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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Suggested Citation:"A New Biology Curriculum 2." National Research Council. 2003. BIO2010: Transforming Undergraduate Education for Future Research Biologists. Washington, DC: The National Academies Press. doi: 10.17226/10497.
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Below is the uncorrected machine-read text of this chapter, intended to provide our own search engines and external engines with highly rich, chapter-representative searchable text of each book. Because it is UNCORRECTED material, please consider the following text as a useful but insufficient proxy for the authoritative book pages.

2 A New Biology Curriculum RECOMMENDATION #1 Given the profound changes in the nature of biology and how biological research is performed and communicated, each institution of higher education should reexamine its current courses and teaching approaches (as described in this report) to see if they meet the needs of today’s undergraduate biology stu- dents. Those selecting the new approaches should consider the importance of building a strong foundation in mathematics, physical, and information sci- ences to prepare students for research that is increasingly interdisciplinary in character. The implementation of new approaches should be accompanied by a parallel process of assessment, to verify that progress is being made toward the institutional goal of student learning. This chapter presents ideas for ways to enhance undergraduate educa- tion in biology. However, the committee recognizes that the specific ex- amples described here are only a subset of the many possible ways to in- crease interdisciplinary learning. The list of concepts that follow are lengthy. There is no way to incorporate all of this material into one or even several courses. The lists are presented as concepts that would be helpful to future biomedical researchers, if they were introduced at some point during a four-year undergraduate program. Many but not all would be helpful to other biology students who are focusing their studies on areas of life sci- ences such as population biology, plant biology, or cognitive science. These non-biomedical biologists would benefit from the increased attention to 27

28 BIO2010 biological concepts in their other science courses. All biology students should study some of the concepts in depth as undergraduates. The spe- cific concepts studied in detail by any individual student will depend on their interests, career goals, and the course offerings and course content available at their own school. Beyond the specific content of what they learn, students need hands-on experience with experimental inquiry and research starting early in their undergraduate careers. Their undergraduate experience should give them a sense of the power and beauty of science that takes full advantage of the richness of ideas and tools provided by a broad range of disciplines. The concepts are presented at the beginning of this chapter and poten- tial curricula at the end. The concepts are presented first so that faculty can consider how they might be incorporated into the courses offered. An evolutionary biologist teaching introductory biology will select different concepts from these lists than a developmental biologist teaching the same course. Either set of choices can improve interdisciplinary training of stu- dents and contribute to the creation of graduates who think more broadly. Ideally the changes will also help students see the connections between their different science courses and relate the topics to their own lives. Most biology students will not take such intensive schedules as presented in the sample curricula, and it is certainly possible to become a biomedical re- searcher without all of this background. However, the committee feels that future biomedical researchers, and possibly many other types of research- ers, would be better prepared to contribute to interdisciplinary break- throughs with such a background. Because of the striking advances in contemporary biology, those who plan to carry out biological research will need to access a broader range of concepts and skills than did past generations. The modern biologist uses a wide array of advanced techniques, ranging from special measuring instru- ments, novel imaging systems, computer methods, and quantitative ana- lytical tools and models. Understanding and effectively applying these tech- niques requires knowledge from outside of the biological sciences. Furthermore, the analysis of biological systems, with their web of complex interactions, will require the design of new theoretical approaches. To meet the challenges of the new biology, the committee believes that all future biological researchers will need concepts and skills drawn from a range of scientific disciplines that must be broader than what has been expected up to now. Because of biology’s great diversity, specific requirements will differ among the various subareas of biological research, and no one individual is

A NEW BIOLOGY CURRICULUM 29 expected to be equally competent in all the relevant areas of physics, chem- istry, mathematics, and engineering. Nevertheless, as a guide to the key biologically relevant ideas in these areas, and to stimulate discussion of what constitutes the core knowledge for the new biological curricula, the report begins by offering what is believed to be the central concepts of chemistry, physics, engineering, and mathematics that are most relevant to biology. Following these concepts are four examples of potential under- graduate biology curricula that would be appropriate for future biomedical researchers. These examples are not meant to discourage the use of alter- nate curricula that also cover the content of mathematics and physical and information sciences. Many of the courses listed have familiar titles in order to illustrate that many of the recommendations found in this report could be implemented through existing courses. However, the content of the courses would likely be altered to increase the integration of the differ- ent sciences. Throughout this report the committee uses the term “quantitative bi- ology” to refer to a biology in which mathematics and computing serve as essential tools in framing experimental questions, analyzing experimental data, generating models, and making predictions that can be tested. In quantitative biology, the multifaceted relationships between molecules, cells, organisms, species, and communities are characterized and compre- hended by finding structure in massive data sets that span different levels of biological organization. It is a science in which new computational, physi- cal, and chemical tools are sought and applied to gain a deeper and more coherent understanding of the biological world that has strong predictive power. Communicating how scientific advances and discoveries are made is a crucial part of undergraduate scientific education. First, exposure to the experimental and conceptual basis of key discoveries gives students a deeper understanding of scientific principles. Reading a classic paper can give stu- dents a sense of scientific inquiry at its best. Students can gain much by considering questions such as: What motivated the study? How were the experiments designed? What new experimental methods or analytical ap- proaches were needed? How surprising was the outcome? How did the discovery influence the future course of science? Second, by exploring how discoveries are made, students acquire an appreciation of the history and culture of science. Science becomes a human endeavor that spans time and space. Third, scientific discoveries are inspirational. They stimulate stu-

30 BIO2010 dents, demonstrate the importance of the prepared mind, and convey a sense of adventure and excitement. Scientific discoveries and how they were made can be communicated in many mutually reinforcing ways. First, lectures can be made more vivid and engaging by presenting carefully chosen exemplars of the process of discovery, such as Darwin’s finches, Mendel’s peas, Morgan’s flies, and McClintock’s maize. Roentgen’s discovery of x-rays, von Laue’s and the Braggs’ use of them to reveal atomic structure, and Watson and Crick’s reading of x-ray diffraction patterns in discovering the DNA double helix could be presented as a remarkable sequence of major scientific advances over more than a half century that led to the birth of a new biology. Sec- ond, many textbooks contain lucid accounts of the process of discovery that are interwoven with expositions of basic principles. Students should also be encouraged to read the full text of classic papers, which can be made accessible by posting them on the Web. Third, problem sets in- cluded in texts or written by instructors for their courses can be choice devices for exploring scientific advances that are inherently quantitative, such as the Hardy-Weinberg equilibrium and Shannon’s measure of infor- mation. Fourth, laboratory courses can motivate an experiment by recount- ing the historical background. For example, a biochemistry laboratory ex- periment on a glycolytic enzyme could begin with the Buchners’ discovery of fermentation in a cell-free yeast extract, a chemistry laboratory experi- ment on halogenation with Scheele’s discovery of chlorine, and a physics laboratory experiment on lasers with Einstein’s prediction of stimulated emission. Indeed, a classic discovery can be the basis of an extended ex- periment in which students explore new terrain, as in the use of the Hill reaction (light-induced electron transfer in illuminated chloroplasts), to find herbicides (an experiment in the interdisciplinary laboratory course described in Case Study #6). Noteworthy current advances should be presented along with classic discoveries. The covers of major journals often have striking images depict- ing important research findings. They can be used as evocative starting points in lectures and group discussions to motivate as well as inform stu- dents. For example, the recent discovery of fossils suggesting that the diver- gence between the human and chimpanzee lineages occurred earlier than previously thought (Brunet et al., 2002) would inform and enliven the teaching of human origins, especially if the paper were contrasted with previous estimates of the time of divergence based on molecular clocks.

