8

Education

Building a new scientific field is a multigenerational project. It is clear that realizing the promise of biological physics depends on what students are taught, and on how this material is presented. Developing effective educational strategies is vital for communicating the enticing intellectual opportunities of the field and for attracting talented aspiring scientists from the broadest possible cross-section of our society. The importance of this challenge is reflected in the fact that the majority of input the committee received from the community—voiced at the two town halls and in writing through the online platform—was about education. This input came from colleagues at all career stages—from senior faculty to beginning students—and from a wide range of institutions, including community colleges, primarily undergraduate institutions, and major research universities.

Science education is about much more than educating scientists. On the largest scale, a crucial responsibility of the scientific community is to contribute to the scientific literacy of the citizenry at large. A successful education transmits not just the importance of science for human health and the economy, but also a sense of wonder at the beauty and intricacy of our world. Science helps us to understand the world, but also holds that understanding to exacting standards, reminding us how difficult it can be to find convincing answers to important questions. Science is not just a foundation for technology and medicine, but part of human culture, and biological physics has a unique role to play in this larger cultural enterprise. The field combines the grandeur of the physicists’ search for unifying principles with our human interest in ourselves; it brings extraordinary instruments that allow us literally to see what has never been seen, while engaging with the remark-

able diversity of life on Earth; and it provides foundations for developments in technology and medicine that have revolutionized our lives. The unique appeal of combining the physicist’s style of inquiry with the striking qualitative phenomena of life confers a special opportunity to attract a broader and more diverse community of students. This chapter explores how the emergence of biological physics fits into the culture of physics education, how biological physics can be integrated into the physics curriculum, and how this field can be leveraged to enhance the education of scientists more generally.

As we explore the educational challenges and opportunities created by the emergence of biological physics, it will be clear that some of these are internal to physics departments, while others involve collaboration between physics faculty and colleagues in other departments. Some are grounded in the frontier questions in our field, while others leverage the lessons of our field to explore foundational topics across disciplines. Some opportunities are in traditional classroom teaching, and others involve integrating teaching with research.

While the opportunities are inspiring, there are also barriers, both structural and perceived. Physics students who become interested in biological physics often wonder to what extent they need to “learn biology,” something viewed as being outside of physics, taught in the equivalent of a foreign language. Conversely, biology students at most colleges and universities are required to have only minimal preparation in mathematics, so that physics is taught in a foreign language for them as well. Importantly, the scale of this cultural divide does not reflect the current state of the scientific enterprise: Biology today is a vastly more quantitative enterprise, more integrated with the mathematical and physical sciences, than it was a generation ago, and the curriculum for biology students has not kept pace. These problems will not be solved by making longer lists of courses from multiple departments and congratulating ourselves for our multi-disciplinarity. They demand a thoughtful approach to integrating biological physics into the fabric of physics education, and science education more generally, in ways that truly add value for all students.

Given the enormous variety of institutions and environments in which education takes place, there is no one-size-fits-all model for addressing the challenges and opportunities identified here, but there are some general principles on which to build these efforts. In defining biological physics, this report has emphasized that physics more generally is distinguished not by the objects or systems that are being studied, but rather by the kinds of questions that are being asked and by the kinds of answers that the physics community finds satisfying. Similarly, the teaching of physics is distinguished by a certain style and ambition: the focus on general principles, and the demonstration of how these principles are used to predict the behavior of particular systems; the sense that numbers matter, and that numerical facts about the world make sense in relation to one another and to the general principles; that one can construct instruments to measure these numbers, reliably; that much is understood

by simplifying, and that sometimes even over-simplification is productive. Time and again, seemingly distant subfields of physics have been found to be connected deeply to one another, emphasizing that well-educated physicists need to develop the intellectual breadth that will prepare them to make and appreciate the next new connections that are discovered. These grand goals are not always achieved, but creating a new subfield of physics does not exempt the community from these aspirations. On the contrary, as physics expands it becomes even more important to transmit this core, unifying culture.

It is essential to acknowledge that any proposal to add something to the curriculum requires that something else be taken out, or at least compressed. This is painful, but crucial, and perhaps quite urgent. Typical core physics curricula today hardly require undergraduates to learn anything that happened after 1950, while modern biology and computer science focus on ideas and results from after 1950. Should we be surprised, then, to hear people speak of physics as the science of the past, while biology and computing are the sciences of the future? The findings, conclusions, and recommendations that emerge from this report address only part of this larger issue.

CURRENT STATE OF EDUCATION IN BIOLOGICAL PHYSICS

The current state of education in biological physics is largely a state of untapped opportunity. While a healthy community of biological physics researchers can be found in graduate schools and at the postdoctoral level, it is quite possible for today’s undergraduate student to earn a degree in physics without ever encountering the physics of living systems. Exposure to the field, if it occurs at all, typically happens in an undergraduate’s junior or senior year, making it difficult for students to engage deeply with the field before they graduate. At the same time, students in other fields, who sample physics only through a single introductory course, may get no hint about the relevance of physics to the phenomena of life.

