-212 Public Economics: Environment, Health, and Inequality (Course not offered this year.)
SOCI-328 The Pandemic (Course not offered this year.)

About Amherst College

About Amherst College

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Biology

Professors Clotfelter‡, Goutte, Hood‡, Miller†, and Temeles; Associate Professors Purdy (Chair) and Trapani; Assistant Professors Edwards, Jeong*, Kim, Ragkousi and Wu Orr; Senior Lecturer Levin†; Visiting Assistant Professor Roche; Visiting Lecturer Heussler; Lab Coordinators Bowers 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 five categories:

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.

Five advanced courses in Biology, three of which must involve laboratories and one of which must be at the 400 level. Note that Special Topics courses 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.

Among the five advanced courses, one course in each of the two core areas of biology should be completed by the end of the junior year:

i) Cellular/Molecular Core: either Cell Structure and Function (BIOL 291) or Molecular Genetics (BIOL 370/371).
ii) Organismal Core: either Ecology (BIOL 230) or Evolutionary Biology (BIOL 320/321).

Two courses in chemistry (CHEM 151 or 155 and CHEM 161 or 165 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 151/155 and/or CHEM 161/165 are requisites for several Biology courses. Students are therefore encouraged to take CHEM 151/155 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 117/124 is recommended for BIOL 351 and BIOL 400.

One course to fulfill the STEM diversity, equity, and inclusion requirement. For the Class of 2023 and beyond, Biology Majors must take one science-oriented course that is approved by the department and that focuses on structural and systemic issues of diversity, equity, and inclusion.

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 usually 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; students placing out of BIOL 191 must take both courses from the Cellular/Molecular Core. 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). Placing out will not reduce the number of courses required for the major.

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 2022-23.†On leave fall semester 2022-23.‡On leave spring semester 2022-2023.

150, 250 Being Human in STEM

(Offered as BIOL-150, CHEM-250) This is an interactive course that combines academic inquiry and community engagement to investigate identity, inequality and representation within Science Technology Engineering and Mathematics (STEM) fields--at Amherst and beyond. We begin the course by grounding our understanding of the STEM experience at Amherst in national and global contexts. We will survey the interdisciplinary literature on the ways in which identity - race, gender, class, ability, sexuality- and geographic context shape STEM persistence and belonging. We will bring this literature into conversation with our own Amherst experiences. These challenging conversations require vulnerability, openness and the ability to tolerate discomfort. We will work from day one to build a brave space whose foundation is trust, accountability and growth. Students will design group projects that apply themes from the literature and our seminar discussions to develop resources and engage the STEM community, whether at the college, local, or national level. Course work includes critical reading and discussion, reflective writing, and collaborative work culminating in community engagement proposals which students will share with the campus and the broader public.

Open to sophomores, juniors and seniors. This course will be taught in two sections

Offered Spring Semester: Professors Jaswal and Trapani.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2020, January 2021, January 2022, Spring 2022, Spring 2024

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 Hood and Temeles; Lab Coordinator Kristensen.

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

2022-23: Offered in Fall 2022
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

181L Adaptation & Organism Lab

Lab 

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2022, Fall 2023

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 or CHEM 165. Limited to 96 students. Fall semester: Professors Goutte and Edwards, Lab Coordinator Bowers. Spring semester: Professors Purdy and Ragkousi, Lab Coordinator Bowers. 

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

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Spring 2019, Fall 2022, Fall 2023, Spring 2024

191L Molecules, Genes, & Cells Lab

Lab

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2022, Fall 2023, Spring 2024

210, 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 110 or equivalent. Limited to 40 students. Spring Semester. Professor Temeles.

2022-23: Offered in Spring 2023
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 2024

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 emphasis 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. Fall and Spring semester. Professors Kim and Roche.

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2019, Fall 2020, Fall 2021, Fall 2022, Fall 2023

221 Developmental Biology with Lab

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 2022. Professor Ragkousi.

 

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2014, Spring 2016, Fall 2022, Spring 2024

241 Genetic Analysis with lab

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 16 students. Not open to first-year students. Spring Semester. Professor Goutte.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2012, Spring 2013, Spring 2014, Spring 2015, Spring 2017, Spring 2018, Spring 2022

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. Fall 2022. Professor Purdy.

2022-23: Offered in Fall 2022
Other years: Offered in Spring 2018, Spring 2019, Fall 2022, Spring 2024

275H Evolutionary Ecology Laboratory

This laboratory explores how underlying evolutionary changes in the morphology, physiology or behavior of organisms is associated with ecological interactions within or between species.  By employing artificial selection, students will determine whether genetic variation associated with herbivore defense exists.  This experiment will be coupled with ecological investigations to explore the potential of herbivory to drive evolutionary change.  Further, using the model system Daphnia, students will have the opportunity to engage in a self-directed, in-depth investigation of selection on morphological and life history traits across varying environmental conditions.  Throughout the course, students will engage with the scientific literature, and develop their skills in identifying relevant and important questions, experimental design and hypothesis formulation, data analysis and scientific communication.  Three hours of laboratory each week.

Requisite: Prior completion of or concurrent registration in either BIOL-230 or BIOL-320. Limited to 16 students. Professors Miller and Temeles and Lab Instructor Kristensen. Spring semester.

2022-23: Offered in Spring 2023

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. Four classroom hours per week.

Requisite: BIOL 181. Limited to 20 students. Fall semester. Professor Clotfelter.

2022-23: Offered in Fall 2022
Other years: Offered in Spring 2015, Fall 2015, Fall 2016, Fall 2017, Spring 2019, Fall 2019, Fall 2022, Fall 2023

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.

In Fall 2022, the laboratory will be structured as a course-based undergraduate research experience (CURE).  The focus will be a semester-long, field-based project on bird behavior in the Wildlife Sanctuary.  Students will be outdoors in the Sanctuary under almost all weather conditions.  Students can register for either laboratory section, both of which will be held at low enrollments to facilitate field research.

Requisite: BIOL 181. Limited to 8 students per lab section. Fall semester. Professor Clotfelter.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2011, Fall 2013, Fall 2022, Fall 2023

290, 290H, 390, 490, 490H Special Topics

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

Fall and spring semesters. The Department.

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2011, Spring 2012, Fall 2012, Spring 2013, Fall 2013, Spring 2014, Fall 2014, Spring 2015, Fall 2015, Spring 2016, Fall 2016, Spring 2017, Fall 2017, Spring 2018, Fall 2018, Spring 2019, Fall 2019, Spring 2020, Fall 2020, Spring 2021, Fall 2021, Spring 2022, Fall 2022, Fall 2023, Spring 2024

291 Cell Structure and Function with Lab

Offered as BIOL 291 and BCBP 291) An analysis of the structure and function of eukaryotic cells. 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. There will be three classroom hours consisting of both lectures and problem-solving sessions, and three hours of laboratory per week. The course will be taught in-person.

Requisite: BIOL 191, CHEM151/155, and CHEM161/165. Limited to 16 students. Spring semester. Professor Edwards.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2012, Spring 2013, Spring 2014, Spring 2015, Spring 2016, Spring 2017, Spring 2018, Spring 2019, Fall 2023

301 Molecular Neurobiology with Lab

(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 and synapse development as well as neuronal 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 supplement the lectures and also emphasize current molecular techniques utilized in the study of the nervous system. Four classroom hours and three hours of laboratory per week.

While enrolled at the college, students may only take either BIOL/NEUR-301 or BIOL/NEUR-303.

Requisite: Bio 191. Bio/Neur 214 strongly suggested.

Limited to 18 students. Spring semester. Professor Roche.

2022-23: Not offered
Other years: Offered in Fall 2011, Fall 2014, Fall 2015, Fall 2016, Spring 2020, Spring 2021, Spring 2022, Fall 2023

303 Molecular Neurobiology: CURE

(Offered as BIOL 303 and NEUR 303) 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. This course is a CURE, a Course-based Undergraduate Research Experience. Students will engage in scientific practice, make novel discoveries, participate in iterative work and collaborate with peers. The goal is to give students an authentic research experience in which they will acquire necessary skills for being a scientist. Four classroom hours and three hours of laboratory per week.

Requisite: BIOL 214 or BIOL 260. Limited to 18 students. Spring semester. Professor Kim.

2022-23: Offered in Spring 2023

311 Small Proteins in Antibiotic Resistance with Lab

This lab course will provide students a hands-on mentored research experience in protein biochemistry and molecular microbiology. Through structured primary literature analysis and studying historic examples, students will learn how to develop scientific questions, structure a convincing scientific argument, design experiments to address those questions, and critically analyze data and present findings to a scientific community. In addition to participation in classroom workshops and discussions, students will work in teams to design and execute a short research project in an area of active faculty research investigating a small protein involved in antibiotic resistance in the bacterial model organism Escherichia coli. Students will present the results of their work in a short research talk and in a poster session. At the end of this course, students will be able to apply the fundamental skills used to conduct scientific research.

Professor Wu Orr

2022-23: Offered in Spring 2023

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; BIOL 191 recommended. Limited to 30 students. Not open to first-year students. Spring semester. Professor Miller

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2024

330 Biochemical Principles of Life at the Molecular Level

(Offered as CHEM 330 and BIOL 330) What are the molecular underpinnings of processes central to life? We will explore the chemical and structural properties of biological molecules and learn the logic used by the cell to build complex structures from a few basic raw materials. Some of these complex structures have evolved to catalyze chemical reactions with an enormous degree of selectivity and specificity, and we seek to discover these enzymatic strategies. We will consider the detailed balance sheet that shows how living things harvest energy from their environment to fuel metabolic processes and to reproduce and grow. Examples of the exquisite control that permits a cell to be responsive and adapt its responses based on input from the environment will be considered. We will also consider some of the means by which cells respond to change and to stress. A student may not receive credit for both CHEM/BIOL 330 and BCBP/BIOL/CHEM 331. Note: BCBP/BIOL/CHEM 331 is a requirement for the biochemistry track of the BCBP major, so prospective BCBP majors should not enroll in CHEM/BIOL 330 if they are considering the biochemistry track of the major.

Requisite: BIOL 191 and CHEM  221. Limited to 40 students with 20 students per discussion section.

Spring Semester: Professor Bishop.

2022-23: Offered in Spring 2023
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Spring 2020, Spring 2021, Spring 2022, Spring 2024

331 Biochemistry with Lab

(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 CHEM/BIOL 330 and BCBP/BIOL/CHEM 331.

BCBP/BIOL/CHEM 331 is a requirement for the biochemistry track of the BCBP major, so prospective BCBP majors should not enroll in CHEM/BIOL 330 if they are considering the biochemistry track of the major.

Requisite: CHEM 231 and BIOL 191; or consent of the instructor. (CHEM 231 may be taken either as a prerequisite or as a co-requisite.) Limited to 30 students. Fall semester. Professors Jaswal and Wu Orr. 

2022-23: Offered in Fall 2022
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

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. 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.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2012, Fall 2013, Fall 2016, Fall 2022

371 Molecular Genetics with 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 30 students. Not open to first-year students. Omitted 2022-23. Visiting Professor Heussler.

2022-23: Not offered
Other years: Offered in Fall 2017, Fall 2018, Fall 2019, Spring 2020, Fall 2020, Fall 2021, Spring 2024

391 Quantitative Imaging with Lab: From Molecules to Cells and Beyond

(Offered as BIOL 391, BCBP 391, and NEUR-391) Fluorescence imaging offers a window to understanding the structure and function of living cells at sub-cellular resolution. This laboratory and lecture course will focus on advanced quantitative fluorescence microscopy techniques used for imaging a range of biological model systems and functions ranging from single molecules to cells to tissues. Students will learn the fundamentals of optics and microscopy and gain hands-on experience with classical and emerging imaging technology including: confocal microscopy, deconvolution and super-resolution methods, and live-cell imaging. Fluorescent probes and genetically encoded biosensors will be discussed and used to study dynamic cellular processes. In addition, the class will explore a range of image processing and analysis workflows. Lectures dealing with the theory, mechanics, and application of fluorescent imaging methods will be intermingled with extended laboratory sessions in our new Amherst Biological Imaging Center.