A NEW BIOLOGY CURRICULUM 31 Future research biologists should also be exposed to scientific controversies and their resolution. CONCEPTS AND SKILLS FOR THE NEW CURRICULUM The concepts presented in this chapter are the end result of the long study process described in Chapter 1. Initially the committee examined the requirements for biology majors at 12 institutions of various types around the country. They compared this information to the requirements for biology majors at their own college or university and discussed some of the similarities and differences. The committee also discussed the desired characteristics for the invited experts who would participate in the panels on Chemistry, Physics and Engineering, and Mathematics and Computer Science. They selected faculty members who covered the subdisciplines within each panel’s charge, and those who are known for their teaching. The following lists of concepts owe much to the ideas shared by the panel members during their respective meetings. Each panel approached its task from a different perspective, and hence created slightly different types of recommendations. The panel members considered the way their discipline is currently taught to biology students, at their own institution and others with which they are familiar. In assembling their recommendations, they considered the course requirements, the content of those courses, the con- tent that is most relevant to biology students, and to some degree the way in which the material is taught (lectures, seminars, laboratories). The com- mittee as a whole went through a similar process to create the list of biology concepts presented below. In preparing the final concept lists for the re- port, the committee has attempted to structure the lists in a way that stresses their pertinence to interdisciplinary research and education. In addition to the concepts presented on the following lists, the com- mittee recognizes that future biologists, and indeed all future workers and citizens, will also need more general skills. Science faculty are not required to leave the teaching of reading, writing, critical thinking, and communica- tion skills solely to the humanities and social sciences faculty. For example, incorporating the writing of grant proposals, or the scientific component of a business proposal for a biotech start-up, into a course provides useful experience requiring knowledge of both scientific ideas and other skills. These types of activities also provide an opportunity for students to con- sider the interplay between scientific discovery and society, including the importance of the scientific method and scientific ethics.

32 BIO2010 Biology RECOMMENDATION #1.1 Understanding the unity and diversity of life requires mastery of a set of fundamental concepts. This understanding will be greatly enhanced if biology courses build on material begun in other science courses to expose students to the ideas of modeling and analyzing biological and other systems. Biological systems show remarkable unity at the molecular and cellular levels, reflecting their common ancestry. Variations on this unity lead to the extraordinary diversity of individual organisms. In order for biology stu- dents to understand the unifying features of the biological concepts listed below, the concepts must be taught in multiple contexts. Biology faculty should consider the various points in their courses at which the concepts will fit. They should also consider the concept lists for chemistry, physics, and mathematics that follow and the ways in which those ideas could be incorporated into biology courses. In order for biology students to receive a truly interdisciplinary education, cooperation between departments will be necessary. It is the responsibility of the biology faculty to make active outreach efforts to colleagues in other departments by offering to work together on mechanisms for incorporating biological concepts and examples into non-biology courses. Concepts of Biology Central Themes • All living things have evolved from a common ancestor, through processes that include natural selection and genetic drift acting on heritable genetic variation. • Biological systems obey the laws of chemistry and physics. • Structural complexity and information content are built up by com- bining simpler subunits into multiple complex combinations. • Understanding biological systems requires both reductionist and holistic thinking because novel properties emerge as simpler units assemble into more complex structures. • Living systems are far from equilibrium. They utilize energy, largely derived from photosynthesis, which is stored in high-energy bonds or ionic concentration gradients. The release of this energy is coupled to thermody- namically unfavorable reactions to drive biological processes.

A NEW BIOLOGY CURRICULUM 33 • Although fundamental molecular and cellular processes are con- served, biological systems and organisms are extraordinarily diverse. Unlike atoms and simple molecules studied in chemistry and physics, no two cells are identical. • Biological systems maintain homeostasis by the action of complex regulatory systems. These are often networks of interconnecting partially redundant systems to make them stable to internal or external changes. • Cells are fundamental units of living systems. Three fundamental cell types have evolved: bacteria, archea, and eukaryotes. • Living organisms have behavior, which can be altered by experience in many species. • Information encoded in DNA is organized into genes. These heri- table units use RNA as informational intermediates to encode protein se- quences, which become functional on folding into distinctive three-dimen- sional structures. In some situations RNA itself has catalytic activity. • Most biological processes are controlled by multiple proteins, which assemble into modular units to carry out and coordinate complex func- tions. • Lipids assemble with proteins to form membranes, which surround cells to separate them from their environment. Membranes also form dis- tinct compartments within eukaryotic cells. • Communication networks within and between cells, and between organisms, enable multicellular organisms to coordinate development and function. • In multicellular organisms, cells divide and differentiate to form tissues, organs, and organ systems with distinct functions. These differ- ences arise primarily from changes in gene expression. • Many diseases arise from disruption of cellular communication and coordination by infection, mutation, chemical insult, or trauma. • Groups of organisms exist as species, which include interbreeding populations sharing a gene pool. • Populations of species interact with one another and the environ- ment to form interdependent ecosystems with flow of energy and materials between multiple levels. • Humans, as well as many other species, are members of multiple ecosystems. They have the capacity to disrupt or preserve the ecosystems upon which they depend.

34 BIO2010 Chemistry RECOMMENDATION #1.2 The committee recommends that biology majors receive a thorough educa- tion in chemistry, including general chemistry and aspects of organic chemistry, physical chemistry, analytical chemistry, and biochemistry, incorporated into a new course or courses. They should master the chemistry concepts listed below. Biology faculty should work in concert with their chemistry colleagues to help design chemistry curricula that will not only foster growth of aspiring chemists, but also stimulate biology majors as well as students majoring in other disci- plines. Furthermore, chemistry faculty must work with biologists to find ways to collaborate on the incorporation of chemistry concepts, and those of other scien- tific disciplines, into their teaching of biology. Learning biology should not be dependent upon chemistry but, rather, integrated with it. Biology students should begin their study of chemistry in the first year so that they will acquire a strong foundation in chemistry before starting their study of chemically based aspects of biology. Chemistry has always been an important sister science to biology, espe- cially to biochemistry and medicine. Today, modern molecular biology and cell biology focus on understanding the chemistry of genes and of cell struc- ture. In the applied area, for example, chemistry is central to modern agriculture. Biomedical engineering draws on chemistry for new materials. It is evident that future research biologists will need to have a thorough grounding in chemistry to make their research possible and to understand the work of others. Such a grounding in general chemistry and organic chemistry has historically required at least three semesters of chemistry courses, but could require fewer following an integrated restructuring. There are many combinations of courses that would allow students to learn these chemical concepts. In the traditional program, a full year of general chemistry is followed by a full year of organic chemistry, and then by physi- cal chemistry. Regardless of when it is taught, organic chemistry should include ma- terial on the principal biomolecules, including heterocyclic chemistry and the chemistry of phosphate esters. The role of these biomolecules in biol- ogy is so important that they should not be omitted, as too frequently occurs. Furthermore, including a description of the biochemical versions of displacement reactions, aldol and Claisen condensations, and free radical reactions will add interest for all students, not just biologists.