These missed opportunities are more an artifact of history than a thoughtful analysis of the best path of study for today’s students. Since the 1960s, many college physics programs have taken a narrower and more focused view of the subject, even as physics itself has become a much broader enterprise. A good illustration of this is provided by the table of contents of Fundamentals of Physics,¹ a textbook widely used for introductory physics courses. The book starts with Newtonian mechanics and builds the subject through electricity and magnetism toward the concluding chapters on nuclear and particle physics and the big bang. There is a short excursion into thermal physics, but the modern view of statistical physics does not make an appearance, nor does Brownian motion, despite its central historical role

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¹ D. Halliday, R. Resnick, and J. Walker, 2013, Fundamentals of Physics, Extended, 10th Edition, John Wiley and Sons, New York.

as the proof that matter is composed of discrete atoms and molecules. There is no effort to situate physics in relation to intellectual challenges outside this limited canon.² Such a structure reinforces the idea that there are two paths in the development of physics as a view of the natural world, one toward the very small (particles) and the other to the very large (the universe).

This tight focus on physics as an exploration of the very small and the very large leaves little room for students to experience the power of physics principles to illuminate the often-dramatic behaviors of systems at each of the many intermediate scales. It completely misses the idea that phenomena at intermediate scales are not merely a consequence of principles from the scale below, but may be a source of fundamental concepts and challenges in and of themselves. Indeed, the largest subfield of physics today, condensed matter physics, is defined by its focus on these emergent phenomena. While one might once have been able to think of this as quantum mechanics “applied” to macroscopic materials, it is now understood that condensed matter gives us concepts of great depth and broad applicability—order parameters, spontaneous symmetry breaking, scaling, the role of topology, and more. While many institutions do better, the typical introductory physics course gives little hint that one could learn anything fundamental by studying a block of metal, let alone a living cell.

The neglect of the living world, and its exploration by the physics community, continues into more advanced physics courses. Discussions of electric circuits and current flow seldom touch on the electrical dynamics of neurons; advanced mechanics courses seldom hint at the challenges of walking; optics courses rarely explain the principles of optical trapping or super-resolution microscopy; and quantum mechanics courses leave as mysteries the broad optical absorption bands of biological molecules, so different from atoms in gas phase but so central to the ability of life on Earth to capture the energy of the sun and to the rich colors that we experience every day. Statistical physics courses miss numerous opportunities to use the phenomena of life as illustrations of basic principles, while thermodynamics typically is presented without mentioning that experiments on animals played a key role in establishing the principle of conservation of energy. Fluid mechanics has slipped away from most core physics curricula, missing the opportunity to explore the surprising restoration of time-reversal invariance in the limit of large viscosity, and the profound implications of this for the movement of single cells. The end result is that physics students can easily get their undergraduate degrees without knowing that biological physics exists.

Of course, some physics programs offer undergraduates the opportunity to take special courses in biological physics or other subfields. However, this opportunity typi-

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² This approach contrasts strongly with that taken in The Feynman Lectures on Physics, based on a course taught precisely 60 years ago (R.P. Feynman, R.B. Leighton, and M. Sands, 1963, The Feynman Lectures on Physics, Addison-Wesley, Reading, MA).

cally arises only after several years of studying Newtonian mechanics, electricity and magnetism, and quantum mechanics. One major obstacle in teaching biological physics earlier is that statistical physics typically is not offered before the junior year. Since the principles of statistical physics are so central to how physicists think about living systems, encountering these principles late in the physics curriculum means most students will have only a limited time window in which to engage with biological physics.

More subtly, conventional undergraduate courses on statistical and thermal physics emphasize non-interacting systems, such as the ideal quantum gases, which can be given an exact microscopic description. In contrast, much of modern statistical physics is concerned with how interactions among many degrees of freedom drive the emergence of qualitatively new macroscopic phenomena, and how these emergent phenomena can be described using models that ignore many microscopic details. The renormalization group explains how this simplification happens, and connects very concrete behaviors of real materials to more general and abstract theoretical principles. Monte Carlo methods make it possible to explore more complex, interacting systems, far beyond the ideal gases that are the focus conventional courses. These ideas are central to the physicist’s exploration of life, both as theoretical methods and conceptual background.

In the same way that statistical physics provides much of the theoretical foundation for biological physics, modern optics is central to experimental biological physics. Optics itself has undergone revolutionary developments—from understanding the forces applied by light and the resulting invention of optical tweezers to the breaking of the diffraction limit to imaging, both recognized by Nobel Prizes—and this has been intertwined, beautifully, with developments in biological physics, as described in detail in Part I of this report. These connections, and even core ideas such laser-scanning imaging and super-resolution microscopy, are absent from the laboratory experience of most undergraduate physics students. There are missed opportunities both for the integration of biological physics into the curriculum and for the presentation of deep and fundamental physics.

In many cases ideas of great relevance for biological physics lie just beyond the bounds of traditional physics courses. Advanced mechanics courses typically do not point to the broader mathematical analysis of dynamical systems. Statistical physics courses, certainly at the undergraduate level, end before students can see Brownian motion as the primordial example of a stochastic process or realize that Monte Carlo simulation provides a path for exploring probabilistic models of systems well beyond thermal equilibrium. While physics teaching is properly focused on core subjects, all students would be well served by seeing that these subjects touch a wider variety of problems.

The problems identified in the undergraduate physics curriculum have a profound impact on efforts to grow the biological physics community. But these problems are more general, and have a much broader impact. The physics community today works on a range of problems that is much broader than what could be

imagined when most of the current curriculum was solidified. The community has allowed a great chasm to develop between how active researchers think about physics and what is conveyed to the typical undergraduate. The emergence of biological physics is just one reason to think more deeply about the core physics curriculum.