Pre requisites : BIOL 191 and PHYS 117 and one of BIOL/BCBP291, BIOL221 or BIOL/NEUR 301, or with instructor permission.

There will be two 80-minute lectures and a weekly laboratory. Limited to 12 students. Fall semester. Professors Edwards and Kim  

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2022, Spring 2024

400 Molecular and Cellular Biophysics

(Offered as PHYS 400, BIOL 400, BCBP 400, and CHEM 400) How do the physical laws that dominate our lives change at the small length and energy scales of individual molecules? What design principles break down at the sub-cellular level and what new chemistry and physics becomes important? We will answer these questions by looking at bio-molecules, cellular substructures, and control mechanisms that work effectively in the microscopic world. How can we understand both the static and dynamic shape of proteins using the laws of thermodynamics and kinetics? How has the basic understanding of the smallest molecular motor in the world, ATP synthase, changed our understanding of friction and torque? We will explore new technologies, such as atomic force and single molecule microscopy that have allowed research into these areas. This course will address topics in each of the three major divisions of Biophysics: bio-molecular structure, biophysical techniques, and biological mechanisms.

Requisite: CHEM 161/165, PHYS 116/123, PHYS 117/124, BIOL 191 or evidence of equivalent coverage in pre-collegiate courses. Spring semester. Professor Carter. 

2022-23: Offered in Spring 2023
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Spring 2015, Spring 2016, Spring 2017, Spring 2018, Spring 2019, Spring 2020, Spring 2021, Spring 2022, Spring 2024

424 Seminar in 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. Spring semester. Prof. Miller. 

2022-23: Offered in Spring 2023
Other years: Offered in Fall 2023

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. Fall 2022. Professor Temeles.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2011, Spring 2013, Fall 2014, Spring 2016, Spring 2019, Spring 2020, Spring 2021, Fall 2022

440, 441 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 close reading and discussion of articles from the primary literature, the course will explore key topics including overexploitation (including connections between the wildlife trade and emergent diseases such as COVID-19), habitat fragmentation, climate change, restoration, protected areas, payments for ecosystem services, as well as how to determine appropriate conservation priorities. Three classroom hours per week.

Requisite: BIOL 230/ENST 210 or BIOL 320, or consent of the instructor. Not open to first-year students. Limited to 14 students. Spring Semester. Senior Lecturer Levin.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2012, Fall 2013, Fall 2015, Fall 2016, Spring 2018, Fall 2019, Fall 2020, Spring 2022

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 BIOL-214, BIOL 260, BIOL 350/351, or consent of the instructor. Limited to 18 students. Not open to first-year students. Spring Semester. Professor Trapani.

2022-23: Not offered
Other years: Offered in Spring 2013, Spring 2017, Spring 2019, Spring 2022, Spring 2024

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 or ENST 200. 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. Omitted 2022-23. Professor Clotfelter and Senior Lecturer Levin.

2022-23: Not offered
Other years: Offered in Spring 2024

498, 499, 499D Senior Departmental Honors

Honors students take three courses of thesis research, usually, but not always, with the double course load in the spring (499D). The work consists of seminar programs, individual research projects, and preparation of a thesis on the research project.

Open to seniors. Fall semester. The Department.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Fall 2019, Fall 2020, Fall 2021, Fall 2022, Fall 2023

Related Courses

AMST-296 Black Women and Reproductive Justice in the African Diaspora (Course not offered this year.)
ANTH-245 Medical Anthropology (Course not offered this year.)
ECON-212 Public Economics: Environment, Health, and Inequality (Course not offered this year.)
SOCI-328 The Pandemic (Course not offered this year.)

Admission & Financial Aid

Admission & Financial Aid

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Biology

Professors Clotfelter‡, Goutte, Hood‡, Miller†, and Temeles; Associate Professors Purdy (Chair) and Trapani; Assistant Professors Edwards, Jeong*, Kim, Ragkousi and Wu Orr; Senior Lecturer Levin†; Visiting Assistant Professor Roche; Visiting Lecturer Heussler; Lab Coordinators Bowers 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 five categories:

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.

Five advanced courses in Biology, three of which must involve laboratories and one of which must be at the 400 level. Note that Special Topics courses 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.

Among the five advanced courses, one course in each of the two core areas of biology should be completed by the end of the junior year:

i) Cellular/Molecular Core: either Cell Structure and Function (BIOL 291) or Molecular Genetics (BIOL 370/371).
ii) Organismal Core: either Ecology (BIOL 230) or Evolutionary Biology (BIOL 320/321).

Two courses in chemistry (CHEM 151 or 155 and CHEM 161 or 165 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 151/155 and/or CHEM 161/165 are requisites for several Biology courses. Students are therefore encouraged to take CHEM 151/155 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 117/124 is recommended for BIOL 351 and BIOL 400.

One course to fulfill the STEM diversity, equity, and inclusion requirement. For the Class of 2023 and beyond, Biology Majors must take one science-oriented course that is approved by the department and that focuses on structural and systemic issues of diversity, equity, and inclusion.

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 usually 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; students placing out of BIOL 191 must take both courses from the Cellular/Molecular Core. 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). Placing out will not reduce the number of courses required for the major.

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 2022-23.†On leave fall semester 2022-23.‡On leave spring semester 2022-2023.

150, 250 Being Human in STEM

(Offered as BIOL-150, CHEM-250) This is an interactive course that combines academic inquiry and community engagement to investigate identity, inequality and representation within Science Technology Engineering and Mathematics (STEM) fields--at Amherst and beyond. We begin the course by grounding our understanding of the STEM experience at Amherst in national and global contexts. We will survey the interdisciplinary literature on the ways in which identity - race, gender, class, ability, sexuality- and geographic context shape STEM persistence and belonging. We will bring this literature into conversation with our own Amherst experiences. These challenging conversations require vulnerability, openness and the ability to tolerate discomfort. We will work from day one to build a brave space whose foundation is trust, accountability and growth. Students will design group projects that apply themes from the literature and our seminar discussions to develop resources and engage the STEM community, whether at the college, local, or national level. Course work includes critical reading and discussion, reflective writing, and collaborative work culminating in community engagement proposals which students will share with the campus and the broader public.

Open to sophomores, juniors and seniors. This course will be taught in two sections

Offered Spring Semester: Professors Jaswal and Trapani.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2020, January 2021, January 2022, Spring 2022, Spring 2024

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 Hood and Temeles; Lab Coordinator Kristensen.

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

2022-23: Offered in Fall 2022
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

181L Adaptation & Organism Lab

Lab 

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2022, Fall 2023

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 or CHEM 165. Limited to 96 students. Fall semester: Professors Goutte and Edwards, Lab Coordinator Bowers. Spring semester: Professors Purdy and Ragkousi, Lab Coordinator Bowers. 

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

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Spring 2019, Fall 2022, Fall 2023, Spring 2024

191L Molecules, Genes, & Cells Lab

Lab

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2022, Fall 2023, Spring 2024

210, 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 110 or equivalent. Limited to 40 students. Spring Semester. Professor Temeles.

2022-23: Offered in Spring 2023
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 2024

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 emphasis 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. Fall and Spring semester. Professors Kim and Roche.

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2019, Fall 2020, Fall 2021, Fall 2022, Fall 2023

221 Developmental Biology with Lab

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 2022. Professor Ragkousi.

 

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2014, Spring 2016, Fall 2022, Spring 2024

241 Genetic Analysis with lab

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 16 students. Not open to first-year students. Spring Semester. Professor Goutte.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2012, Spring 2013, Spring 2014, Spring 2015, Spring 2017, Spring 2018, Spring 2022

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. Fall 2022. Professor Purdy.

2022-23: Offered in Fall 2022
Other years: Offered in Spring 2018, Spring 2019, Fall 2022, Spring 2024

275H Evolutionary Ecology Laboratory

This laboratory explores how underlying evolutionary changes in the morphology, physiology or behavior of organisms is associated with ecological interactions within or between species.  By employing artificial selection, students will determine whether genetic variation associated with herbivore defense exists.  This experiment will be coupled with ecological investigations to explore the potential of herbivory to drive evolutionary change.  Further, using the model system Daphnia, students will have the opportunity to engage in a self-directed, in-depth investigation of selection on morphological and life history traits across varying environmental conditions.  Throughout the course, students will engage with the scientific literature, and develop their skills in identifying relevant and important questions, experimental design and hypothesis formulation, data analysis and scientific communication.  Three hours of laboratory each week.

Requisite: Prior completion of or concurrent registration in either BIOL-230 or BIOL-320. Limited to 16 students. Professors Miller and Temeles and Lab Instructor Kristensen. Spring semester.

2022-23: Offered in Spring 2023

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. Four classroom hours per week.

Requisite: BIOL 181. Limited to 20 students. Fall semester. Professor Clotfelter.

2022-23: Offered in Fall 2022
Other years: Offered in Spring 2015, Fall 2015, Fall 2016, Fall 2017, Spring 2019, Fall 2019, Fall 2022, Fall 2023

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.

In Fall 2022, the laboratory will be structured as a course-based undergraduate research experience (CURE).  The focus will be a semester-long, field-based project on bird behavior in the Wildlife Sanctuary.  Students will be outdoors in the Sanctuary under almost all weather conditions.  Students can register for either laboratory section, both of which will be held at low enrollments to facilitate field research.

Requisite: BIOL 181. Limited to 8 students per lab section. Fall semester. Professor Clotfelter.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2011, Fall 2013, Fall 2022, Fall 2023

290, 290H, 390, 490, 490H Special Topics

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

Fall and spring semesters. The Department.

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2011, Spring 2012, Fall 2012, Spring 2013, Fall 2013, Spring 2014, Fall 2014, Spring 2015, Fall 2015, Spring 2016, Fall 2016, Spring 2017, Fall 2017, Spring 2018, Fall 2018, Spring 2019, Fall 2019, Spring 2020, Fall 2020, Spring 2021, Fall 2021, Spring 2022, Fall 2022, Fall 2023, Spring 2024

291 Cell Structure and Function with Lab

Offered as BIOL 291 and BCBP 291) An analysis of the structure and function of eukaryotic cells. 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. There will be three classroom hours consisting of both lectures and problem-solving sessions, and three hours of laboratory per week. The course will be taught in-person.

Requisite: BIOL 191, CHEM151/155, and CHEM161/165. Limited to 16 students. Spring semester. Professor Edwards.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2012, Spring 2013, Spring 2014, Spring 2015, Spring 2016, Spring 2017, Spring 2018, Spring 2019, Fall 2023

301 Molecular Neurobiology with Lab

(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 and synapse development as well as neuronal 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 supplement the lectures and also emphasize current molecular techniques utilized in the study of the nervous system. Four classroom hours and three hours of laboratory per week.

While enrolled at the college, students may only take either BIOL/NEUR-301 or BIOL/NEUR-303.

Requisite: Bio 191. Bio/Neur 214 strongly suggested.

Limited to 18 students. Spring semester. Professor Roche.

2022-23: Not offered
Other years: Offered in Fall 2011, Fall 2014, Fall 2015, Fall 2016, Spring 2020, Spring 2021, Spring 2022, Fall 2023

303 Molecular Neurobiology: CURE

(Offered as BIOL 303 and NEUR 303) 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. This course is a CURE, a Course-based Undergraduate Research Experience. Students will engage in scientific practice, make novel discoveries, participate in iterative work and collaborate with peers. The goal is to give students an authentic research experience in which they will acquire necessary skills for being a scientist. Four classroom hours and three hours of laboratory per week.