A NEW BIOLOGY CURRICULUM 35 Concepts of Chemistry Atoms • Periodic table, trends (size, electronic properties, isoelectronic sys- tems) • Orbitals and electronic configuration • Nuclear chemistry Molecules • Lewis structures • Molecular properties (shape, dipole moments, bond energies) • Bonding models (valence bond theory, molecular orbital theory) • Molecular interactions (ion pair, hydrogen bond, van der Waals) • Metal ions and metal complexes • Resonance and electron delocalization • Computational methods and modeling Water and Aqueous Solutions • Structure and polarity of liquid water • Ionic compounds in aqueous solutions • Acid-base equilibria, pH, pK, indicators • Hydrophobic effect Chemical Reactions • Stoichiometry • Hydrocarbons, heterocycles, and functional groups • Reaction types (acid-base, redox, addition, elimination, substitu- tion) • Reactive intermediates: carbocations, carbanions, enols, enolates, free radicals • Mechanisms of selected classes of chemical reactions Energetics and Equilibria • Enthalpy, entropy, and free energy • Equilibrium constant • Temperature dependence of equilibria • Electrochemistry, electrochemical cells, Nernst equation • Boltzmann distribution

36 BIO2010 Reaction Kinetics • Reaction rate laws and kinetic order • Transition states • Temperature dependence of kinetics • Catalysis, enzyme-catalyzed reactions, and the Michaelis-Menten equation • Diffusion-limited reactions • Thermodynamic versus kinetic stability Biomolecules • Building blocks: amino acids, nucleotides, carbohydrates, fatty acids • Biopolymers: proteins, nucleic acids, polysaccharides • Three-dimensional structure of biological macromolecules • Molecular assemblies: micelles, monolayers, biological membranes • Solid-phase synthesis of oligonucleotides and peptides • Combinatorial synthesis • Spectroscopic reporters Analyzing Molecules and Reactions • Mass spectrometry • Absorption and emission spectroscopy (UV, visible, infrared) • NMR spectroscopy • Diffraction (x-ray, neutron, electron) • Electron microscopy and scanning probe microscopy • Separation methods: chromatography, electrophoresis, and centrifu- gation • Isotopic tracers and radioactivity Materials • Metals • Properties and synthesis of polymers • Conductors, insulators, and semiconductors A list of questions useful in teaching these concepts is presented in Appendix D.

A NEW BIOLOGY CURRICULUM 37 Physics RECOMMENDATION #1.3 The principles of physics are central to the understanding of biological pro- cesses, and are increasingly important in sophisticated measurements in biology. The committee recommends that life science majors master the key physics con- cepts listed below. Experience with these principles provides a simple context in which to learn the relationship between observations and mathematical de- scription and modeling. The typical calculus-based introductory physics course taught today was designed to serve the needs of physics, mathematics, and engineering students. It allocates a major block of time to electromagnetic theory and to many details of classical mechanics. In so doing, it does not provide the time needed for in-depth descriptions of the equally basic physics on which students can build an understanding of biology. By emphasizing exactly solvable problems, the course rarely illustrates the ways that physics can be applied to more recalcitrant problems. Illustrations involving modern bi- ology are rarely given, and computer simulations are usually absent. Collec- tive behaviors and systems far from equilibrium are not a traditional part of introductory physics. However, the whole notion of emergent behavior, pattern formation, and dynamical networks is so central to understanding biology, where it occurs in an extremely complex context, that it should be introduced first in physical systems, where all interactions and parameters can be clearly specified, and quantitative study is possible. Concepts of Physics Motion, Dynamics, and Force Laws • Measurement: physical quantities, units, time/length/mass, preci- sion • Equations of motion: position, velocity, acceleration, motion under gravity • Newton’s laws: force, mass, acceleration, springs and related mate- rial: stiffness, damping, exponential decay, harmonic motion • Gravitational and spring potential energy, kinetic energy, power, heat from dissipation, work • Electrostatic forces, charge, conductors/insulators, Coulomb’s law • Electric potential, current, units, Ohm’s law

38 BIO2010 • Capacitors, R and RC circuits • Magnetic forces and magnetic fields • Magnetic induction and induced currents Conservation Laws and Gobal Constraints • Conservation of energy and momentum • Conservation of charge • First and Second Laws of thermodynamics Thermal Processes at the Molecular Level • Thermal motions: Brownian motion, thermal force (collisions), temperature, equilibrium • Boltzmann’s law, kT, examples • Ideal gas statistical concepts using Boltzmann’s law, pressure • Diffusion limited dynamics, population dynamics Waves, Light, Optics, and Imaging • Oscillators and waves • Geometrical optics: rays, lenses, mirrors • Optical instruments: microscopes and microscopy • Physical optics: interference and diffraction • X-ray scattering and structure determination • Particle in a box; energy levels; spectroscopy from a quantum view- point • Other microscopies: electron, scanning tunneling, atomic force Collective Behaviors and Systems far from Equilibrium • Liquids, laminar flow, viscosity, turbulence • Phase transitions, pattern formation, and symmetry breaking • Dynamical networks: electrical, neural, chemical, genetic Engineering RECOMMENDATION #1.4 The committee recommends that life science majors be exposed to engineer- ing principles and analysis. This does not necessarily require that they take a course in a school of engineering; courses in physics, biology, and other depart- ments can provide exposure to these concepts. Students should have the opportu- nity to participate in laboratories that give them hands-on experience, so that