Beyond the core curriculum, physics departments typically offer courses in the subfields of physics, both at the undergraduate and the graduate level. These courses play an important role in educating the students who will do research in these fields. Some departments also insist that graduate students take a number of these courses outside their research field in order to broaden their physics culture. It is important that while many areas of physics have strong connections to other disciplines, these courses are taught at a level suitable for advanced physics students. As an example, it is helpful for students interested in condensed matter physics to understand how their field is connected to areas of materials science, electrical engineering, and chemistry, but a multidisciplinary course that is built around these connections would not be an effective substitute for a course on condensed matter physics itself. It is even more important for students interested in biological physics to understand how their field is connected to many different parts of biology, but again exploring these connections cannot substitute for an advanced course on the physics of living systems itself. Such courses still are quite rare.

The structure of the undergraduate physics curriculum also influences how physics is viewed and understood by students and scientists in other disciplines. For the biological physics community the most important of these connections is with students and colleagues in the life sciences. For the vast majority of these students, their only interaction with physics is through an introductory “physics for life scientists” or “physics for premedical students” course. A traditional distinction between these courses and the introductory courses aimed at physicists and engineers is that the course for biologists does not make use of calculus, even when teaching mechanics (for which calculus was developed). In many cases, this course is required but not functionally prerequisite to other courses, and students therefore wait to take the course in their final year of undergraduate study. This structure completely misses the opportunity to convey how physics principles bear on the variety of biological problems that life science students confront, what the methods and concepts of physics have taught us about life’s mechanisms, or more generally, how the physicist’s perspective on the mathematical description of nature could guide further explorations.

In the same way that the phenomena of life are absent from the physics curriculum, the concepts and methods of physics are absent from the biology curriculum. In many ways, this gap is more surprising, since these concepts and methods have played such a crucial role in so many parts of biology. Addressing these issues requires appreciating the great breadth of the biological sciences, and situating the role of physics in the larger project of building a more quantitative biology. These topics are addressed in the next section.

In contrast to the situation for undergraduates, engagement with biological physics in graduate-level physics education is relatively strong. The National Center for Science and Engineering Statistics (NCSES) tracks the awarding of PhDs in the United States by field and subfield; since 2004 NCSES has tracked biological physics as a subfield of physics, with the results shown in Figure 8.1. Although many people have the sense that biological physics is a nascent or minor activity in the physics community, in fact, the number of students receiving PhDs and doing their thesis research in biological physics now is comparable to the numbers in well-established subfields, and this has happened in just 15 years. Biological physics today is producing the same number of new PhDs as did elementary particle physics in the years 2000–2005, and is growing.

In many physics departments, applicants to the PhD program are expected to articulate an area of interest. While this declaration is not binding, it does influence the admission process. Since many physics students receive their undergraduate degrees without learning that biological physics is a branch of physics, there is an obvious problem.

At the graduate level, biological physics education in physics programs—what is counted in Figure 8.1—coexists with a wide range of programs in the biological

sciences that have some overlap with the field. It is important to keep in mind that biology is a much larger enterprise than physics, producing, for example, nearly five times as many PhDs per year, spread across many more distinct subfields. As emphasized throughout this report, many of these subfields—molecular biology, structural biology, cell biology, systems biology, neurobiology, and more—have had, and continue to have, important input from the ideas and methods of physics; many of these activities are identified as biophysics. Although it might be more accurate to view all of this activity as a continuum, the NCSES tracks the number of PhDs given in “biophysics (biological sciences)” as well as in “biophysics (physics),” as shown in Figure 8.2. Over the past decade, the number of PhDs granted in biophysics (biological sciences) has declined slowly, while the number in biophysics (physics) has increased, with the total remaining relatively constant. Thus, where physics students who became fascinated by the phenomena of life once saw themselves as becoming biologists, today they can retain their identity as physicists.

Finding: There has been considerable growth in the number of PhD students working in biological physics, so that the field now is comparable in size to well-established subfields of physics. This growth has occurred in less than a generation, and is continuing.

Finding: Biological physics remains poorly represented in the core undergraduate physics curriculum, and few students have opportunities for specialized courses that convey the full breadth and depth of the field.

Conclusion: The current physics curriculum misses opportunities to convey both the coherence of biological physics as a part of physics and its impact on biology.

STRENGTHENING BIOLOGICAL PHYSICS EDUCATION

The missed opportunities in biological physics education occur at every stage of the educational pipeline, from K–12 education to the launch of a scientific career. There are numerous opportunities to strengthen the effort, both within physics departments and at areas of intersection with other fields. This discussion of education is in the context of the first conclusion, from Part I of this report:

Conclusion: Biological physics, or the physics of living systems, now has emerged fully as a field of physics, alongside more traditional fields of astrophysics and cosmology; atomic, molecular, and optical physics; condensed matter physics; nuclear physics; particle physics; and plasma physics.

At research universities, the ideal is that teaching and research missions are aligned and synergistic. In that context, it is useful to recall our first recommendation, again from Part I:

General Recommendation: Physics departments at research universities should have identifiable efforts in the physics of living systems, alongside groups in more traditional subfields of physics.

Certainly a group of faculty who are active members of the biological physics research community will play a critical role in responding to the educational challenges identified here. On the other hand, these challenges arise in many different parts of the physics and biology curricula, and responses cannot be segregated. Universities, and their physics departments, cannot assume that a small group of biological physics faculty will solve these problems on their own, without engaging a broader range of colleagues and without perturbing physics and biology education more generally.