Requisite: BIOL 214 or BIOL 260. Limited to 18 students. Spring semester. Professor Kim.

2022-23: Offered in Spring 2023

311 Small Proteins in Antibiotic Resistance with Lab

This lab course will provide students a hands-on mentored research experience in protein biochemistry and molecular microbiology. Through structured primary literature analysis and studying historic examples, students will learn how to develop scientific questions, structure a convincing scientific argument, design experiments to address those questions, and critically analyze data and present findings to a scientific community. In addition to participation in classroom workshops and discussions, students will work in teams to design and execute a short research project in an area of active faculty research investigating a small protein involved in antibiotic resistance in the bacterial model organism Escherichia coli. Students will present the results of their work in a short research talk and in a poster session. At the end of this course, students will be able to apply the fundamental skills used to conduct scientific research.

Professor Wu Orr

2022-23: Offered in Spring 2023

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; BIOL 191 recommended. Limited to 30 students. Not open to first-year students. Spring semester. Professor Miller

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2024

330 Biochemical Principles of Life at the Molecular Level

(Offered as CHEM 330 and BIOL 330) What are the molecular underpinnings of processes central to life? We will explore the chemical and structural properties of biological molecules and learn the logic used by the cell to build complex structures from a few basic raw materials. Some of these complex structures have evolved to catalyze chemical reactions with an enormous degree of selectivity and specificity, and we seek to discover these enzymatic strategies. We will consider the detailed balance sheet that shows how living things harvest energy from their environment to fuel metabolic processes and to reproduce and grow. Examples of the exquisite control that permits a cell to be responsive and adapt its responses based on input from the environment will be considered. We will also consider some of the means by which cells respond to change and to stress. A student may not receive credit for both CHEM/BIOL 330 and BCBP/BIOL/CHEM 331. Note: BCBP/BIOL/CHEM 331 is a requirement for the biochemistry track of the BCBP major, so prospective BCBP majors should not enroll in CHEM/BIOL 330 if they are considering the biochemistry track of the major.

Requisite: BIOL 191 and CHEM  221. Limited to 40 students with 20 students per discussion section.

Spring Semester: Professor Bishop.

2022-23: Offered in Spring 2023
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Spring 2020, Spring 2021, Spring 2022, Spring 2024

331 Biochemistry with Lab

(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 CHEM/BIOL 330 and BCBP/BIOL/CHEM 331.

BCBP/BIOL/CHEM 331 is a requirement for the biochemistry track of the BCBP major, so prospective BCBP majors should not enroll in CHEM/BIOL 330 if they are considering the biochemistry track of the major.

Requisite: CHEM 231 and BIOL 191; or consent of the instructor. (CHEM 231 may be taken either as a prerequisite or as a co-requisite.) Limited to 30 students. Fall semester. Professors Jaswal and Wu Orr. 

2022-23: Offered in Fall 2022
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

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. 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.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2012, Fall 2013, Fall 2016, Fall 2022

371 Molecular Genetics with 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 30 students. Not open to first-year students. Omitted 2022-23. Visiting Professor Heussler.

2022-23: Not offered
Other years: Offered in Fall 2017, Fall 2018, Fall 2019, Spring 2020, Fall 2020, Fall 2021, Spring 2024

391 Quantitative Imaging with Lab: From Molecules to Cells and Beyond

(Offered as BIOL 391, BCBP 391, and NEUR-391) Fluorescence imaging offers a window to understanding the structure and function of living cells at sub-cellular resolution. This laboratory and lecture course will focus on advanced quantitative fluorescence microscopy techniques used for imaging a range of biological model systems and functions ranging from single molecules to cells to tissues. Students will learn the fundamentals of optics and microscopy and gain hands-on experience with classical and emerging imaging technology including: confocal microscopy, deconvolution and super-resolution methods, and live-cell imaging. Fluorescent probes and genetically encoded biosensors will be discussed and used to study dynamic cellular processes. In addition, the class will explore a range of image processing and analysis workflows. Lectures dealing with the theory, mechanics, and application of fluorescent imaging methods will be intermingled with extended laboratory sessions in our new Amherst Biological Imaging Center.

Pre requisites : BIOL 191 and PHYS 117 and one of BIOL/BCBP291, BIOL221 or BIOL/NEUR 301, or with instructor permission.

There will be two 80-minute lectures and a weekly laboratory. Limited to 12 students. Fall semester. Professors Edwards and Kim  

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2022, Spring 2024

400 Molecular and Cellular Biophysics

(Offered as PHYS 400, BIOL 400, BCBP 400, and CHEM 400) How do the physical laws that dominate our lives change at the small length and energy scales of individual molecules? What design principles break down at the sub-cellular level and what new chemistry and physics becomes important? We will answer these questions by looking at bio-molecules, cellular substructures, and control mechanisms that work effectively in the microscopic world. How can we understand both the static and dynamic shape of proteins using the laws of thermodynamics and kinetics? How has the basic understanding of the smallest molecular motor in the world, ATP synthase, changed our understanding of friction and torque? We will explore new technologies, such as atomic force and single molecule microscopy that have allowed research into these areas. This course will address topics in each of the three major divisions of Biophysics: bio-molecular structure, biophysical techniques, and biological mechanisms.

Requisite: CHEM 161/165, PHYS 116/123, PHYS 117/124, BIOL 191 or evidence of equivalent coverage in pre-collegiate courses. Spring semester. Professor Carter. 

2022-23: Offered in Spring 2023
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Spring 2015, Spring 2016, Spring 2017, Spring 2018, Spring 2019, Spring 2020, Spring 2021, Spring 2022, Spring 2024

424 Seminar in 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. Spring semester. Prof. Miller. 

2022-23: Offered in Spring 2023
Other years: Offered in Fall 2023

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. Fall 2022. Professor Temeles.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2011, Spring 2013, Fall 2014, Spring 2016, Spring 2019, Spring 2020, Spring 2021, Fall 2022

440, 441 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 close reading and discussion of articles from the primary literature, the course will explore key topics including overexploitation (including connections between the wildlife trade and emergent diseases such as COVID-19), habitat fragmentation, climate change, restoration, protected areas, payments for ecosystem services, as well as how to determine appropriate conservation priorities. Three classroom hours per week.

Requisite: BIOL 230/ENST 210 or BIOL 320, or consent of the instructor. Not open to first-year students. Limited to 14 students. Spring Semester. Senior Lecturer Levin.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2012, Fall 2013, Fall 2015, Fall 2016, Spring 2018, Fall 2019, Fall 2020, Spring 2022

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 BIOL-214, BIOL 260, BIOL 350/351, or consent of the instructor. Limited to 18 students. Not open to first-year students. Spring Semester. Professor Trapani.

2022-23: Not offered
Other years: Offered in Spring 2013, Spring 2017, Spring 2019, Spring 2022, Spring 2024

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 or ENST 200. 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. Omitted 2022-23. Professor Clotfelter and Senior Lecturer Levin.

2022-23: Not offered
Other years: Offered in Spring 2024

498, 499, 499D Senior Departmental Honors

Honors students take three courses of thesis research, usually, but not always, with the double course load in the spring (499D). The work consists of seminar programs, individual research projects, and preparation of a thesis on the research project.

Open to seniors. Fall semester. The Department.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Fall 2019, Fall 2020, Fall 2021, Fall 2022, Fall 2023

Related Courses

AMST-296 Black Women and Reproductive Justice in the African Diaspora (Course not offered this year.)
ANTH-245 Medical Anthropology (Course not offered this year.)
ECON-212 Public Economics: Environment, Health, and Inequality (Course not offered this year.)
SOCI-328 The Pandemic (Course not offered this year.)

Regulations & Requirements

Regulations & Requirements

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Biology

Professors Clotfelter‡, Goutte, Hood‡, Miller†, and Temeles; Associate Professors Purdy (Chair) and Trapani; Assistant Professors Edwards, Jeong*, Kim, Ragkousi and Wu Orr; Senior Lecturer Levin†; Visiting Assistant Professor Roche; Visiting Lecturer Heussler; Lab Coordinators Bowers 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 five categories:

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.

Five advanced courses in Biology, three of which must involve laboratories and one of which must be at the 400 level. Note that Special Topics courses 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.

Among the five advanced courses, one course in each of the two core areas of biology should be completed by the end of the junior year:

i) Cellular/Molecular Core: either Cell Structure and Function (BIOL 291) or Molecular Genetics (BIOL 370/371).
ii) Organismal Core: either Ecology (BIOL 230) or Evolutionary Biology (BIOL 320/321).

Two courses in chemistry (CHEM 151 or 155 and CHEM 161 or 165 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 151/155 and/or CHEM 161/165 are requisites for several Biology courses. Students are therefore encouraged to take CHEM 151/155 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 117/124 is recommended for BIOL 351 and BIOL 400.

One course to fulfill the STEM diversity, equity, and inclusion requirement. For the Class of 2023 and beyond, Biology Majors must take one science-oriented course that is approved by the department and that focuses on structural and systemic issues of diversity, equity, and inclusion.

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 usually 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; students placing out of BIOL 191 must take both courses from the Cellular/Molecular Core. 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). Placing out will not reduce the number of courses required for the major.

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 2022-23.†On leave fall semester 2022-23.‡On leave spring semester 2022-2023.

150, 250 Being Human in STEM

(Offered as BIOL-150, CHEM-250) This is an interactive course that combines academic inquiry and community engagement to investigate identity, inequality and representation within Science Technology Engineering and Mathematics (STEM) fields--at Amherst and beyond. We begin the course by grounding our understanding of the STEM experience at Amherst in national and global contexts. We will survey the interdisciplinary literature on the ways in which identity - race, gender, class, ability, sexuality- and geographic context shape STEM persistence and belonging. We will bring this literature into conversation with our own Amherst experiences. These challenging conversations require vulnerability, openness and the ability to tolerate discomfort. We will work from day one to build a brave space whose foundation is trust, accountability and growth. Students will design group projects that apply themes from the literature and our seminar discussions to develop resources and engage the STEM community, whether at the college, local, or national level. Course work includes critical reading and discussion, reflective writing, and collaborative work culminating in community engagement proposals which students will share with the campus and the broader public.

Open to sophomores, juniors and seniors. This course will be taught in two sections

Offered Spring Semester: Professors Jaswal and Trapani.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2020, January 2021, January 2022, Spring 2022, Spring 2024

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 Hood and Temeles; Lab Coordinator Kristensen.

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

2022-23: Offered in Fall 2022
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

181L Adaptation & Organism Lab

Lab 

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2022, Fall 2023

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 or CHEM 165. Limited to 96 students. Fall semester: Professors Goutte and Edwards, Lab Coordinator Bowers. Spring semester: Professors Purdy and Ragkousi, Lab Coordinator Bowers. 

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

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Spring 2019, Fall 2022, Fall 2023, Spring 2024

191L Molecules, Genes, & Cells Lab

Lab

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2022, Fall 2023, Spring 2024

210, 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 110 or equivalent. Limited to 40 students. Spring Semester. Professor Temeles.

2022-23: Offered in Spring 2023
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 2024

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 emphasis 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. Fall and Spring semester. Professors Kim and Roche.

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2019, Fall 2020, Fall 2021, Fall 2022, Fall 2023

221 Developmental Biology with Lab

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 2022. Professor Ragkousi.