A NEW BIOLOGY CURRICULUM 39 they may learn about the functioning of complex systems, especially as they relate to the basic principles of physical science, mathematics, and modeling. Basic courses in physics and engineering should be developed specifically for life sci- ences students; these courses could be taught in either the physics or the biology department. This could be complemented exceptionally well by biology lecture or laboratory courses that assist students in their understanding of principles of physics and engineering (e.g., a unit on biomechanics taught in a physiology or anatomy course). Biology increasingly involves the analysis of complex systems. Under- standing function at the systems level requires a way of thinking that is common to engineers. Creating (or re-creating) function by building a com- plex system and getting it to work provides compelling proof that the scien- tist understands the essential building blocks and how they work in syn- chrony. Organisms can be analyzed in terms of subsystems having particular functions. To understand system function in biology in a predictive and quantitative fashion, it is necessary to describe and model how the system function results from the properties of its constituent elements. One ap- proach to the study of biology is as a problem in reverse engineering. Manu- factured systems are easier to understand than biological systems, because they have no unknown components, and their design principles can be explicitly stated. It is easiest to learn how to analyze systems through inves- tigating how manufactured systems achieve their designed purpose, how their function depends on properties of their components, and how func- tion can be reliable even with imperfect components. As an example, a quantitative understanding of a cell-signaling chemical network involves the concepts of negative feedback, gain, signal-to-noise, bandwidth, and cross-talk. These concepts are simple to experience in the context of how an electrical amplifier can be built from components. Similarly, an effort to understand the locomotion of insects might be preceded by a laboratory involving an analysis of a simple legged robot. In such a system, the de- scription of the muscles (activators) and control signals is completely known, and the relation between the laws of physics and the problem of controlling directed movements can be seen clearly. Examples of Engineering Topics Suitable for Inclusion in a Biology Cur- riculum • The blood circulatory system and its control; fluid dynamics; pres- sure and force balance.

40 BIO2010 • Swimming, flying, walking, dynamical description, energy require- ments, actuators, control. Material properties of biological systems and how their structure relates to their function (e.g., wood, hair, cell membranes, cartilage). • Long range neuron signals; physical necessity of repeaters (e.g., nodes of Ranvier), engineering advantage of pulse coding, action potential generation, information transmission and errors. • Shapes of cells: force balance, hydrostatic pressure, elasticity of membrane and effects of the spatial dependence of elasticity; cytoskeletal force effects on shape. One such effort illustrates the interactions of the engineering and sci- ence involved, and makes it clear that the subject can be examined in enough detail to teach essential ideas honestly. A “long range neural sig- nals” section might begin with the electrical conductivity of salt water, of the lipid cell membrane, and the electrical capacitance of the cell mem- brane. It would next develop the simple equations for the attenuation of a voltage applied across the membrane at one end of an axon “cylinder” with distance down the axon, and the effect of membrane capacitance on signal dynamics for time-varying signals. After substituting numbers, it becomes clear that amplifiers will be essential. Real systems are always noisy and imperfect; amplifiers have limited dynamical range; and the combination of these facts makes sending of an analog voltage signal through a large number of amplifiers essentially impossible. Pulse coding information es- capes that problem (all long distance communication is digital these days). How are “pulses” generated by a cell? This would lead to the power supply needed by an amplifier—ion pumps, and the Nernst potential. How are action potentials generated? A first example of the transduction of an ana- log quantity into pulses might be stick-slip fraction, in which a block rest- ing on a table and pulled by a weak spring whose end is steadily moved, moves in “jumps” whose distance is always the same. This introduction to nonlinear dynamics contains the essence of how an action potential is gen- erated. The “negative resistance” of the sodium channels in a neuron mem- brane provides the same kind of “breakdown” phenomenon. Stability and instabilities (static and dynamic) of nonlinear dynamical systems can be analyzed, and finally the Hodgkin Huxley equations illustrated. The mate- rial is an excellent source of imaginative laboratories involving electrical measurements, circuits, dynamical systems, batteries and the Nernst poten- tial, and information and noise, and classical mechanics. It has great po-

A NEW BIOLOGY CURRICULUM 41 tential for simulations of systems a little too complicated for complete math- ematical analysis, and thus is ideal for teaching simulation as a tool for understanding. Many topics in biology interact with the engineering viewpoint in such a fashion. Mathematics and Computer Science RECOMMENDATION #1.5 Quantitative analysis, modeling, and prediction play increasingly signifi- cant day-to-day roles in today’s biomedical research. To prepare for this sea change in activities, biology majors headed for research careers need to be edu- cated in a more quantitative manner, and such quantitative education may require the development of new types of courses. The committee recommends that all biology majors master the concepts listed below. In addition, the com- mittee recommends that life science majors become sufficiently familiar with the elements of programming to carry out simulations of physiological, ecological, and evolutionary processes. They should be adept at using computers to acquire and process data, carry out statistical characterization of the data and perform statistical tests, and graphically display data in a variety of representations. Furthermore, students should also become skilled at using the Internet to carry out literature searches, locate published articles, and access major databases. The elucidation of the sequence of the human genome has opened new vistas and has highlighted the increasing importance of mathematics and computer science in biology. The intense interest in genetic, metabolic, and neural networks reflects the need of biologists to view and understand the coordinated activities of large numbers of components of the complex systems underlying life. Biology students should be prepared to carry out in silico (computer) experiments to complement in vitro and in vivo experi- ments. It is essential that biology undergraduates become quantitatively literate. The concepts of rate of change, modeling, equilibria and stability, structure of a system, interactions among components, data and measure- ment, visualizing, and algorithms are among those most important to the curriculum. Every student should acquire the ability to analyze issues aris- ing in these contexts in some depth, using analytical methods (e.g., pencil and paper), appropriate computational tools, or both. The course of study would include aspects of probability, statistics, discrete models, linear alge- bra, calculus and differential equations, modeling, and programming.

42 BIO2010 Concepts of Mathematics and Computer Science Calculus • Complex numbers • Functions • Limits • Continuity • The integral • The derivative and linearization • Elementary functions • Fourier series • Multidimensional calculus: linear approximations, integration over multiple variables Linear Algebra • Scalars, vectors, matrices • Linear transformations • Eeigenvalues and eigenvectors • Invariant subspaces Dynamical Systems • Continuous time dynamics—equations of motion and their trajec- tories • Test points, limit cycles, and stability around them • Phase plane analysis • Cooperativity, positive feedback, and negative feedback • Multistability • Discrete time dynamics — mappings, stable points, and stable cycles • Sensitivity to initial conditions and chaos Probability and Statistics • Probability distributions • Random numbers and stochastic processes • Covariation, correlation, and independence • Error likelihood Information and Computation • Algorithms (with examples) • Computability

A NEW BIOLOGY CURRICULUM 43 • Optimization in mathematics and computation • ”Bits”: information and mutual information Data Structures • Metrics: generalized ‘distance’ and sequence comparisons • Clustering • Tree-relationships • Graphics: visualizing and displaying data and models for concep- tual understanding Additional Quantitative Principles Useful to Biology Students Rate of Change • This can be a specific (e.g., per capita) rate of change or a total rate of change of some system component. • Discrete rates of change arise in difference equations, which have associated with them an inherent time-scale. • Continuous rates of change arise as derivatives or partial derivatives, representing instantaneous (relative to the units in which time is scaled) rates. Modeling • The process of abstracting certain aspects of reality to include in the simplifications of reality we call models. • Scale (spatial and temporal)—different questions arise on different scales. • What is included (system variables) depends on the questions ad- dressed, as does the hierarchical level in which the problem is framed (e.g., molecular, cellular, organismal). • There are trade-offs in modeling—no one model can address all questions. These trade-offs are between generality, precision, and realism. • Evaluating models depends in part on the purpose for which the model was constructed. Models oriented toward prediction of spe- cific phenomena may require formal statistical validation methods, while models that wish to elucidate general patterns of system re- sponse may require corroboration with the available observed pat- terns.