Nearly half of physics education in the United States happens outside the research intensive, PhD granting institutions. Quantitatively,³ of the nearly 9,000 physics bachelor’s degrees conferred in the United States in 2018, 45 percent were from institutions that are focused on undergraduate education and do not grant doctorates, and 15 percent of recipients started their educations at community col-

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³ P.J. Mulvey and S. Nicholson, 2020, “Physics Bachelor’s Degrees: 2018,” American Institute of Physics, https://www.aip.org/statistics/reports/physics-bachelors-degrees-2018.

leges, presumably taking their introductory physics courses at those institutions. Far beyond the number of students who receive physics degrees, nearly 250,000 students are enrolled in calculus-based introductory physics courses each year. These observations on the scale and breadth of physics education emphasize that integration of the physics of living systems into undergraduate physics education cannot be done solely by the relatively small number of faculty who identify as part of the biological physics research community.

Educational challenges do not have one-size-fits-all solutions, not least because the environment for teaching varies enormously across institutions. Faculty need to be empowered—and given the necessary resources—to develop curricula that are most appropriate for their institutions, and the recommendations that follow are meant to provide guidance and support rather than prescriptions. As a practical matter, the discussion begins with issues that can be addressed within physics departments, then moves outward to issues that engage educational institutions and their supporting agencies more broadly, and then to considering the integration of education with research. Throughout these efforts, it is crucial to ground education in biological physics firmly in the intellectual framework and principles of physics, even as we draw examples and inspiration from the fields with which it intersects.

Biological Physics in the Physics Curriculum

The different fields of physics often are represented by specialized courses aimed at advanced undergraduates or graduate students. These courses extend and reinforce the core physics curriculum and give students a view of where the subject is today. But it is difficult to imagine a student receiving a physics degree and not realizing that there is something called elementary particle physics, even if they never take a specialized course in the subject, and similarly for other well-established fields. In different ways, the results and goals of these different parts of physics are not just appended to the curriculum, but integrated into the core.

General Recommendation: All universities and colleges should integrate biological physics into the mainstream physics curriculum, at all levels.

This integration will have different implications for different groups of students. Some students will find themselves electrified by the field, sparking an interest in pursuing biological physics as part of their undergraduate degree (if the option is available) or in graduate school. For others, the field could act as a gateway, drawing their interest into physics from other disciplines. For still others, the field will form one part of their general physics education and help to deepen and inform their understanding of physics principles and other physics subfields. Serving the needs of all of these groups of students requires a multifaceted approach. Timing

matters. Exposing students to biological physics too late in the undergraduate course of study can close off their options for specialization. Conversely, an early but narrow focus on biological physics risks compromising students’ foundational knowledge of physics, and they will end up less well prepared to embrace and address the complexity of the living world. The goal is to give students paths for exploring the field in ways that reinforce, rather than sacrifice, the depth and breadth of a general physics education. We emphasize once again that there is no unique solution to these problems, and that workable solutions must be tuned to the context at each individual institution. It also is crucial that curricular innovation receive institutional support.

Students will not become better prepared to “do physics” in the more complex context of living systems by learning less physics. Indeed, biological physics is not the only part of physics now addressing phenomena of greater complexity. While physics continues to be characterized by the search for simplicity, the community now searches in more complex contexts, whether using machine learning in the search for elusive particles or in the analysis of images of living cells, perhaps using similar analysis methods; this trend toward the exploration of more complex phenomena is accelerating. Ideally, a physics degree will prepare our students for physics as it is practiced today, and as it is likely to be practiced in the next generation, starting from the beginning and continuing through the entire curriculum.

The Core Curriculum

The most straightforward way to expose physics students to living systems early in their education is to weave topics from biological physics into introductory courses. Standing waves literally come to life when explaining the physics of how Escherichia coli finds its middle via Min protein oscillations (Chapter 1); the physics of diffusion leads to fundamental limits on how well bacteria can sense their environment (Chapter 1); the statistical mechanics of two-level systems can be used to address single molecule experiments on ion channels (Chapter 2); and the mechanics of a bouncing pogo stick can rationalize aspects of animal movement, from cockroaches to kangaroos (Chapter 7). Using examples from living systems as a teaching tool in physics can provide early exposure to the field while simultaneously introducing key principles that form the foundation of a physics education.

Specific Recommendation: Physics courses and textbooks should illustrate major principles with examples from biological physics, in all courses from introductory to advanced levels.

Examples from biological physics can be used to teach topics in the core physics curriculum well beyond the introductory level. To revisit the list of missed op-

portunities from above, but now in positive terms: flying, swimming, and walking provide an engaging universe of examples of classical mechanics; the dynamics of neurons provide examples of electric circuits and current flow; optical trapping and super-resolution microscopy illustrate deep principles of electromagnetism and optics; and the broad optical absorption bands of biological molecules, which literally give color to much of our world, provide opportunities to build quantum mechanical intuition beyond the energy levels of isolated atoms. The concepts and methods of statistical physics, in particular, are illustrated by numerous phenomena from the living world, on all scales from protein folding to flocking and swarming, as seen in detail in Chapter 3. In an important counter to the impression that experiments on biological systems are messy, some of the most quantitative tests of simple polymer physics models, and the notion of entropic elasticity, have been done with DNA (see Figure 5.1).

For both the introductory and more advanced courses, the community of biological physicists has a special role to play in identifying good examples. But integrating these examples into the canon of physics teaching is a project that needs to be adopted by the broader community of physics faculty.

Conclusion: There is a need to develop, collect, and disseminate resources showing how examples from biological physics can be used to teach core physics principles.