 

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2014, Spring 2016, Fall 2022, Spring 2024

241 Genetic Analysis with lab

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 16 students. Not open to first-year students. Spring Semester. Professor Goutte.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2012, Spring 2013, Spring 2014, Spring 2015, Spring 2017, Spring 2018, Spring 2022

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. Fall 2022. Professor Purdy.

2022-23: Offered in Fall 2022
Other years: Offered in Spring 2018, Spring 2019, Fall 2022, Spring 2024

275H Evolutionary Ecology Laboratory

This laboratory explores how underlying evolutionary changes in the morphology, physiology or behavior of organisms is associated with ecological interactions within or between species.  By employing artificial selection, students will determine whether genetic variation associated with herbivore defense exists.  This experiment will be coupled with ecological investigations to explore the potential of herbivory to drive evolutionary change.  Further, using the model system Daphnia, students will have the opportunity to engage in a self-directed, in-depth investigation of selection on morphological and life history traits across varying environmental conditions.  Throughout the course, students will engage with the scientific literature, and develop their skills in identifying relevant and important questions, experimental design and hypothesis formulation, data analysis and scientific communication.  Three hours of laboratory each week.

Requisite: Prior completion of or concurrent registration in either BIOL-230 or BIOL-320. Limited to 16 students. Professors Miller and Temeles and Lab Instructor Kristensen. Spring semester.

2022-23: Offered in Spring 2023

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. Four classroom hours per week.

Requisite: BIOL 181. Limited to 20 students. Fall semester. Professor Clotfelter.

2022-23: Offered in Fall 2022
Other years: Offered in Spring 2015, Fall 2015, Fall 2016, Fall 2017, Spring 2019, Fall 2019, Fall 2022, Fall 2023

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.

In Fall 2022, the laboratory will be structured as a course-based undergraduate research experience (CURE).  The focus will be a semester-long, field-based project on bird behavior in the Wildlife Sanctuary.  Students will be outdoors in the Sanctuary under almost all weather conditions.  Students can register for either laboratory section, both of which will be held at low enrollments to facilitate field research.

Requisite: BIOL 181. Limited to 8 students per lab section. Fall semester. Professor Clotfelter.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2011, Fall 2013, Fall 2022, Fall 2023

290, 290H, 390, 490, 490H Special Topics

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

Fall and spring semesters. The Department.

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2011, Spring 2012, Fall 2012, Spring 2013, Fall 2013, Spring 2014, Fall 2014, Spring 2015, Fall 2015, Spring 2016, Fall 2016, Spring 2017, Fall 2017, Spring 2018, Fall 2018, Spring 2019, Fall 2019, Spring 2020, Fall 2020, Spring 2021, Fall 2021, Spring 2022, Fall 2022, Fall 2023, Spring 2024

291 Cell Structure and Function with Lab

Offered as BIOL 291 and BCBP 291) An analysis of the structure and function of eukaryotic cells. 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. There will be three classroom hours consisting of both lectures and problem-solving sessions, and three hours of laboratory per week. The course will be taught in-person.

Requisite: BIOL 191, CHEM151/155, and CHEM161/165. Limited to 16 students. Spring semester. Professor Edwards.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2012, Spring 2013, Spring 2014, Spring 2015, Spring 2016, Spring 2017, Spring 2018, Spring 2019, Fall 2023

301 Molecular Neurobiology with Lab

(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 and synapse development as well as neuronal 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 supplement the lectures and also emphasize current molecular techniques utilized in the study of the nervous system. Four classroom hours and three hours of laboratory per week.

While enrolled at the college, students may only take either BIOL/NEUR-301 or BIOL/NEUR-303.

Requisite: Bio 191. Bio/Neur 214 strongly suggested.

Limited to 18 students. Spring semester. Professor Roche.

2022-23: Not offered
Other years: Offered in Fall 2011, Fall 2014, Fall 2015, Fall 2016, Spring 2020, Spring 2021, Spring 2022, Fall 2023

303 Molecular Neurobiology: CURE

(Offered as BIOL 303 and NEUR 303) 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. This course is a CURE, a Course-based Undergraduate Research Experience. Students will engage in scientific practice, make novel discoveries, participate in iterative work and collaborate with peers. The goal is to give students an authentic research experience in which they will acquire necessary skills for being a scientist. Four classroom hours and three hours of laboratory per week.

Requisite: BIOL 214 or BIOL 260. Limited to 18 students. Spring semester. Professor Kim.

2022-23: Offered in Spring 2023

311 Small Proteins in Antibiotic Resistance with Lab

This lab course will provide students a hands-on mentored research experience in protein biochemistry and molecular microbiology. Through structured primary literature analysis and studying historic examples, students will learn how to develop scientific questions, structure a convincing scientific argument, design experiments to address those questions, and critically analyze data and present findings to a scientific community. In addition to participation in classroom workshops and discussions, students will work in teams to design and execute a short research project in an area of active faculty research investigating a small protein involved in antibiotic resistance in the bacterial model organism Escherichia coli. Students will present the results of their work in a short research talk and in a poster session. At the end of this course, students will be able to apply the fundamental skills used to conduct scientific research.

Professor Wu Orr

2022-23: Offered in Spring 2023

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; BIOL 191 recommended. Limited to 30 students. Not open to first-year students. Spring semester. Professor Miller

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2024

330 Biochemical Principles of Life at the Molecular Level

(Offered as CHEM 330 and BIOL 330) What are the molecular underpinnings of processes central to life? We will explore the chemical and structural properties of biological molecules and learn the logic used by the cell to build complex structures from a few basic raw materials. Some of these complex structures have evolved to catalyze chemical reactions with an enormous degree of selectivity and specificity, and we seek to discover these enzymatic strategies. We will consider the detailed balance sheet that shows how living things harvest energy from their environment to fuel metabolic processes and to reproduce and grow. Examples of the exquisite control that permits a cell to be responsive and adapt its responses based on input from the environment will be considered. We will also consider some of the means by which cells respond to change and to stress. A student may not receive credit for both CHEM/BIOL 330 and BCBP/BIOL/CHEM 331. Note: BCBP/BIOL/CHEM 331 is a requirement for the biochemistry track of the BCBP major, so prospective BCBP majors should not enroll in CHEM/BIOL 330 if they are considering the biochemistry track of the major.

Requisite: BIOL 191 and CHEM  221. Limited to 40 students with 20 students per discussion section.

Spring Semester: Professor Bishop.

2022-23: Offered in Spring 2023
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Spring 2020, Spring 2021, Spring 2022, Spring 2024

331 Biochemistry with Lab

(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 CHEM/BIOL 330 and BCBP/BIOL/CHEM 331.

BCBP/BIOL/CHEM 331 is a requirement for the biochemistry track of the BCBP major, so prospective BCBP majors should not enroll in CHEM/BIOL 330 if they are considering the biochemistry track of the major.

Requisite: CHEM 231 and BIOL 191; or consent of the instructor. (CHEM 231 may be taken either as a prerequisite or as a co-requisite.) Limited to 30 students. Fall semester. Professors Jaswal and Wu Orr. 

2022-23: Offered in Fall 2022
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

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. 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.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2012, Fall 2013, Fall 2016, Fall 2022

371 Molecular Genetics with 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 30 students. Not open to first-year students. Omitted 2022-23. Visiting Professor Heussler.

2022-23: Not offered
Other years: Offered in Fall 2017, Fall 2018, Fall 2019, Spring 2020, Fall 2020, Fall 2021, Spring 2024

391 Quantitative Imaging with Lab: From Molecules to Cells and Beyond

(Offered as BIOL 391, BCBP 391, and NEUR-391) Fluorescence imaging offers a window to understanding the structure and function of living cells at sub-cellular resolution. This laboratory and lecture course will focus on advanced quantitative fluorescence microscopy techniques used for imaging a range of biological model systems and functions ranging from single molecules to cells to tissues. Students will learn the fundamentals of optics and microscopy and gain hands-on experience with classical and emerging imaging technology including: confocal microscopy, deconvolution and super-resolution methods, and live-cell imaging. Fluorescent probes and genetically encoded biosensors will be discussed and used to study dynamic cellular processes. In addition, the class will explore a range of image processing and analysis workflows. Lectures dealing with the theory, mechanics, and application of fluorescent imaging methods will be intermingled with extended laboratory sessions in our new Amherst Biological Imaging Center.

Pre requisites : BIOL 191 and PHYS 117 and one of BIOL/BCBP291, BIOL221 or BIOL/NEUR 301, or with instructor permission.

There will be two 80-minute lectures and a weekly laboratory. Limited to 12 students. Fall semester. Professors Edwards and Kim  

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2022, Spring 2024

400 Molecular and Cellular Biophysics

(Offered as PHYS 400, BIOL 400, BCBP 400, and CHEM 400) How do the physical laws that dominate our lives change at the small length and energy scales of individual molecules? What design principles break down at the sub-cellular level and what new chemistry and physics becomes important? We will answer these questions by looking at bio-molecules, cellular substructures, and control mechanisms that work effectively in the microscopic world. How can we understand both the static and dynamic shape of proteins using the laws of thermodynamics and kinetics? How has the basic understanding of the smallest molecular motor in the world, ATP synthase, changed our understanding of friction and torque? We will explore new technologies, such as atomic force and single molecule microscopy that have allowed research into these areas. This course will address topics in each of the three major divisions of Biophysics: bio-molecular structure, biophysical techniques, and biological mechanisms.

Requisite: CHEM 161/165, PHYS 116/123, PHYS 117/124, BIOL 191 or evidence of equivalent coverage in pre-collegiate courses. Spring semester. Professor Carter. 

2022-23: Offered in Spring 2023
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Spring 2015, Spring 2016, Spring 2017, Spring 2018, Spring 2019, Spring 2020, Spring 2021, Spring 2022, Spring 2024

424 Seminar in 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. Spring semester. Prof. Miller. 

2022-23: Offered in Spring 2023
Other years: Offered in Fall 2023

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. Fall 2022. Professor Temeles.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2011, Spring 2013, Fall 2014, Spring 2016, Spring 2019, Spring 2020, Spring 2021, Fall 2022

440, 441 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 close reading and discussion of articles from the primary literature, the course will explore key topics including overexploitation (including connections between the wildlife trade and emergent diseases such as COVID-19), habitat fragmentation, climate change, restoration, protected areas, payments for ecosystem services, as well as how to determine appropriate conservation priorities. Three classroom hours per week.

Requisite: BIOL 230/ENST 210 or BIOL 320, or consent of the instructor. Not open to first-year students. Limited to 14 students. Spring Semester. Senior Lecturer Levin.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2012, Fall 2013, Fall 2015, Fall 2016, Spring 2018, Fall 2019, Fall 2020, Spring 2022

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 BIOL-214, BIOL 260, BIOL 350/351, or consent of the instructor. Limited to 18 students. Not open to first-year students. Spring Semester. Professor Trapani.

2022-23: Not offered
Other years: Offered in Spring 2013, Spring 2017, Spring 2019, Spring 2022, Spring 2024

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 or ENST 200. 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. Omitted 2022-23. Professor Clotfelter and Senior Lecturer Levin.

2022-23: Not offered
Other years: Offered in Spring 2024

498, 499, 499D Senior Departmental Honors

Honors students take three courses of thesis research, usually, but not always, with the double course load in the spring (499D). The work consists of seminar programs, individual research projects, and preparation of a thesis on the research project.

Open to seniors. Fall semester. The Department.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Fall 2019, Fall 2020, Fall 2021, Fall 2022, Fall 2023

Related Courses

AMST-296 Black Women and Reproductive Justice in the African Diaspora (Course not offered this year.)
ANTH-245 Medical Anthropology (Course not offered this year.)
ECON-212 Public Economics: Environment, Health, and Inequality (Course not offered this year.)
SOCI-328 The Pandemic (Course not offered this year.)