44 BIO2010 Equilibria and Stability • Equilibria arise when a process (or several processes) rate of change is zero. • There can be more than one equilibrium. Multiple stable states (e.g., long-term patterns that are returned to following a perturbation of the system) are typical of biological systems. The system dynamics may drive the process to any of these depending on initial condi- tions and history (e.g., the order of any sequence of changes in the system may affect the outcomes). • Equilibria can be dynamic, so that a periodic pattern of system re- sponse may arise. This period pattern may be stable in that for some range of initial conditions, the system approaches this period pat- tern. • There are numerous notions of stability, including not just whether a system that is perturbed from an equilibrium returns to it, but also how the system returns (e.g., how rapidly it does so). • Modifying some system components can lead to destabilization of a previously stable equilibrium, possibly generating entirely new equi- libria with differing stability characteristics. These bifurcations of equilibria arise in many nonlinear systems typical in biology. Structure • Grouping components of a system affects the kinds of questions addressed and the data required to parameterize the system. • Choosing different aggregated formulations (by sex, age, size, physi- ological state, activity state) can expand or limit the questions that can be addressed, and data availability can limit the ability to inves- tigate effects of structure. • Geometry of the aggregation can affect the resulting formulation. • Symmetry can be useful in many biological contexts to reduce the complexity of the problem, and situations in which symmetry is lost (symmetry-breaking) can aid in understanding system response. Interactions • There are relatively few ways for system components to interact. Negative feedbacks arise through competitive and predator-prey type interactions, positive feedback through mutualistic or com- mensal ones. • Some general properties can be derived based upon these (e.g., two-

A NEW BIOLOGY CURRICULUM 45 species competitive interactions), but even relatively few interacting system components can lead to complex dynamics. • Though ultimately everything is hitched to everything else, signifi- cant effects are not automatically transferred through a connected system of interacting components—locality can matter. • Sequences of interactions can determine outcomes—program order matters. Data and Measurement • Only a few basic data types arise (numeric, ordinal, categorical), but these will often be interconnected and expanded (e.g., as vectors or arrays). • Consistency of the units with which one measures a system is im- portant. • A variety of statistical methods exist to characterize single data sets and to make comparisons between data sets. Using such methods with discernment takes practice. Stochasticity • In a stochastic process, individual outcomes cannot be predicted with certainty. Rather, these outcomes are determined randomly according to a probability distribution that arises from the underly- ing mechanisms of the process. Probabilities for measurements that are continuous (height, weight, etc.), and those that are discrete (sex, cell type) arise in many biological contexts. • Risk can be identified and estimated. • There are ways to determine if an experimental result is significant. • There are instances when stochasticity is significant and averages are not sufficient. Visualizing • There are diverse methods to display data. • Simple line and bar graphs are often not sufficient. • Nonlinear transformations can yield new insights. Algorithms • These are rules that determine the types of interactions in a system, how decisions are made, and the time course of system response. • These can be thought of as a sequence of actions similar to a com-

46 BIO2010 puter program, with all the associated options such as assignments, if-then loops, and while-loops. Using Computers Many of the concepts above deal with types of analysis and modeling that require knowledge of computer programming. However, there is an- other aspect of computing that is important for the future research biolo- gist: the use of computers as tools. Computer use is a fact of life for all modern life scientists. Exposure during the early years of their undergradu- ate careers will help life science students use current computer methods and learn how to exploit emerging computer technologies as they arise. As com- puter power continues to grow rapidly, applications that were available only on supercomputers a few years ago can now be used on relatively inexpen- sive personal computers. Computers are essential today for obtaining infor- mation from databases (e.g., genetic data from Genbank), establishing rela- tionships (e.g., using the BLAST algorithm to quantitate the similarity of a given DNA or protein sequence to all known sequences), deducing pat- terns (e.g., clustering genes that are regulated in concert), carrying out sta- tistical tests, preparing plots and other graphics for presentation, and writ- ing manuscripts for publication. Furthermore, computers are playing a central role in the laboratory in controlling equipment, obtaining data from measuring devices, and carrying out real-time analysis (e.g., image acquisi- tion in confocal fluorescence microscopy). Research biologists are increas- ingly acquiring and analyzing vast amounts of data (e.g., the degree of expression of tens of thousands of genes in multiple cellular states). They will need to be conversant with new theoretical and modeling approaches to come to grips with the interplay of many simultaneously interacting components of complex systems. Many analyses of biological data can be accomplished with existing programs (e.g., BLAST). However, being able to modify or construct ap- plications is necessary in many research areas. Learning how a computer application is developed provides students with insight into the software they use. Computer understanding can be taught by providing experiences in computer programming, teaching about computer algorithms, and how to construct simple simulations. This familiarity could be accomplished by exposing students to programming in higher-level languages such as Matlab, Perl, or C. The Internet is increasingly becoming the primary source of informa- tion for life scientists. Databases in a variety of areas (e.g., genomics, global

A NEW BIOLOGY CURRICULUM 47 warming, population dynamics) provide integrative frameworks that are valuable for addressing important biological issues. Becoming fully con- versant with databases such as the National Center for Biotechnology In- formation (NCBI) is important for all biology majors. NCBI’s mission is to develop new information technologies to aid in the understanding of fun- damental molecular and genetic processes that control health and disease. Searchable databases at NCBI’s Web site (http://www.ncbi.nlm.nih.gov) include Genbank (all publicly available DNA sequences), PubMed (access to more than 11 million Medline citations of biomedical literature, includ- ing links to full text articles), BLAST (Basic Local Alignment Search Tool for carrying out similarity searches of DNA or protein query sequences), Taxonomy (a wide range of taxonomic information at the molecular level), and Structure (database of three-dimensional structure of biological macro- molecules and tools for visualization and comparative analysis). Major model organism databases such as Fly Base (www.flybase.org) are useful, and The Interactive Fly (http://sdb.bio.purdue.edu/fly/aimain/laahome.htm) is a related learning tool. Sites such as PubMed are essential for searching the literature and valu- able for linking to full-text publications. Students should learn how to ob- tain different kinds of information from Web sites (e.g., DNA and protein sequences, atomic coordinates, phylogenetic relationships, functional anatomy, and biogeographic ecosystem data) and how to make informa- tion available to others over the Web (e.g., depositing new DNA sequences in Genbank). In addition, students should learn about mechanisms (e.g., peer review) of evaluating and increasing the reliability of information ob- tained on the Web. Students should have experience operating lab equipment controlled by computer, and observe or attempt modification of the settings or the programming to fit the needs of the experiment. This type of experience is important for demonstrating that biological research is not constrained to the use of preexisting applications and materials. New approaches and equipment are developed regularly. DESIGNING NEW CURRICULA SUITABLE FOR VARIOUS TYPES OF INSTITUTIONS RECOMMENDATION #2 Concepts, examples, and techniques from mathematics, and the physical and information sciences should be included in biology courses, and biological concepts and examples should be included in other science courses. Faculty in