There have been several good starts in this direction, but the committee concludes that much more is needed.

As noted above, two topics in the core physics curriculum stand out for their great relevance to biological physics—statistical physics and optics. Unfortunately, these topics also stand out for the size of the gap between the typical presentation to undergraduates and our modern understanding of the subjects. The committee believes that these problems are important not just for the progress of biological physics, but for the progress of physics more generally.

Finding: Current undergraduate courses in statistical mechanics often do not reflect our modern understanding of the subject, or even its full historical role in the development of physics. Among other neglected topics, Brownian motion, Monte Carlo simulation, and the renormalization group all belong in the undergraduate curriculum.

Finding: Statistical mechanics courses typically come late in the undergraduate curriculum, limiting the window in which students can explore biological physics with an adequate foundation.

Specific Recommendation: Physics faculty should modernize the presentation of statistical physics to undergraduates, find ways of moving at least parts of the subject earlier in the curriculum, and highlight connections to biological physics.

Finding: Current treatment of optics in the undergraduate physics curriculum does not reflect modern developments, many of which have strong connections to biological physics. Among other neglected topics, optical traps and tweezers, laser scanning, nonlinear optical imaging modalities, and imaging beyond the diffraction limit all belong in the undergraduate curriculum.

Specific Recommendation: Physics faculty should modernize undergraduate laboratory courses to include modules on light microscopy that emphasize recent developments, and highlight connections to biological physics.

Specialized Coursework in Biological Physics

Beyond exposing students to biological physics in the core of their physics education, creating opportunities for interested students to delve more deeply raises a variety of additional considerations: What mathematics background and physics experience are needed before taking a specialized course on biological physics? What is the appropriate balance between biological physics coursework and general physics coursework for students who choose to specialize in this field? What are the appropriate roles for laboratory research experience, inquiry-based learning, and computational approaches? Should biological physics be offered as a separate major or track, or folded into the traditional physics program? Many of these questions arise for other fields of physics. While there are no universal answers to these questions, it is possible to identify a few guiding principles.

A well-educated physicist, regardless of specialization, is able to “think like a physicist” and make connections between different subfields of physics. One risk of creating an overly specialized program is that it can become obsolete, for example, as technology changes. Even when graduate-level courses in biological physics are offered, they are often rather narrow in scope. Emphasizing the approach to the living world through a physics mindset provides students with a flexible foundation that can later be adapted as fields and technologies evolve. Ensuring the breadth and depth of the general physics education is equally important for students who wish to pursue special study in biological physics as it is for those interested in any other subfield of physics. This foundation of general physics principles is crucial to a deep and productive exploration into the complexity of the living world.

Traditional biophysics or biological physics courses often are fragmented along lines defined by the subfields and history of biology. In practice, such a course

might include protein structure but not the dynamics of neurons and networks, it may cover the mechanics of the cytoskeleton but not the collective behavior of flocks and swarms, and so on. To be consistent with the rest of the physics curriculum, teaching biological physics needs to be organized around conceptual questions and general principles—with diverse case studies as manifestations of those principles—rather than along a succession of disconnected biological topics. In addition to reinforcing the general physics culture, this approach will help physics students, who may not have extensive previous knowledge of biology, not to get lost in a sea of biological details.

Conclusion: The great breadth of the field poses a challenge in teaching an introduction to biological physics for advanced undergraduates or beginning graduate students.

General Recommendation: Physics faculty should organize biological physics coursework around general principles, and ensure that students specializing in biological physics receive a broad and deep general physics education.

An important part of the physicist’s approach to nature, which also is central to the teaching of biological physics, is that our understanding is expressed in mathematical terms and tested in quantitative experiments. This interaction between theory and experiment can reach extraordinary precision, but physics crucially also is about simple approximate arguments. When something is finally understood, it is possible both to give order of magnitude estimates on the back of an envelope and to predict the results of detailed experiments, and the path to understanding often involves an interplay between these different approaches. Understanding also generates the ability to engineer new and often simplified systems, capturing the essence of what we see in nature; engineering in turn probes the limits of our understanding. In discussing the differences between physics and biology education, emphasis often is placed on the role of sophisticated mathematical analysis. This indeed is essential for physics, but focusing on this alone misses the roles of both quantitative experiment and simple models. Physics has a culture of quantitative measurement so pervasive that it can be taken for granted, and a taste for simple arguments that can feel more like art than science. But there are often explicit claims that biology is different, and that the complexity of life is both irreducible and irreducibly messy. Education needs to confront this problem, explicitly.

One of the basic conclusions from the vast array of work reviewed in Part I of this report is that the complex phenomena of life can be tamed, resulting in the sorts of reproducible, quantitative experiments that are the norm in the rest of physics. There are examples of this on all scales, from single molecules to populations of organisms, and in many cases, taming the complexity has involved building

new instruments and introducing methods of data analysis that are grounded in more abstract theoretical principles. These results exemplify what is possible when the community holds to high standards for quantitative measurements and, more deeply, in the comparison between theory and experiment. Students need to be taught that the complexity of living systems is not an excuse to be satisfied with lower quality data, or with merely qualitative comparisons between theory and experiment, and that complexity itself is not an argument against the exploration of simple models.