Amherst College Courses

Amherst College Courses

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Biology

Professors Clotfelter‡, Goutte, Hood‡, Miller†, and Temeles; Associate Professors Purdy (Chair) and Trapani; Assistant Professors Edwards, Jeong*, Kim, Ragkousi and Wu Orr; Senior Lecturer Levin†; Visiting Assistant Professor Roche; Visiting Lecturer Heussler; Lab Coordinators Bowers 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 five categories:

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.

Five advanced courses in Biology, three of which must involve laboratories and one of which must be at the 400 level. Note that Special Topics courses 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.

Among the five advanced courses, one course in each of the two core areas of biology should be completed by the end of the junior year:

i) Cellular/Molecular Core: either Cell Structure and Function (BIOL 291) or Molecular Genetics (BIOL 370/371).
ii) Organismal Core: either Ecology (BIOL 230) or Evolutionary Biology (BIOL 320/321).

Two courses in chemistry (CHEM 151 or 155 and CHEM 161 or 165 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 151/155 and/or CHEM 161/165 are requisites for several Biology courses. Students are therefore encouraged to take CHEM 151/155 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 117/124 is recommended for BIOL 351 and BIOL 400.

One course to fulfill the STEM diversity, equity, and inclusion requirement. For the Class of 2023 and beyond, Biology Majors must take one science-oriented course that is approved by the department and that focuses on structural and systemic issues of diversity, equity, and inclusion.

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 usually 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; students placing out of BIOL 191 must take both courses from the Cellular/Molecular Core. 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). Placing out will not reduce the number of courses required for the major.

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 2022-23.†On leave fall semester 2022-23.‡On leave spring semester 2022-2023.

150, 250 Being Human in STEM

(Offered as BIOL-150, CHEM-250) This is an interactive course that combines academic inquiry and community engagement to investigate identity, inequality and representation within Science Technology Engineering and Mathematics (STEM) fields--at Amherst and beyond. We begin the course by grounding our understanding of the STEM experience at Amherst in national and global contexts. We will survey the interdisciplinary literature on the ways in which identity - race, gender, class, ability, sexuality- and geographic context shape STEM persistence and belonging. We will bring this literature into conversation with our own Amherst experiences. These challenging conversations require vulnerability, openness and the ability to tolerate discomfort. We will work from day one to build a brave space whose foundation is trust, accountability and growth. Students will design group projects that apply themes from the literature and our seminar discussions to develop resources and engage the STEM community, whether at the college, local, or national level. Course work includes critical reading and discussion, reflective writing, and collaborative work culminating in community engagement proposals which students will share with the campus and the broader public.

Open to sophomores, juniors and seniors. This course will be taught in two sections

Offered Spring Semester: Professors Jaswal and Trapani.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2020, January 2021, January 2022, Spring 2022, Spring 2024

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 Hood and Temeles; Lab Coordinator Kristensen.

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

2022-23: Offered in Fall 2022
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

181L Adaptation & Organism Lab

Lab 

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2022, Fall 2023

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 or CHEM 165. Limited to 96 students. Fall semester: Professors Goutte and Edwards, Lab Coordinator Bowers. Spring semester: Professors Purdy and Ragkousi, Lab Coordinator Bowers. 

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

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Spring 2019, Fall 2022, Fall 2023, Spring 2024

191L Molecules, Genes, & Cells Lab

Lab

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2022, Fall 2023, Spring 2024

210, 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 110 or equivalent. Limited to 40 students. Spring Semester. Professor Temeles.

2022-23: Offered in Spring 2023
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 2024

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 emphasis 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. Fall and Spring semester. Professors Kim and Roche.

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2019, Fall 2020, Fall 2021, Fall 2022, Fall 2023

221 Developmental Biology with Lab

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 2022. Professor Ragkousi.

 

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2014, Spring 2016, Fall 2022, Spring 2024

241 Genetic Analysis with lab

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 16 students. Not open to first-year students. Spring Semester. Professor Goutte.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2012, Spring 2013, Spring 2014, Spring 2015, Spring 2017, Spring 2018, Spring 2022

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. Fall 2022. Professor Purdy.

2022-23: Offered in Fall 2022
Other years: Offered in Spring 2018, Spring 2019, Fall 2022, Spring 2024

275H Evolutionary Ecology Laboratory

This laboratory explores how underlying evolutionary changes in the morphology, physiology or behavior of organisms is associated with ecological interactions within or between species.  By employing artificial selection, students will determine whether genetic variation associated with herbivore defense exists.  This experiment will be coupled with ecological investigations to explore the potential of herbivory to drive evolutionary change.  Further, using the model system Daphnia, students will have the opportunity to engage in a self-directed, in-depth investigation of selection on morphological and life history traits across varying environmental conditions.  Throughout the course, students will engage with the scientific literature, and develop their skills in identifying relevant and important questions, experimental design and hypothesis formulation, data analysis and scientific communication.  Three hours of laboratory each week.

Requisite: Prior completion of or concurrent registration in either BIOL-230 or BIOL-320. Limited to 16 students. Professors Miller and Temeles and Lab Instructor Kristensen. Spring semester.

2022-23: Offered in Spring 2023

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. Four classroom hours per week.

Requisite: BIOL 181. Limited to 20 students. Fall semester. Professor Clotfelter.

2022-23: Offered in Fall 2022
Other years: Offered in Spring 2015, Fall 2015, Fall 2016, Fall 2017, Spring 2019, Fall 2019, Fall 2022, Fall 2023

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.

In Fall 2022, the laboratory will be structured as a course-based undergraduate research experience (CURE).  The focus will be a semester-long, field-based project on bird behavior in the Wildlife Sanctuary.  Students will be outdoors in the Sanctuary under almost all weather conditions.  Students can register for either laboratory section, both of which will be held at low enrollments to facilitate field research.

Requisite: BIOL 181. Limited to 8 students per lab section. Fall semester. Professor Clotfelter.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2011, Fall 2013, Fall 2022, Fall 2023

290, 290H, 390, 490, 490H Special Topics

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

Fall and spring semesters. The Department.

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2011, Spring 2012, Fall 2012, Spring 2013, Fall 2013, Spring 2014, Fall 2014, Spring 2015, Fall 2015, Spring 2016, Fall 2016, Spring 2017, Fall 2017, Spring 2018, Fall 2018, Spring 2019, Fall 2019, Spring 2020, Fall 2020, Spring 2021, Fall 2021, Spring 2022, Fall 2022, Fall 2023, Spring 2024

291 Cell Structure and Function with Lab

Offered as BIOL 291 and BCBP 291) An analysis of the structure and function of eukaryotic cells. 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. There will be three classroom hours consisting of both lectures and problem-solving sessions, and three hours of laboratory per week. The course will be taught in-person.

Requisite: BIOL 191, CHEM151/155, and CHEM161/165. Limited to 16 students. Spring semester. Professor Edwards.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2012, Spring 2013, Spring 2014, Spring 2015, Spring 2016, Spring 2017, Spring 2018, Spring 2019, Fall 2023

301 Molecular Neurobiology with Lab

(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 and synapse development as well as neuronal 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 supplement the lectures and also emphasize current molecular techniques utilized in the study of the nervous system. Four classroom hours and three hours of laboratory per week.

While enrolled at the college, students may only take either BIOL/NEUR-301 or BIOL/NEUR-303.

Requisite: Bio 191. Bio/Neur 214 strongly suggested.

Limited to 18 students. Spring semester. Professor Roche.

2022-23: Not offered
Other years: Offered in Fall 2011, Fall 2014, Fall 2015, Fall 2016, Spring 2020, Spring 2021, Spring 2022, Fall 2023

303 Molecular Neurobiology: CURE

(Offered as BIOL 303 and NEUR 303) 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. This course is a CURE, a Course-based Undergraduate Research Experience. Students will engage in scientific practice, make novel discoveries, participate in iterative work and collaborate with peers. The goal is to give students an authentic research experience in which they will acquire necessary skills for being a scientist. Four classroom hours and three hours of laboratory per week.

Requisite: BIOL 214 or BIOL 260. Limited to 18 students. Spring semester. Professor Kim.

2022-23: Offered in Spring 2023

311 Small Proteins in Antibiotic Resistance with Lab

This lab course will provide students a hands-on mentored research experience in protein biochemistry and molecular microbiology. Through structured primary literature analysis and studying historic examples, students will learn how to develop scientific questions, structure a convincing scientific argument, design experiments to address those questions, and critically analyze data and present findings to a scientific community. In addition to participation in classroom workshops and discussions, students will work in teams to design and execute a short research project in an area of active faculty research investigating a small protein involved in antibiotic resistance in the bacterial model organism Escherichia coli. Students will present the results of their work in a short research talk and in a poster session. At the end of this course, students will be able to apply the fundamental skills used to conduct scientific research.

Professor Wu Orr

2022-23: Offered in Spring 2023

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; BIOL 191 recommended. Limited to 30 students. Not open to first-year students. Spring semester. Professor Miller

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2024

330 Biochemical Principles of Life at the Molecular Level

(Offered as CHEM 330 and BIOL 330) What are the molecular underpinnings of processes central to life? We will explore the chemical and structural properties of biological molecules and learn the logic used by the cell to build complex structures from a few basic raw materials. Some of these complex structures have evolved to catalyze chemical reactions with an enormous degree of selectivity and specificity, and we seek to discover these enzymatic strategies. We will consider the detailed balance sheet that shows how living things harvest energy from their environment to fuel metabolic processes and to reproduce and grow. Examples of the exquisite control that permits a cell to be responsive and adapt its responses based on input from the environment will be considered. We will also consider some of the means by which cells respond to change and to stress. A student may not receive credit for both CHEM/BIOL 330 and BCBP/BIOL/CHEM 331. Note: BCBP/BIOL/CHEM 331 is a requirement for the biochemistry track of the BCBP major, so prospective BCBP majors should not enroll in CHEM/BIOL 330 if they are considering the biochemistry track of the major.

Requisite: BIOL 191 and CHEM  221. Limited to 40 students with 20 students per discussion section.

Spring Semester: Professor Bishop.

2022-23: Offered in Spring 2023
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Spring 2020, Spring 2021, Spring 2022, Spring 2024

331 Biochemistry with Lab

(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 CHEM/BIOL 330 and BCBP/BIOL/CHEM 331.

BCBP/BIOL/CHEM 331 is a requirement for the biochemistry track of the BCBP major, so prospective BCBP majors should not enroll in CHEM/BIOL 330 if they are considering the biochemistry track of the major.

Requisite: CHEM 231 and BIOL 191; or consent of the instructor. (CHEM 231 may be taken either as a prerequisite or as a co-requisite.) Limited to 30 students. Fall semester. Professors Jaswal and Wu Orr. 

2022-23: Offered in Fall 2022
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

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. 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.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2012, Fall 2013, Fall 2016, Fall 2022

371 Molecular Genetics with 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 30 students. Not open to first-year students. Omitted 2022-23. Visiting Professor Heussler.

2022-23: Not offered
Other years: Offered in Fall 2017, Fall 2018, Fall 2019, Spring 2020, Fall 2020, Fall 2021, Spring 2024

391 Quantitative Imaging with Lab: From Molecules to Cells and Beyond

(Offered as BIOL 391, BCBP 391, and NEUR-391) Fluorescence imaging offers a window to understanding the structure and function of living cells at sub-cellular resolution. This laboratory and lecture course will focus on advanced quantitative fluorescence microscopy techniques used for imaging a range of biological model systems and functions ranging from single molecules to cells to tissues. Students will learn the fundamentals of optics and microscopy and gain hands-on experience with classical and emerging imaging technology including: confocal microscopy, deconvolution and super-resolution methods, and live-cell imaging. Fluorescent probes and genetically encoded biosensors will be discussed and used to study dynamic cellular processes. In addition, the class will explore a range of image processing and analysis workflows. Lectures dealing with the theory, mechanics, and application of fluorescent imaging methods will be intermingled with extended laboratory sessions in our new Amherst Biological Imaging Center.