48 BIO2010 biology, mathematics, and physical sciences must work collaboratively to find ways of integrating mathematics and physical sciences into life science courses as well as providing avenues for incorporating life science examples that reflect the emerging nature of the discipline into courses taught in mathematics and physi- cal sciences. Suggestions are provided here for integrating physical science and mathematics more fully into a biology education. Each institution will need to evaluate these recommendations in light of its own particular cir- cumstances. Decisions will be influenced by many factors, including the size and expertise of the faculty, number of life science majors, and num- ber of students from other science majors enrolled in biology courses. Con- sideration will also need to be given to the available resources, cooperation from other departments and the administration, and the need for curricu- lar change to keep up with the dynamic growth of the discipline of biol- ogy. Regardless of individual circumstances, all institutions are capable of beginning the process of change by adding interdisciplinary examples to existing courses in relevant disciplines to emphasize the integrative nature of the biological sciences with mathematics and physical science. Chapter 3 presents case studies and ideas for courses that promote interdisciplinary learning. The courses required of a biology major today typically consist of one year of physics, with lab; 2.5 years of chemistry, some with labs; some calculus and possibly some statistics; and a variety of biology courses. The remainder of undergraduate courses would be in disciplines outside of the sciences. A study of the “core” or required biology courses for undergradu- ate biology majors was carried out by Dominick Marocco . He states that required courses reveal “the consensus of the faculty at an institution that the subject matter of the core is central to the education of a biologist.” He concludes that a consensus core based on the requirements at the 104 schools surveyed would include genetics, biochemistry, cell biology, micro- biology, evolution/ecology, and a seminar. Another major impact on today’s curriculum are requirements for admission to medical school. This issue is discussed further in Recommendation #7, found in Chapter 6. The physical sciences and mathematics background of biology majors can best be strengthened by integrated teaching rather than by the addition of courses taught in isolation of biology. Though all of the topics found on the concept lists are offered in most universities and colleges, it is difficult for life science students to master the essential ones without taking a larger

A NEW BIOLOGY CURRICULUM 49 number of courses than can be accommodated in a biology major. Hence, the committee recommends the creation of new courses (or revamping of old courses) to cover the most pertinent part of this material in less time and with examples geared toward biology. Furthermore, as with key con- cepts in the physical sciences that are relevant for the study of biological systems, biology faculty can further enhance students’ understanding of the connections between mathematics, computer science, and biology by in- troducing these concepts into courses in the biology curriculum. Relevant courses might be taught by faculty from mathematics, computer science, or biology, or by a collaborating team of faculty from multiple departments. Outside input should be sought if the course is to be taught by a biologist who does not have extensive interdisciplinary experience. A mathemati- cian or computer scientist might also be invited to give a guest lecture or two. Similarly, biologists should provide assistance to the mathematics and computer science faculty in designing biological examples for use in their courses. One aspect of reform is the reevaluation of the topics covered in intro- ductory courses. Is some material covered just because it is in the textbook or has “traditionally” been taught in this course? Are there other topics that would be more useful or more relevant or interesting to the students cur- rently enrolled in the course? By adding modules and redesigning courses, a department can make its curriculum more interdisciplinary without any increase in the number of courses required. The order in which the material is taught should be carefully consid- ered in relation to the rest of the curriculum. For example, the early intro- duction of statistics and discrete mathematics could be beneficial for biol- ogy courses. This is the type of change that should be assessed after implementation to see if it is beneficial to student learning. While a sub- stantial part of the material in the concept lists can be taught as mathemat- ics, chemistry, or physics (with biological examples), some of the more advanced and more specifically biological material might instead be cov- ered in a biology course or an interdepartmental course, depending on the teaching resources and interests of the particular departments. For example, a course on modeling could be taught in many different departments, or modules on modeling could be added to preexisting courses. Those biology students who wish to eventually work at the interface of biology and physi- cal, mathematical or information sciences will need to become more expert in those fields, and may want to take some of the standard courses offered in those disciplines that provide a more rigorous foundation. The integra-

50 BIO2010 tion of disciplines may also be well served through the development of an interdisciplinary concentration in mathematics or physics, so that biology and other faculty and departments can work more closely together, through shared resources and curriculum, to develop and maintain a program that is best tailored to address student needs. In the traditional program, a full year of general chemistry is followed by a full year of organic chemistry, and then by physical chemistry. Some institutions are now adopting nontraditional plans, in which organic chem- istry is taught earlier. Several have experimented with organic chemistry as the first course; for biology students, the advantage is they can start study- ing biochemistry in their second year with the chemical background needed to understand it. Earlier knowledge of biochemistry is useful in many biology courses, ranging from genetics to development. Another way to allow students to learn biochemistry earlier is to restructure the introduc- tory chemistry course so that only one semester is required before students begin organic chemistry. This plan is well suited to biology majors who can take both general chemistry and half of organic chemistry in their first year, preparing them for chemistry-based biology in their second year. One-se- mester courses to follow organic chemistry could include concepts of physi- cal chemistry, perhaps focusing on solution chemistry; an introduction to analytical chemistry; or biochemistry at a chemically sophisticated level (i.e., where biomolecular structure and reaction mechanisms are presented in considerable depth). Relevant biological examples should be part of these courses, and indeed part of the organic and general chemistry courses as well. Restructuring chemistry courses along these lines would be compatible with the needs of physicists, geologists, and nonchemical engineers who often need to take one year of chemistry. A yearlong course covering both inorganic and organic chemistry would also be useful for humanities and social science students seeking an overview of chemistry to meet their sci- ence requirements. It would be more demanding than many of the courses currently offered to nonscience majors, but potentially more appealing be- cause of its increased use of applied examples that students are more easily able to relate to their own lives and surroundings. A first semester of or- ganic chemistry, given in the spring, could include a general survey of the properties of the major classes of organic compounds and their key reac- tions, so those students not going further in chemistry would still have a reasonable picture of the subject. A second semester of organic chemistry, given in the fall of the second year for chemists, biologists, and chemical