Mathematical Methods

Physics has a special relationship to mathematics, and this is true for all parts of the discipline including biological physics. As with the core physics curriculum itself, there is a canon of mathematical methods for physics. Some of this is conveyed in a collection of courses, taught in the mathematics department and often designed for the first 2 years of undergraduate education, moving from single variable to multivariable calculus, linear algebra, and differential equations. But the more advanced parts of even the undergraduate physics curriculum draw on eigenfunction expansions, complex analysis, asymptotic approximation methods, Fourier methods, and more. Individual physicists differ in their relationship to this material, and this diversity of views is transmitted to the students. Physics departments might require their undergraduates to take particular advanced courses in applicable mathematics, they might offer their own courses on the mathematical methods of physics, or they might assume that more advanced methods are taught as part of physics courses; many institutions offer a mix of these approaches.

For students interested in deeper exploration of biological physics, what is missing from the conventional collection of mathematical methods is not so much particular topics as an understanding that these methods fit into larger and more generally applicable structures. As an example, physics students take courses on differential equations and advanced classical mechanics, but these typically stop short of introducing more general ideas about nonlinear dynamical systems. This is important because many of the dynamical systems relevant to the living world—from networks of biochemical reactions in a single cell to networks of neurons in the brain to interactions among species in an ecosystem—do not have the symmetries and conservation laws that are central to classical mechanics. This broader notion of nonlinear dynamical systems also is relevant for many other areas of physics, is connected to the foundations of statistical mechanics, and provides accessible examples of universality and renormalization. Physicists in general would benefit from knowing that these topics exist, just beyond the bounds of their traditional courses.

In a similar spirit, elementary statistical mechanics courses often do not emphasize that it is a fundamentally probabilistic description of the world. In fact, everything we observe—even the pressure that an ideal gas exerts on its container—is predicted to fluctuate, and the analysis of these fluctuations has been crucial at many stages in the development of the subject. Leaving fluctuations as an advanced topic, often beyond the core undergraduate course, also misses the opportunity to situate statistical mechanics in the larger context of probabilistic models and stochastic processes, many of which are relevant to our description of the living world. Indeed, the exploration of probabilistic models is a huge field, with applications to an ever growing array of problems, from economics to health care, machine learning, and more. Many of the concepts and methods in this field have their roots in statistical mechanics, with the Boltzmann distribution as the primordial example of a probabilistic model. All physics students would benefit from knowing that these connections exist, and the physics community as a whole would benefit from reclaiming some of this larger field, now belonging primarily to computer science and applied mathematics, where physicists have made many contributions.

Conventional boundaries for the mathematical methods of physics were established before computers became widely available. Today, the role of computation in the practice of science cannot be overstated, and biological physics is no exception. As is well known, physicists use computation in two very different ways. First is the simulation of models, using computing as an extension of theory, to explore phenomena that are not yet captured with analytic methods. Second is the analysis of data, using computing as an extension of experiment to extract meaning from ever larger data sets.

Simulation provides a path for students to explore theoretical questions even when they are not fully prepared for the analytic theory; as an example, some statistical physics courses now use Monte Carlo simulation for this purpose. Importantly, simulation can close the gap between more traditional physics problems, such as random walks, and biological physics problems, such as the “run and tumble” behavior of bacteria. The cost of making simulations more realistic can be very low, allowing students to go beyond the proverbial spherical cow while still seeing connections to simple models. Higher-level languages with relatively transparent syntax, such as MATLAB and Python, reduce the barriers to getting started on computational projects, and Jupyter notebooks provide a structure for collaboration and the sharing of resources online. While using these tools to help students explore more widely, confidently, and even playfully, it also is important to convey the lessons from generations of computational physicists: that simulations have something in common with experiments, and need to be analyzed carefully; that the ease of simulating complex models does not replace our search for principled, simplified descriptions, ultimately expressible with pen and paper; that simulation is one path to understanding rather than an end in itself.

Almost all fields of science are being revolutionized by the opportunity to gather “big data.” While this often is presented as recent development, experimental high energy physics and cosmological surveys entered the big data era before it had a name. Today biological physics is following a similar path, with even modest experiments generating terabytes of data in an afternoon, and many experiments reaching the petabyte scale.

Conclusion: Biological physics, and physics more generally, face a challenge in embracing the excitement that surrounds big data, while maintaining the unique physics culture of interaction between experiment and theory.

Coda

Taken together, the recommendations above point toward a more general aspect of physics culture. Physics and physicists have played central roles in developments spanning the full range of science, technology, and policy, from plate tectonics and global climate to the semiconductor industry and the World Wide Web, from energy production to arms control. This engagement is an important part of physics culture, and is transmitted to students both informally and as side commentary in core courses and their textbooks. Despite the enormous impact that physics has had on our exploration of the living world, from basic science to the practice of medicine and care for public health, the phenomena of life remain largely absent from the broader notion of what physics is and what physicists do, that we transmit to our students. The recommendations here provide a path to closing this gap.

Biological Physics and Cross-Disciplinary Education

This report has emphasized biological physics as a branch of physics. This stands in contrast to the view of the field as the application of physics to biology, or as some interdisciplinary amalgam. The current state of the field emerged from rich interactions, over the course of a century, among distinct disciplines of physics and biology, as well as chemistry, psychology, and more. It is especially important to respect the richness of these interactions when teaching, while still conveying what is new and exciting about the physicist’s perspective on the phenomena of life. A central theme in this history is that many scientists have made progress by crossing traditional boundaries between disciplines. These boundary-crossing events have many different outcomes: In some cases, an individual scientist changes fields; in other cases, attention is drawn to the opportunities for interdisciplinary collaboration; and finally, the boundaries can move, as has happened in biological physics. Students need to understand not so much the sociology of these boundary--

crossing events, but rather that the ability to cross boundaries will expand their opportunities to formulate and solve problems that matter to them as individuals, to the scientific community, and to society as a whole.