Pre requisites : BIOL 191 and PHYS 117 and one of BIOL/BCBP291, BIOL221 or BIOL/NEUR 301, or with instructor permission.

There will be two 80-minute lectures and a weekly laboratory. Limited to 12 students. Fall semester. Professors Edwards and Kim  

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2022, Spring 2024

400 Molecular and Cellular Biophysics

(Offered as PHYS 400, BIOL 400, BCBP 400, and CHEM 400) How do the physical laws that dominate our lives change at the small length and energy scales of individual molecules? What design principles break down at the sub-cellular level and what new chemistry and physics becomes important? We will answer these questions by looking at bio-molecules, cellular substructures, and control mechanisms that work effectively in the microscopic world. How can we understand both the static and dynamic shape of proteins using the laws of thermodynamics and kinetics? How has the basic understanding of the smallest molecular motor in the world, ATP synthase, changed our understanding of friction and torque? We will explore new technologies, such as atomic force and single molecule microscopy that have allowed research into these areas. This course will address topics in each of the three major divisions of Biophysics: bio-molecular structure, biophysical techniques, and biological mechanisms.

Requisite: CHEM 161/165, PHYS 116/123, PHYS 117/124, BIOL 191 or evidence of equivalent coverage in pre-collegiate courses. Spring semester. Professor Carter. 

2022-23: Offered in Spring 2023
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Spring 2015, Spring 2016, Spring 2017, Spring 2018, Spring 2019, Spring 2020, Spring 2021, Spring 2022, Spring 2024

424 Seminar in 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. Spring semester. Prof. Miller. 

2022-23: Offered in Spring 2023
Other years: Offered in Fall 2023

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. Fall 2022. Professor Temeles.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2011, Spring 2013, Fall 2014, Spring 2016, Spring 2019, Spring 2020, Spring 2021, Fall 2022

440, 441 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 close reading and discussion of articles from the primary literature, the course will explore key topics including overexploitation (including connections between the wildlife trade and emergent diseases such as COVID-19), habitat fragmentation, climate change, restoration, protected areas, payments for ecosystem services, as well as how to determine appropriate conservation priorities. Three classroom hours per week.

Requisite: BIOL 230/ENST 210 or BIOL 320, or consent of the instructor. Not open to first-year students. Limited to 14 students. Spring Semester. Senior Lecturer Levin.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2012, Fall 2013, Fall 2015, Fall 2016, Spring 2018, Fall 2019, Fall 2020, Spring 2022

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 BIOL-214, BIOL 260, BIOL 350/351, or consent of the instructor. Limited to 18 students. Not open to first-year students. Spring Semester. Professor Trapani.

2022-23: Not offered
Other years: Offered in Spring 2013, Spring 2017, Spring 2019, Spring 2022, Spring 2024

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 or ENST 200. 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. Omitted 2022-23. Professor Clotfelter and Senior Lecturer Levin.

2022-23: Not offered
Other years: Offered in Spring 2024

498, 499, 499D Senior Departmental Honors

Honors students take three courses of thesis research, usually, but not always, with the double course load in the spring (499D). The work consists of seminar programs, individual research projects, and preparation of a thesis on the research project.

Open to seniors. Fall semester. The Department.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Fall 2019, Fall 2020, Fall 2021, Fall 2022, Fall 2023

Related Courses

AMST-296 Black Women and Reproductive Justice in the African Diaspora (Course not offered this year.)
ANTH-245 Medical Anthropology (Course not offered this year.)
ECON-212 Public Economics: Environment, Health, and Inequality (Course not offered this year.)
SOCI-328 The Pandemic (Course not offered this year.)

Five College Programs & Certificates

Five College Programs & Certificates

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Biology

Professors Clotfelter‡, Goutte, Hood‡, Miller†, and Temeles; Associate Professors Purdy (Chair) and Trapani; Assistant Professors Edwards, Jeong*, Kim, Ragkousi and Wu Orr; Senior Lecturer Levin†; Visiting Assistant Professor Roche; Visiting Lecturer Heussler; Lab Coordinators Bowers 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 five categories:

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.

Five advanced courses in Biology, three of which must involve laboratories and one of which must be at the 400 level. Note that Special Topics courses 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.

Among the five advanced courses, one course in each of the two core areas of biology should be completed by the end of the junior year:

i) Cellular/Molecular Core: either Cell Structure and Function (BIOL 291) or Molecular Genetics (BIOL 370/371).
ii) Organismal Core: either Ecology (BIOL 230) or Evolutionary Biology (BIOL 320/321).

Two courses in chemistry (CHEM 151 or 155 and CHEM 161 or 165 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 151/155 and/or CHEM 161/165 are requisites for several Biology courses. Students are therefore encouraged to take CHEM 151/155 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 117/124 is recommended for BIOL 351 and BIOL 400.

One course to fulfill the STEM diversity, equity, and inclusion requirement. For the Class of 2023 and beyond, Biology Majors must take one science-oriented course that is approved by the department and that focuses on structural and systemic issues of diversity, equity, and inclusion.

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 usually 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; students placing out of BIOL 191 must take both courses from the Cellular/Molecular Core. 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). Placing out will not reduce the number of courses required for the major.

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 2022-23.†On leave fall semester 2022-23.‡On leave spring semester 2022-2023.

150, 250 Being Human in STEM

(Offered as BIOL-150, CHEM-250) This is an interactive course that combines academic inquiry and community engagement to investigate identity, inequality and representation within Science Technology Engineering and Mathematics (STEM) fields--at Amherst and beyond. We begin the course by grounding our understanding of the STEM experience at Amherst in national and global contexts. We will survey the interdisciplinary literature on the ways in which identity - race, gender, class, ability, sexuality- and geographic context shape STEM persistence and belonging. We will bring this literature into conversation with our own Amherst experiences. These challenging conversations require vulnerability, openness and the ability to tolerate discomfort. We will work from day one to build a brave space whose foundation is trust, accountability and growth. Students will design group projects that apply themes from the literature and our seminar discussions to develop resources and engage the STEM community, whether at the college, local, or national level. Course work includes critical reading and discussion, reflective writing, and collaborative work culminating in community engagement proposals which students will share with the campus and the broader public.

Open to sophomores, juniors and seniors. This course will be taught in two sections

Offered Spring Semester: Professors Jaswal and Trapani.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2020, January 2021, January 2022, Spring 2022, Spring 2024

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 Hood and Temeles; Lab Coordinator Kristensen.

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

2022-23: Offered in Fall 2022
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

181L Adaptation & Organism Lab

Lab 

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2022, Fall 2023

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 or CHEM 165. Limited to 96 students. Fall semester: Professors Goutte and Edwards, Lab Coordinator Bowers. Spring semester: Professors Purdy and Ragkousi, Lab Coordinator Bowers. 

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

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Spring 2019, Fall 2022, Fall 2023, Spring 2024

191L Molecules, Genes, & Cells Lab

Lab

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2022, Fall 2023, Spring 2024

210, 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 110 or equivalent. Limited to 40 students. Spring Semester. Professor Temeles.

2022-23: Offered in Spring 2023
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 2024

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 emphasis 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. Fall and Spring semester. Professors Kim and Roche.

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2019, Fall 2020, Fall 2021, Fall 2022, Fall 2023

221 Developmental Biology with Lab

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 2022. Professor Ragkousi.

 

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2014, Spring 2016, Fall 2022, Spring 2024

241 Genetic Analysis with lab

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 16 students. Not open to first-year students. Spring Semester. Professor Goutte.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2012, Spring 2013, Spring 2014, Spring 2015, Spring 2017, Spring 2018, Spring 2022

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. Fall 2022. Professor Purdy.

2022-23: Offered in Fall 2022
Other years: Offered in Spring 2018, Spring 2019, Fall 2022, Spring 2024

275H Evolutionary Ecology Laboratory

This laboratory explores how underlying evolutionary changes in the morphology, physiology or behavior of organisms is associated with ecological interactions within or between species.  By employing artificial selection, students will determine whether genetic variation associated with herbivore defense exists.  This experiment will be coupled with ecological investigations to explore the potential of herbivory to drive evolutionary change.  Further, using the model system Daphnia, students will have the opportunity to engage in a self-directed, in-depth investigation of selection on morphological and life history traits across varying environmental conditions.  Throughout the course, students will engage with the scientific literature, and develop their skills in identifying relevant and important questions, experimental design and hypothesis formulation, data analysis and scientific communication.  Three hours of laboratory each week.

Requisite: Prior completion of or concurrent registration in either BIOL-230 or BIOL-320. Limited to 16 students. Professors Miller and Temeles and Lab Instructor Kristensen. Spring semester.

2022-23: Offered in Spring 2023

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. Four classroom hours per week.

Requisite: BIOL 181. Limited to 20 students. Fall semester. Professor Clotfelter.

2022-23: Offered in Fall 2022
Other years: Offered in Spring 2015, Fall 2015, Fall 2016, Fall 2017, Spring 2019, Fall 2019, Fall 2022, Fall 2023

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.

In Fall 2022, the laboratory will be structured as a course-based undergraduate research experience (CURE).  The focus will be a semester-long, field-based project on bird behavior in the Wildlife Sanctuary.  Students will be outdoors in the Sanctuary under almost all weather conditions.  Students can register for either laboratory section, both of which will be held at low enrollments to facilitate field research.

Requisite: BIOL 181. Limited to 8 students per lab section. Fall semester. Professor Clotfelter.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2011, Fall 2013, Fall 2022, Fall 2023

290, 290H, 390, 490, 490H Special Topics

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

Fall and spring semesters. The Department.

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2011, Spring 2012, Fall 2012, Spring 2013, Fall 2013, Spring 2014, Fall 2014, Spring 2015, Fall 2015, Spring 2016, Fall 2016, Spring 2017, Fall 2017, Spring 2018, Fall 2018, Spring 2019, Fall 2019, Spring 2020, Fall 2020, Spring 2021, Fall 2021, Spring 2022, Fall 2022, Fall 2023, Spring 2024

291 Cell Structure and Function with Lab

Offered as BIOL 291 and BCBP 291) An analysis of the structure and function of eukaryotic cells. 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. There will be three classroom hours consisting of both lectures and problem-solving sessions, and three hours of laboratory per week. The course will be taught in-person.

Requisite: BIOL 191, CHEM151/155, and CHEM161/165. Limited to 16 students. Spring semester. Professor Edwards.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2012, Spring 2013, Spring 2014, Spring 2015, Spring 2016, Spring 2017, Spring 2018, Spring 2019, Fall 2023

301 Molecular Neurobiology with Lab

(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 and synapse development as well as neuronal 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 supplement the lectures and also emphasize current molecular techniques utilized in the study of the nervous system. Four classroom hours and three hours of laboratory per week.

While enrolled at the college, students may only take either BIOL/NEUR-301 or BIOL/NEUR-303.

Requisite: Bio 191. Bio/Neur 214 strongly suggested.

Limited to 18 students. Spring semester. Professor Roche.

2022-23: Not offered
Other years: Offered in Fall 2011, Fall 2014, Fall 2015, Fall 2016, Spring 2020, Spring 2021, Spring 2022, Fall 2023

303 Molecular Neurobiology: CURE

(Offered as BIOL 303 and NEUR 303) 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. This course is a CURE, a Course-based Undergraduate Research Experience. Students will engage in scientific practice, make novel discoveries, participate in iterative work and collaborate with peers. The goal is to give students an authentic research experience in which they will acquire necessary skills for being a scientist. Four classroom hours and three hours of laboratory per week.