A NEW BIOLOGY CURRICULUM 51 engineers, could then be a more advanced treatment, with more informa- tion on mechanism and synthesis than in the first semester. The typical two-semester introductory physics course with calculus, which has changed rather little over more than a quarter-century, is often the only option for a biology student who wants a strong physics prepara- tion. One way to teach the material on the physics concept list, described earlier in the chapter, would be as a three-semester sequence. However, there are other ways that such material could be covered. For example, the more conventional physics topics might be covered by a one-year course within a physics department while the other materials (which more specifi- cally bridge biology and physics) might then be part of another course, in either the physics or biology department; in fact, some of it is appropriate for a physical chemistry course. The choice of department and number of semesters would vary from institution to institution, and depend to some degree on the expertise of the faculty in each department. Alternatively the material could be taught as an interdepartmental course. While all the topics listed have direct relevance to biology, the emphasis in course design should be on learning and developing the relationship between observa- tions and mathematical description and modeling, rather than on slavishly covering every topic. An attractive option for quantitative literacy, mathematics, and com- puter science at some institutions might be the development of an inte- grated course to teach quantitative approaches and tools for research, as has been successfully developed at the University of Tennessee (see Case Study #4.) This innovative two-semester course designed for life science majors replaces the traditional calculus course. It introduces topics such as the mathematics of discrete variables, linear algebra, statistics, programming, and modeling early in the course, to provide completely new material for well-prepared students. These topics are then connected to applied aspects of calculus. It should be noted that this course makes extensive use of graduate students in Tennessee’s mathematical and computational ecology program. These graduate students are well positioned to explain the con- nections between mathematics and biology. A two-semester quantitative course such as the one at Tennessee ex- poses students to many mathematical ideas but is too brief to provide much depth in many of them. A more intensive alternative would be a four- semester series. Two semesters could deal with calculus (single and multi- variate), quantitative differential equations (including phase plane analy- sis), and the relevant elementary linear algebra, taught in the context of

52 BIO2010 biological applications. A third semester might be on biostatistics, empha- sizing different ways to analyze and interpret data. A fourth semester could include discrete math and algorithms and could be taught in the context of biological issues, including those arising in genomics. In summary, for the future biomedical researcher, the committee pro- poses: • A reorganization of the chemistry offerings to allow for the early presentation of organic chemistry and the addition of some analytical and physical chemistry to the organic and inorganic courses. One potential ar- rangement of courses would be for students to start with a one-semester introductory inorganic course (rather than the two currently taught at many institutions), followed by two semesters of organic, one (or two) of bio- chemistry and then a combined physical and analytical course. • An expansion of the physics offerings to include a third semester that incorporates engineering principles into the syllabus in order to assist students in becoming familiar with modeling and analysis of biological and other systems. Other topics might include molecular physics, biospectroscopies, and dynamical networks. • A new mathematics sequence that exposes students to statistics, probability, discrete math, linear algebra, calculus, and modeling without requiring that a full semester be spent on each topic. A brief overview of these topics could be presented in two semesters, but a full introduction and the inclusion of more computer science would more likely take four semesters. Potential Curricula Four quite different examples of a modernized four-year curriculum for a biology major are presented below to stimulate discussion among faculty. These tables represent various course options a student might take. They do not represent proposed requirements for a major. At first glance the courses in the tables may not look so different from the current offer- ings at some colleges. The idea here is to incorporate some of the concepts presented earlier in the chapter into each of these science courses. Another change from the current practice at some universities would be the in- creased incorporation of teaching techniques such as inquiry-based learn- ing and approaches such as those presented in the next two chapters. Many institutions would need to revamp their course offerings in order to allow

A NEW BIOLOGY CURRICULUM 53 their students to create this type of course mix. A student taking all the courses listed in one of the following examples would likely exceed the institution’s requirements for a biology major. Different choices will be made by different schools and different students. For example, the content of mathematics courses may be influenced by the types of material covered in that school’s biology courses. Opportunities to learn mathematical skills in a rich content context will enhance conceptual understanding and pro- cedural fluency. The committee envisions two levels of potential changes that could facilitate interdisciplinary learning. In the first level of change, the goal would be on increasing communication between science departments and working together to develop and integrate modules into preexisting courses. The following chapters of the report present some examples of potential modules that could be used to provide students with real-world examples of how mathematics, chemistry, physics, computer science, and engineering are useful in the study of biology. In the second level of change, interdisciplinary courses could be developed (possibly using team teaching approaches) or biology-focused science or mathematics courses could be developed. The committee recognizes that it may be difficult for some schools, particularly small ones, to add new courses unless they re- place preexisting course offerings. However, these same schools may have other advantages, such as a small science faculty that is used to working with colleagues outside their own immediate area of specialization that would facilitate the creation of modules or increase the feasibility of team teaching. Some aspects of curriculum A are more complex than can be repre- sented in the table that follows: The yearlong mathematics sequence sug- gested for first-year students could be a newly designed course modeled after Case Study #4 taught at the University of Tennessee, or one that cov- ers selected aspects of calculus, differential equations, linear algebra, and statistics. At some schools, students will continue to take traditional math- ematics courses. For some of those students, calculus would be appropri- ate, others will need remedial mathematics courses, still others will enter with calculus and might enroll in discrete math and/or computer science courses. For more ideas, see Appendix F: Mathematics and Computer Science Panel Summary. Possible biology electives (for the senior year) in- clude Bioinformatics and Computational Biology, Mechanics of Organ- isms (see Case Study #5), Organismal Physiology, Comparative or Human Anatomy, Toxicology, Neurobiology, and Environmental Biochemistry. At

54 BIO2010 Potential Curriculum A Fall Spring First year Introductory Biology I (and lab) Introductory Biology II (and lab) Inorganic Chemistry (and lab) Organic Chemistry I (and lab) Introductory Math Ia Introductory Math II Faculty Research Seminar General Education Elective General Education Elective General Education Elective Sophomore Molecular Biology Cell and Developmental Biology Organic Chemistry II (and lab) Biochemistry Introductory Physics I (and lab) Introductory Physics II (and Engineering lab) General Education Elective General Education Elective General Education Elective General Education Elective Junior Analytical/Physical Chemistry Evolutionary Biology/Ecology (and lab) Biology Laboratory Course Genetics General Education Elective General Education Elective General Education Elective General Education Elective Independent Laboratory Research Independent Laboratory Research Senior Biology Elective Biology Elective Science Elective Science Elective Faculty Research Seminar General Education Elective General Education Elective General Education Elective Independent Laboratory Research Independent Laboratory Research aFor more ideas, see Appendix F: Mathematics and Computer Science Panel Summary.