One reaction to the complex history of biological physics, and to other examples of intellectual boundary crossing, is to emphasize that boundaries are artificial. Indeed, the phenomena in nature are not labeled intrinsically as being biology, chemistry, or physics. But faced with the same phenomena, biologists, chemists, and physicists will ask different questions, and expect different kinds of answers, as will applied mathematicians, computer scientists, and engineers. It is not reasonable to ask that these cultural differences be obliterated, any more than it would be reasonable to insist that all novels around the world be written in a single language. How should colleges and universities prepare students for a world in which their scientific interests will lead them to the edges of their chosen disciplines? What are the best ways to “translate” the culture of one discipline, making it accessible for others?

The analogy to language suggests that crossing boundaries between scientific disciplines is easiest if one starts early. Ideally, students would be exposed to biological physics at a pre-college level, potentially as early as middle school. In this early context, the interplay between physical laws and constraints and properties of living systems can motivate questions in both biology and physics. At the high school level, biological physics can be an integral part of both physics and biology education. This does not require a separate course; rather the relevance of physical considerations in understanding living systems can be taught as a part of high school biology, and high school physics courses can include examples drawn from biological systems, as described above for university courses. While the challenges of K–12 education are beyond the scope of the task for this report (Appendix A), the committee views them as crucial for progress in our field, and in science more broadly.

For most science students, their only contact with physics is through a single introductory course. Although sometimes referred to as “service courses,” these courses offer a great opportunity. A meaningful goal is to convey to each student how the concepts and methods of physics provide productive tools for exploring the parts of the world that they find most interesting—this is the true service that can be performed in these courses. In addition, these courses can aspire to convey something of the beauty and grandeur of physics itself. The previous section emphasized the opportunities for integrating biological physics into introductory courses for physics students, and the same arguments apply even more strongly to physics courses for students in the life sciences. Examples from biological physics illustrate many core principles of physics more generally, and the notion that these principles are relevant to the phenomena of life is itself an important fact, one that can change a young student’s view of the intellectual landscape. To make this work,

the community needs to develop a catalogue of illustrative biological examples that serve as equivalents to the inclined plane, simple pendulum, planetary motion, and so on, emphasizing our earlier conclusion on updates to the curriculum:

Conclusion: There is a need to develop, collect, and disseminate resources showing how examples from biological physics can be used to teach core physics principles.

Many widely used biology textbooks have zero equations. To the extent that mathematical analysis appears in the teaching of biology, it often focuses on the reliability of inference from limited data rather than on the phenomena themselves. This approach sends a clear message to students that numbers are irrelevant to the exploration of life, despite many counterexamples from the history of biology. By the end of the 20th century, it was clear to many people that this approach would not prepare students for the future of biology. This perspective was summarized in the BIO 2010 report.⁴

BIO 2010 was motivated by the observation that the practice of biology was changing rapidly: Instead of studying individual genes, the field was moving to studying whole genomes; instead of probing single neurons in the brain, it was becoming possible to monitor large populations of cells; and more. At a purely practical level, the scale of biological data was reaching the point that individual scientists could no longer reason “by hand” about their results, but instead needed to formalize their data analysis in algorithmic and ultimately mathematical terms. More broadly, much of biomedical research was characterized as being at the interface between biology and the physical, mathematical, and information sciences. BIO 2010 emphasized that revolutionary changes in the research environment should drive comparably profound changes in teaching. In particular, it was necessary to push biology students to develop more quantitative skills, and to be sure that biology courses draw on these skills throughout the curriculum. Making these changes would require new resources, both from the federal government and from individual academic institutions, and collaboration among faculty from multiple departments.

How far has the community come in responding to BIO 2010? Almost all research universities now have visible programs in areas that can be described as “quantitative biology,” although exactly what this means is different at different institutions, and the extent to which these programs are accessible to undergraduates also varies. Interestingly, many institutions have programs in biophysics

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⁴ National Research Council, 2003, BIO 2010: Transforming Undergraduate Education for Future Research Biologists, The National Academies Press, Washington, DC.

that anticipated the goals of BIO 2010 by decades. These programs have played an important role in helping students who have strong physics preparation engage with the frontier of biological research. In particular, biophysics programs have been a major source of students working on X-ray crystallography, nuclear magnetic resonance, and cryogenic electron microscopy approaches to the structure of biological molecules, well before these approaches merged into structural biology.

Even with the growth of quantitative biology programs, the basic requirements for traditional biology undergraduates remain light in mathematics and the physical sciences, and this has consequences for how more advanced biology students engage with central topics in the field. Thus, modern molecular biology and biochemistry courses are built around molecular structures, but little can be said about the experimental methods that make it possible to visualize these structures. Similarly, core neurobiology courses describe the central role of ion channels in brain function, but typically do not make reference to the equations that describe the dynamics of these molecules or to the deeply quantitative analyses that led to their discovery. Despite dramatic changes in the practice of biology as a science, this curriculum continues to send the message that mathematical approaches are optional rather than integral to our exploration of life.

Conclusion: There still is room to improve the integration of quantitative methods and theoretical ideas into the core biology curriculum, continuing the spirit of BIO 2010. This remains crucial in preparing students for the biomedical sciences as they are practiced today, and as they are likely to evolve over the coming generation.