Requisite: BIOL 214 or BIOL 260. Limited to 18 students. Spring semester. Professor Kim.

2022-23: Offered in Spring 2023

311 Small Proteins in Antibiotic Resistance with Lab

This lab course will provide students a hands-on mentored research experience in protein biochemistry and molecular microbiology. Through structured primary literature analysis and studying historic examples, students will learn how to develop scientific questions, structure a convincing scientific argument, design experiments to address those questions, and critically analyze data and present findings to a scientific community. In addition to participation in classroom workshops and discussions, students will work in teams to design and execute a short research project in an area of active faculty research investigating a small protein involved in antibiotic resistance in the bacterial model organism Escherichia coli. Students will present the results of their work in a short research talk and in a poster session. At the end of this course, students will be able to apply the fundamental skills used to conduct scientific research.

Professor Wu Orr

2022-23: Offered in Spring 2023

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; BIOL 191 recommended. Limited to 30 students. Not open to first-year students. Spring semester. Professor Miller

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2024

330 Biochemical Principles of Life at the Molecular Level

(Offered as CHEM 330 and BIOL 330) What are the molecular underpinnings of processes central to life? We will explore the chemical and structural properties of biological molecules and learn the logic used by the cell to build complex structures from a few basic raw materials. Some of these complex structures have evolved to catalyze chemical reactions with an enormous degree of selectivity and specificity, and we seek to discover these enzymatic strategies. We will consider the detailed balance sheet that shows how living things harvest energy from their environment to fuel metabolic processes and to reproduce and grow. Examples of the exquisite control that permits a cell to be responsive and adapt its responses based on input from the environment will be considered. We will also consider some of the means by which cells respond to change and to stress. A student may not receive credit for both CHEM/BIOL 330 and BCBP/BIOL/CHEM 331. Note: BCBP/BIOL/CHEM 331 is a requirement for the biochemistry track of the BCBP major, so prospective BCBP majors should not enroll in CHEM/BIOL 330 if they are considering the biochemistry track of the major.

Requisite: BIOL 191 and CHEM  221. Limited to 40 students with 20 students per discussion section.

Spring Semester: Professor Bishop.

2022-23: Offered in Spring 2023
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Spring 2020, Spring 2021, Spring 2022, Spring 2024

331 Biochemistry with Lab

(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 CHEM/BIOL 330 and BCBP/BIOL/CHEM 331.

BCBP/BIOL/CHEM 331 is a requirement for the biochemistry track of the BCBP major, so prospective BCBP majors should not enroll in CHEM/BIOL 330 if they are considering the biochemistry track of the major.

Requisite: CHEM 231 and BIOL 191; or consent of the instructor. (CHEM 231 may be taken either as a prerequisite or as a co-requisite.) Limited to 30 students. Fall semester. Professors Jaswal and Wu Orr. 

2022-23: Offered in Fall 2022
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

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. 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.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2012, Fall 2013, Fall 2016, Fall 2022

371 Molecular Genetics with 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 30 students. Not open to first-year students. Omitted 2022-23. Visiting Professor Heussler.

2022-23: Not offered
Other years: Offered in Fall 2017, Fall 2018, Fall 2019, Spring 2020, Fall 2020, Fall 2021, Spring 2024

391 Quantitative Imaging with Lab: From Molecules to Cells and Beyond

(Offered as BIOL 391, BCBP 391, and NEUR-391) Fluorescence imaging offers a window to understanding the structure and function of living cells at sub-cellular resolution. This laboratory and lecture course will focus on advanced quantitative fluorescence microscopy techniques used for imaging a range of biological model systems and functions ranging from single molecules to cells to tissues. Students will learn the fundamentals of optics and microscopy and gain hands-on experience with classical and emerging imaging technology including: confocal microscopy, deconvolution and super-resolution methods, and live-cell imaging. Fluorescent probes and genetically encoded biosensors will be discussed and used to study dynamic cellular processes. In addition, the class will explore a range of image processing and analysis workflows. Lectures dealing with the theory, mechanics, and application of fluorescent imaging methods will be intermingled with extended laboratory sessions in our new Amherst Biological Imaging Center.

Pre requisites : BIOL 191 and PHYS 117 and one of BIOL/BCBP291, BIOL221 or BIOL/NEUR 301, or with instructor permission.

There will be two 80-minute lectures and a weekly laboratory. Limited to 12 students. Fall semester. Professors Edwards and Kim  

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2022, Spring 2024

400 Molecular and Cellular Biophysics

(Offered as PHYS 400, BIOL 400, BCBP 400, and CHEM 400) How do the physical laws that dominate our lives change at the small length and energy scales of individual molecules? What design principles break down at the sub-cellular level and what new chemistry and physics becomes important? We will answer these questions by looking at bio-molecules, cellular substructures, and control mechanisms that work effectively in the microscopic world. How can we understand both the static and dynamic shape of proteins using the laws of thermodynamics and kinetics? How has the basic understanding of the smallest molecular motor in the world, ATP synthase, changed our understanding of friction and torque? We will explore new technologies, such as atomic force and single molecule microscopy that have allowed research into these areas. This course will address topics in each of the three major divisions of Biophysics: bio-molecular structure, biophysical techniques, and biological mechanisms.

Requisite: CHEM 161/165, PHYS 116/123, PHYS 117/124, BIOL 191 or evidence of equivalent coverage in pre-collegiate courses. Spring semester. Professor Carter. 

2022-23: Offered in Spring 2023
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Spring 2015, Spring 2016, Spring 2017, Spring 2018, Spring 2019, Spring 2020, Spring 2021, Spring 2022, Spring 2024

424 Seminar in 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. Spring semester. Prof. Miller. 

2022-23: Offered in Spring 2023
Other years: Offered in Fall 2023

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. Fall 2022. Professor Temeles.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2011, Spring 2013, Fall 2014, Spring 2016, Spring 2019, Spring 2020, Spring 2021, Fall 2022

440, 441 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 close reading and discussion of articles from the primary literature, the course will explore key topics including overexploitation (including connections between the wildlife trade and emergent diseases such as COVID-19), habitat fragmentation, climate change, restoration, protected areas, payments for ecosystem services, as well as how to determine appropriate conservation priorities. Three classroom hours per week.

Requisite: BIOL 230/ENST 210 or BIOL 320, or consent of the instructor. Not open to first-year students. Limited to 14 students. Spring Semester. Senior Lecturer Levin.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2012, Fall 2013, Fall 2015, Fall 2016, Spring 2018, Fall 2019, Fall 2020, Spring 2022

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 BIOL-214, BIOL 260, BIOL 350/351, or consent of the instructor. Limited to 18 students. Not open to first-year students. Spring Semester. Professor Trapani.

2022-23: Not offered
Other years: Offered in Spring 2013, Spring 2017, Spring 2019, Spring 2022, Spring 2024

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 or ENST 200. 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. Omitted 2022-23. Professor Clotfelter and Senior Lecturer Levin.

2022-23: Not offered
Other years: Offered in Spring 2024

498, 499, 499D Senior Departmental Honors

Honors students take three courses of thesis research, usually, but not always, with the double course load in the spring (499D). The work consists of seminar programs, individual research projects, and preparation of a thesis on the research project.

Open to seniors. Fall semester. The Department.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Fall 2019, Fall 2020, Fall 2021, Fall 2022, Fall 2023

Related Courses

AMST-296 Black Women and Reproductive Justice in the African Diaspora (Course not offered this year.)
ANTH-245 Medical Anthropology (Course not offered this year.)
ECON-212 Public Economics: Environment, Health, and Inequality (Course not offered this year.)
SOCI-328 The Pandemic (Course not offered this year.)

Honors & Fellowships

Honors & Fellowships

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Biology

Professors Clotfelter‡, Goutte, Hood‡, Miller†, and Temeles; Associate Professors Purdy (Chair) and Trapani; Assistant Professors Edwards, Jeong*, Kim, Ragkousi and Wu Orr; Senior Lecturer Levin†; Visiting Assistant Professor Roche; Visiting Lecturer Heussler; Lab Coordinators Bowers 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 five categories:

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.

Five advanced courses in Biology, three of which must involve laboratories and one of which must be at the 400 level. Note that Special Topics courses 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.

Among the five advanced courses, one course in each of the two core areas of biology should be completed by the end of the junior year:

i) Cellular/Molecular Core: either Cell Structure and Function (BIOL 291) or Molecular Genetics (BIOL 370/371).
ii) Organismal Core: either Ecology (BIOL 230) or Evolutionary Biology (BIOL 320/321).

Two courses in chemistry (CHEM 151 or 155 and CHEM 161 or 165 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 151/155 and/or CHEM 161/165 are requisites for several Biology courses. Students are therefore encouraged to take CHEM 151/155 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 117/124 is recommended for BIOL 351 and BIOL 400.

One course to fulfill the STEM diversity, equity, and inclusion requirement. For the Class of 2023 and beyond, Biology Majors must take one science-oriented course that is approved by the department and that focuses on structural and systemic issues of diversity, equity, and inclusion.

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 usually 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; students placing out of BIOL 191 must take both courses from the Cellular/Molecular Core. 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). Placing out will not reduce the number of courses required for the major.

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 2022-23.†On leave fall semester 2022-23.‡On leave spring semester 2022-2023.

150, 250 Being Human in STEM

(Offered as BIOL-150, CHEM-250) This is an interactive course that combines academic inquiry and community engagement to investigate identity, inequality and representation within Science Technology Engineering and Mathematics (STEM) fields--at Amherst and beyond. We begin the course by grounding our understanding of the STEM experience at Amherst in national and global contexts. We will survey the interdisciplinary literature on the ways in which identity - race, gender, class, ability, sexuality- and geographic context shape STEM persistence and belonging. We will bring this literature into conversation with our own Amherst experiences. These challenging conversations require vulnerability, openness and the ability to tolerate discomfort. We will work from day one to build a brave space whose foundation is trust, accountability and growth. Students will design group projects that apply themes from the literature and our seminar discussions to develop resources and engage the STEM community, whether at the college, local, or national level. Course work includes critical reading and discussion, reflective writing, and collaborative work culminating in community engagement proposals which students will share with the campus and the broader public.

Open to sophomores, juniors and seniors. This course will be taught in two sections

Offered Spring Semester: Professors Jaswal and Trapani.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2020, January 2021, January 2022, Spring 2022, Spring 2024

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 Hood and Temeles; Lab Coordinator Kristensen.

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

2022-23: Offered in Fall 2022
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

181L Adaptation & Organism Lab

Lab 

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2022, Fall 2023

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 or CHEM 165. Limited to 96 students. Fall semester: Professors Goutte and Edwards, Lab Coordinator Bowers. Spring semester: Professors Purdy and Ragkousi, Lab Coordinator Bowers. 

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

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Spring 2019, Fall 2022, Fall 2023, Spring 2024

191L Molecules, Genes, & Cells Lab

Lab

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2022, Fall 2023, Spring 2024

210, 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 110 or equivalent. Limited to 40 students. Spring Semester. Professor Temeles.

2022-23: Offered in Spring 2023
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 2024

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 emphasis 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. Fall and Spring semester. Professors Kim and Roche.

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2019, Fall 2020, Fall 2021, Fall 2022, Fall 2023

221 Developmental Biology with Lab

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 2022. Professor Ragkousi.

 

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2014, Spring 2016, Fall 2022, Spring 2024

241 Genetic Analysis with lab

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 16 students. Not open to first-year students. Spring Semester. Professor Goutte.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2012, Spring 2013, Spring 2014, Spring 2015, Spring 2017, Spring 2018, Spring 2022

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. Fall 2022. Professor Purdy.