A NEW BIOLOGY CURRICULUM 55 Potential Curriculum B Fall Spring First year Introductory Biology I (and lab) Introductory Biology II (and lab) Inorganic Chemistry (and lab) Probability and BioStatistics Introductory Math I Introductory Math II Faculty Research Seminar General Education Elective General Education Elective General Education Elective Sophomore Molecular Biology Cell and Developmental Biology Differential Equations Organic Chemistry I (and lab) Introductory Physics I (and lab) Physics II (and Engineering lab) General Education Elective General Education Elective General Education Elective General Education Elective Junior Genetics Evolutionary Biology/Ecology Organic Chemistry II (and lab) Biology Laboratory Course Physics III (and Engineering lab) Biochemistry General Education Elective General Education Elective Independent Laboratory Research Independent Laboratory Research Senior Biology Elective Advanced Mathematics (e.g., Science/Biology Elective discrete math that builds on Analytical/Physical Chemistry genetics already learned) (and lab) Science/Biology Elective Faculty Research Seminar General Education Elective General Education Elective Independent Laboratory Research Independent Laboratory Research

56 BIO2010 least some of the upper-level biology courses should include labs. For ex- ample, students might take a lab along with genetics, molecular biology, or biochemistry, but not necessarily with all three courses. Alternatively, a more quantitative track could be designed as an option for students who are interested in exploring the interfaces between biology, mathematics, computer science, and the physical sciences (Curriculum B). A more radical change in undergraduate biology proposal appears as Potential Curriculum C below. The key idea is that contemporary biology cannot be taught effectively until students have a sufficiently strong back- ground in chemistry, physics, math, and computer science. Consequently, biology is not taught in the first year, apart from a seminar designed to whet the appetite of students for biological research and stimulate their acquisition of a strong background in the physical sciences. Rather, the first year is devoted to providing students with the requisite background in the physical sciences and mathematics. It is difficult to teach chemistry, physics, math, and computer science all in the first year. To succeed, the content of these courses has to be quite different from that of traditional courses in these areas. Also, the notion that an introductory course must occupy two semesters in the same aca- demic year would have to be put aside. The primary objective of the first year would be to provide students with the physical science knowledge and tools needed to effectively study biology starting in the second year at a level that prepares them for contemporary biological research as it is being carried out today. In the proposed curriculum, Chemistry I and II would introduce students to inorganic chemistry, organic chemistry, and key as- pects of biomolecular interactions. Math I would deal with differential cal- culus and elementary linear algebra, and Math II with integral calculus, probability, and statistics. Computer Science I would teach algorithms, simulation of dynamical systems, string (sequence) comparisons, and clus- tering; a high-level language such as Matlab or Mathematica would be used. Physics I would present mechanics, followed by equilibrium statistical phys- ics. Waves, electrostatics, and collective phenomena would be presented in Physics II, followed by signal analysis and processing, basic quantum me- chanics, and spectroscopy in Physics III. The four-semester core biology sequence (Molecular Biology, Cell and Developmental Biology, Genetics, and Evolutionary Biology/Ecology) start- ing in the sophomore year could be taught with a quantitative emphasis that would draw more heavily than now on the physical sciences, math- ematics, and computer science. For example, emergent system properties at

A NEW BIOLOGY CURRICULUM 57 Potential Curriculum C Fall Spring First year Biology Seminar Physics I (and lab) Chemistry I (and lab) Chemistry II (and lab) Math Ia Computer Science I General Education Elective General Education Elective General Education Elective General Education Elective Sophomore Molecular Biology Cell and Developmental Biology Math II Biophysical Chemistry Physics II (and lab) Physics III (and Engineering lab) General Education Elective General Education Elective General Education Elective General Education Elective Junior Genetics Evolutionary Biology/Ecology Biochemistry Biology Laboratory Course Biology Elective General Education Elective General Education Elective General Education Elective Independent Laboratory Research Independent Laboratory Research Senior Biology Elective Math or Computer Science Chemistry Elective Elective Science/Biology Elective General Education Elective Faculty Research Seminar General Education Elective General Education Elective Independent Laboratory Research Independent Laboratory Research aFor more ideas, see Appendix F: Mathematics and Computer Science Panel Summary.

58 BIO2010 Potential Curriculum D Fall Spring First year Introductory Biology I (and lab) Introductory Biology II (and lab) Inorganic Chemistry (and lab) Organic/Biochemistry I (and lab) Calculus and Differential Calculus and Differential Equations I Equations II General Education Elective Faculty Research Seminar General Education Elective General Education Elective Sophomore Molecular Biology Cell and Developmental Biology Organic/Biochemistry Biostatistics II (and lab) Introductory Physics I (and lab) Introductory Physics II (and lab) General Education Elective General Education Elective General Education Elective General Education Elective Junior Genetics (and lab) Evolutionary Biology Computer Science Biology Laboratory Course General Education Elective General Education Elective General Education Elective General Education Elective Independent Laboratory Independent Laboratory Research Research Senior Biology Elective Biology Elective Science Elective Science Elective Faculty Research Seminar General Education Elective General Education Elective General Education Elective Independent Laboratory Independent Laboratory Research Research

A NEW BIOLOGY CURRICULUM 59 all levels of biological organization (e.g., in signal transduction cascades, genetic regulatory circuits, and ecosystems) could be taught making exten- sive use of quantitative models. The fourth potential curriculum is intended for students who are espe- cially interested in evolution, ecology, and systematics. It assumes students enter already having taken calculus and calls for specific courses in biosta- tistics and computer science, essential tools for the study of evolution. Stu- dents focusing on evolution may go on to pursue many types of activities, ranging from field research to clinical research. As discussed earlier, the connections between different types of biology are growing stronger just as the connections between different sciences are growing. Biology is an increasingly complex science that is truly an integrative discipline in which many aspects of mathematics and physical science con- verge to address biological issues. For biology majors to receive an optimal education, the content of their curriculum must be updated to address the interdisciplinary nature of the field. At many institutions, this will mean changes in the course offerings so that those who will become future bio- medical researchers learn more mathematics and more physical and infor- mation sciences than is currently required. It continues to make sense for biology majors to take introductory courses in chemistry and physics and to enroll in courses in the mathematics department. However, for this practice to be most useful, the students must learn how to relate the mate- rial they learn in those courses to biology and how to relate the material they learn in biology courses to chemistry and physics. Perhaps of equal importance, students majoring in mathematics and physical sciences should learn how to relate the material they learn to issues of biology. The recommendations of this report will not be achieved solely by transforming an undergraduate’s schedule into one of the curricular ex- amples shown above. However, much can be accomplished without alter- ing the current list of course titles. The content of the courses must change to incorporate the concepts presented in the first half of this chapter. Dif- ferent schools will likely create different sets of courses. Incorporating these themes into biology courses and ensuring that they are covered in other science courses taken by biologists will greatly benefit the education of biology majors, as well as, the committee believes, other undergraduates who are enrolled in these courses.

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Biological sciences have been revolutionized, not only in the way research is conducted—with the introduction of techniques such as recombinant DNA and digital technology—but also in how research findings are communicated among professionals and to the public. Yet, the undergraduate programs that train biology researchers remain much the same as they were before these fundamental changes came on the scene.

This new volume provides a blueprint for bringing undergraduate biology education up to the speed of today's research fast track. It includes recommendations for teaching the next generation of life science investigators, through:

  • Building a strong interdisciplinary curriculum that includes physical science, information technology, and mathematics.
  • Eliminating the administrative and financial barriers to cross-departmental collaboration.
  • Evaluating the impact of medical college admissions testing on undergraduate biology education.
  • Creating early opportunities for independent research.
  • Designing meaningful laboratory experiences into the curriculum.

The committee presents a dozen brief case studies of exemplary programs at leading institutions and lists many resources for biology educators. This volume will be important to biology faculty, administrators, practitioners, professional societies, research and education funders, and the biotechnology industry.

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