2022-23: Offered in Fall 2022
Other years: Offered in Spring 2018, Spring 2019, Fall 2022, Spring 2024

275H Evolutionary Ecology Laboratory

This laboratory explores how underlying evolutionary changes in the morphology, physiology or behavior of organisms is associated with ecological interactions within or between species.  By employing artificial selection, students will determine whether genetic variation associated with herbivore defense exists.  This experiment will be coupled with ecological investigations to explore the potential of herbivory to drive evolutionary change.  Further, using the model system Daphnia, students will have the opportunity to engage in a self-directed, in-depth investigation of selection on morphological and life history traits across varying environmental conditions.  Throughout the course, students will engage with the scientific literature, and develop their skills in identifying relevant and important questions, experimental design and hypothesis formulation, data analysis and scientific communication.  Three hours of laboratory each week.

Requisite: Prior completion of or concurrent registration in either BIOL-230 or BIOL-320. Limited to 16 students. Professors Miller and Temeles and Lab Instructor Kristensen. Spring semester.

2022-23: Offered in Spring 2023

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. Four classroom hours per week.

Requisite: BIOL 181. Limited to 20 students. Fall semester. Professor Clotfelter.

2022-23: Offered in Fall 2022
Other years: Offered in Spring 2015, Fall 2015, Fall 2016, Fall 2017, Spring 2019, Fall 2019, Fall 2022, Fall 2023

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.

In Fall 2022, the laboratory will be structured as a course-based undergraduate research experience (CURE).  The focus will be a semester-long, field-based project on bird behavior in the Wildlife Sanctuary.  Students will be outdoors in the Sanctuary under almost all weather conditions.  Students can register for either laboratory section, both of which will be held at low enrollments to facilitate field research.

Requisite: BIOL 181. Limited to 8 students per lab section. Fall semester. Professor Clotfelter.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2011, Fall 2013, Fall 2022, Fall 2023

290, 290H, 390, 490, 490H Special Topics

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

Fall and spring semesters. The Department.

2022-23: Offered in Fall 2022, Spring 2023
Other years: Offered in Fall 2011, Spring 2012, Fall 2012, Spring 2013, Fall 2013, Spring 2014, Fall 2014, Spring 2015, Fall 2015, Spring 2016, Fall 2016, Spring 2017, Fall 2017, Spring 2018, Fall 2018, Spring 2019, Fall 2019, Spring 2020, Fall 2020, Spring 2021, Fall 2021, Spring 2022, Fall 2022, Fall 2023, Spring 2024

291 Cell Structure and Function with Lab

Offered as BIOL 291 and BCBP 291) An analysis of the structure and function of eukaryotic cells. 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. There will be three classroom hours consisting of both lectures and problem-solving sessions, and three hours of laboratory per week. The course will be taught in-person.

Requisite: BIOL 191, CHEM151/155, and CHEM161/165. Limited to 16 students. Spring semester. Professor Edwards.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2012, Spring 2013, Spring 2014, Spring 2015, Spring 2016, Spring 2017, Spring 2018, Spring 2019, Fall 2023

301 Molecular Neurobiology with Lab

(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 and synapse development as well as neuronal 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 supplement the lectures and also emphasize current molecular techniques utilized in the study of the nervous system. Four classroom hours and three hours of laboratory per week.

While enrolled at the college, students may only take either BIOL/NEUR-301 or BIOL/NEUR-303.

Requisite: Bio 191. Bio/Neur 214 strongly suggested.

Limited to 18 students. Spring semester. Professor Roche.

2022-23: Not offered
Other years: Offered in Fall 2011, Fall 2014, Fall 2015, Fall 2016, Spring 2020, Spring 2021, Spring 2022, Fall 2023

303 Molecular Neurobiology: CURE

(Offered as BIOL 303 and NEUR 303) 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. This course is a CURE, a Course-based Undergraduate Research Experience. Students will engage in scientific practice, make novel discoveries, participate in iterative work and collaborate with peers. The goal is to give students an authentic research experience in which they will acquire necessary skills for being a scientist. Four classroom hours and three hours of laboratory per week.

Requisite: BIOL 214 or BIOL 260. Limited to 18 students. Spring semester. Professor Kim.

2022-23: Offered in Spring 2023

311 Small Proteins in Antibiotic Resistance with Lab

This lab course will provide students a hands-on mentored research experience in protein biochemistry and molecular microbiology. Through structured primary literature analysis and studying historic examples, students will learn how to develop scientific questions, structure a convincing scientific argument, design experiments to address those questions, and critically analyze data and present findings to a scientific community. In addition to participation in classroom workshops and discussions, students will work in teams to design and execute a short research project in an area of active faculty research investigating a small protein involved in antibiotic resistance in the bacterial model organism Escherichia coli. Students will present the results of their work in a short research talk and in a poster session. At the end of this course, students will be able to apply the fundamental skills used to conduct scientific research.

Professor Wu Orr

2022-23: Offered in Spring 2023

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; BIOL 191 recommended. Limited to 30 students. Not open to first-year students. Spring semester. Professor Miller

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2024

330 Biochemical Principles of Life at the Molecular Level

(Offered as CHEM 330 and BIOL 330) What are the molecular underpinnings of processes central to life? We will explore the chemical and structural properties of biological molecules and learn the logic used by the cell to build complex structures from a few basic raw materials. Some of these complex structures have evolved to catalyze chemical reactions with an enormous degree of selectivity and specificity, and we seek to discover these enzymatic strategies. We will consider the detailed balance sheet that shows how living things harvest energy from their environment to fuel metabolic processes and to reproduce and grow. Examples of the exquisite control that permits a cell to be responsive and adapt its responses based on input from the environment will be considered. We will also consider some of the means by which cells respond to change and to stress. A student may not receive credit for both CHEM/BIOL 330 and BCBP/BIOL/CHEM 331. Note: BCBP/BIOL/CHEM 331 is a requirement for the biochemistry track of the BCBP major, so prospective BCBP majors should not enroll in CHEM/BIOL 330 if they are considering the biochemistry track of the major.

Requisite: BIOL 191 and CHEM  221. Limited to 40 students with 20 students per discussion section.

Spring Semester: Professor Bishop.

2022-23: Offered in Spring 2023
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Spring 2020, Spring 2021, Spring 2022, Spring 2024

331 Biochemistry with Lab

(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 CHEM/BIOL 330 and BCBP/BIOL/CHEM 331.

BCBP/BIOL/CHEM 331 is a requirement for the biochemistry track of the BCBP major, so prospective BCBP majors should not enroll in CHEM/BIOL 330 if they are considering the biochemistry track of the major.

Requisite: CHEM 231 and BIOL 191; or consent of the instructor. (CHEM 231 may be taken either as a prerequisite or as a co-requisite.) Limited to 30 students. Fall semester. Professors Jaswal and Wu Orr. 

2022-23: Offered in Fall 2022
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

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. 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.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2012, Fall 2013, Fall 2016, Fall 2022

371 Molecular Genetics with 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 30 students. Not open to first-year students. Omitted 2022-23. Visiting Professor Heussler.

2022-23: Not offered
Other years: Offered in Fall 2017, Fall 2018, Fall 2019, Spring 2020, Fall 2020, Fall 2021, Spring 2024

391 Quantitative Imaging with Lab: From Molecules to Cells and Beyond

(Offered as BIOL 391, BCBP 391, and NEUR-391) Fluorescence imaging offers a window to understanding the structure and function of living cells at sub-cellular resolution. This laboratory and lecture course will focus on advanced quantitative fluorescence microscopy techniques used for imaging a range of biological model systems and functions ranging from single molecules to cells to tissues. Students will learn the fundamentals of optics and microscopy and gain hands-on experience with classical and emerging imaging technology including: confocal microscopy, deconvolution and super-resolution methods, and live-cell imaging. Fluorescent probes and genetically encoded biosensors will be discussed and used to study dynamic cellular processes. In addition, the class will explore a range of image processing and analysis workflows. Lectures dealing with the theory, mechanics, and application of fluorescent imaging methods will be intermingled with extended laboratory sessions in our new Amherst Biological Imaging Center.

Pre requisites : BIOL 191 and PHYS 117 and one of BIOL/BCBP291, BIOL221 or BIOL/NEUR 301, or with instructor permission.

There will be two 80-minute lectures and a weekly laboratory. Limited to 12 students. Fall semester. Professors Edwards and Kim  

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2022, Spring 2024

400 Molecular and Cellular Biophysics

(Offered as PHYS 400, BIOL 400, BCBP 400, and CHEM 400) How do the physical laws that dominate our lives change at the small length and energy scales of individual molecules? What design principles break down at the sub-cellular level and what new chemistry and physics becomes important? We will answer these questions by looking at bio-molecules, cellular substructures, and control mechanisms that work effectively in the microscopic world. How can we understand both the static and dynamic shape of proteins using the laws of thermodynamics and kinetics? How has the basic understanding of the smallest molecular motor in the world, ATP synthase, changed our understanding of friction and torque? We will explore new technologies, such as atomic force and single molecule microscopy that have allowed research into these areas. This course will address topics in each of the three major divisions of Biophysics: bio-molecular structure, biophysical techniques, and biological mechanisms.

Requisite: CHEM 161/165, PHYS 116/123, PHYS 117/124, BIOL 191 or evidence of equivalent coverage in pre-collegiate courses. Spring semester. Professor Carter. 

2022-23: Offered in Spring 2023
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Spring 2015, Spring 2016, Spring 2017, Spring 2018, Spring 2019, Spring 2020, Spring 2021, Spring 2022, Spring 2024

424 Seminar in 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. Spring semester. Prof. Miller. 

2022-23: Offered in Spring 2023
Other years: Offered in Fall 2023

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. Fall 2022. Professor Temeles.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2011, Spring 2013, Fall 2014, Spring 2016, Spring 2019, Spring 2020, Spring 2021, Fall 2022

440, 441 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 close reading and discussion of articles from the primary literature, the course will explore key topics including overexploitation (including connections between the wildlife trade and emergent diseases such as COVID-19), habitat fragmentation, climate change, restoration, protected areas, payments for ecosystem services, as well as how to determine appropriate conservation priorities. Three classroom hours per week.

Requisite: BIOL 230/ENST 210 or BIOL 320, or consent of the instructor. Not open to first-year students. Limited to 14 students. Spring Semester. Senior Lecturer Levin.

2022-23: Offered in Spring 2023
Other years: Offered in Spring 2012, Fall 2013, Fall 2015, Fall 2016, Spring 2018, Fall 2019, Fall 2020, Spring 2022

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 BIOL-214, BIOL 260, BIOL 350/351, or consent of the instructor. Limited to 18 students. Not open to first-year students. Spring Semester. Professor Trapani.

2022-23: Not offered
Other years: Offered in Spring 2013, Spring 2017, Spring 2019, Spring 2022, Spring 2024

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 or ENST 200. 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. Omitted 2022-23. Professor Clotfelter and Senior Lecturer Levin.

2022-23: Not offered
Other years: Offered in Spring 2024

498, 499, 499D Senior Departmental Honors

Honors students take three courses of thesis research, usually, but not always, with the double course load in the spring (499D). The work consists of seminar programs, individual research projects, and preparation of a thesis on the research project.

Open to seniors. Fall semester. The Department.

2022-23: Offered in Fall 2022
Other years: Offered in Fall 2011, Fall 2012, Fall 2013, Fall 2014, Fall 2015, Fall 2016, Fall 2017, Fall 2018, Fall 2019, Fall 2020, Fall 2021, Fall 2022, Fall 2023

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