While the BIO 2010 report referred broadly to the physical, mathematical, and information sciences, physicists in particular have a crucial role to play in the education of more quantitative biologists. The business of physics is the development of instruments for the quantitative observation of the natural world and the development of mathematical structures that rationalize these data, allowing understanding, prediction, and design. Physicists have been in the forefront of collecting and analyzing large data sets, and in building collaborations that create capabilities far beyond what can be accomplished by single investigators. Because of these traditions, physics departments have developed a substantial infrastructure for quantitative education—in the laboratory, at the blackboard, and at the computer. The emergence of biological physics as a branch of physics has made clear how all of this can be brought to bear on the phenomena of life.

Integrating the principles of biological physics into the education of more quantitative biologists needs to happen at many levels, and at all stages of educa-

tion. As noted above, the traditional model has been that biology students see physics only in an introductory course, and attention has therefore been focused on improving these courses. But students will not develop a more quantitative approach to the life sciences if equations never appear in their subsequent biology courses. Diverse educational experiments at institutions around the country provide examples for how to proceed:

• Physics, chemistry, and biology faculty can collaborate to offer an integrated introduction to the natural sciences, providing an alternative to separate courses in the individual disciplines.
• In the tradition of courses on the mathematical methods of physics, departments can offer courses on mathematical methods in biology, or in subfields of biology.
• Laboratory courses can introduce experimental methods from physics, the building of experimental apparatus, and the physics culture of connecting theory to experiment, while remaining focused on biological systems.
• Intense summer courses can bring together students from across the divide between physics and biology. Sustained over many years, these programs can help to close the gap between the disciplines.

What all these approaches to the education of quantitative biologists have in common is that they require collaboration among faculty from multiple departments. As was noted in the BIO 2010 report, such collaborative teaching often faces administrative obstacles.

Conclusion: The biological physics community has a central role to play in initiatives for multidisciplinary education in quantitative biology, bioengineering, and related areas.

General Recommendation: University and college administrators should allocate resources to physics departments as part of their growing educational and research initiatives in quantitative biology and biological engineering, acknowledging the central role of biological physics in these fields.

It is widely appreciated, following the arguments of BIO 2010, that the education of quantitative biologists must engage faculty in multiple departments. We would add that real progress requires departing from the model in which courses taught by one department are in “service” to the educational agenda of another department. University and college administrators must create funding structures that support genuine collaboration and equal partnership among all relevant faculty.

Coupling Education and Research

In the academic world, research and education are linked. Yet, only a small fraction of institutions require undergraduates to engage with research as part of their degree program, although many more offer research opportunities. Larger, research-intensive institutions, both universities and national laboratories, have summer programs to welcome visiting students who might not have comparable opportunities at their home institutions. It is generally agreed that engaging with research is enormously beneficial for undergraduates, and in some cases it is not an exaggeration to say that such experiences are life changing.

Biological physics research groups have a special role to play in the ecosystem of undergraduate research experiences. Many experimental groups in the field are small and focus on “table top” experiments, providing a more intimate community for young students. Theorists work on problems ranging from data analysis through simulation to abstract theory, providing opportunities for students to enter with varying levels of background knowledge. The same research groups can appeal to students planning a range of undergraduate majors, not just physics and biology but also chemistry and many fields of engineering. Seeing directly, in the laboratory and at the blackboard, how the physicist’s approach illuminates the phenomena of life can reignite students’ interests in physics itself.

Finding: Meaningful engagement with research plays a crucial role in awakening and maintaining undergraduate student interest in the sciences.

Conclusion: Biological physics presents unique opportunities for the involvement of undergraduates in research at the frontier of our understanding, offering more intimate communities through smaller research groups and providing opportunities for students to enter with varying levels of background knowledge and from a range of undergraduate majors.

The widespread enthusiasm for engaging undergraduates in research has been supported by federal funding agencies for a very long time. The National Science Foundation supported an Undergraduate Research Participation Program starting in 1958; cut from the federal budget in 1982, it was revitalized in 1987 as Research Experiences for Undergraduates (REU). The REU program has touched tens of thousands of students, and by now, almost every federal agency has mechanisms aimed specifically at supporting undergraduate involvement in research projects.

An unintended consequence of broader support for undergraduate research experience is that this experience has become a de facto requirement for admission into highly ranked doctoral programs. As a result, the community needs to attend

not just to the total amount of support for such programs, but to the equality of opportunity in access to these programs.

It is important to emphasize that meaningful student engagement in research builds on the foundation provided by the core curriculum. If the goal is to have students get involved in science as soon as possible, ideally in the summer after their first year of undergraduate study, then it is necessary to ask if the introductory courses are preparing our students properly for this experience. The recommendations above reveal additional considerations for the goals of introductory courses. Do laboratory modules instill a taste for measurement that prepares students to move toward the frontier of the subject? Do homework exercises hone the theoretical and computational skills that provide a starting point for engaging with data and models beyond our current understanding? These are challenging questions, and the path to more satisfying answers will necessitate resources beyond those that are currently allocated.

Institutions vary widely in the resources that they bring in support of introductory undergraduate physics courses. It is not unusual for these courses to involve a single faculty member lecturing to hundreds of students, perhaps with problem-solving sessions led by graduate student teaching assistants; institutions without robust doctoral programs may not be able to staff these smaller group discussions. In physics, perhaps more than in biology, one course builds on another, so that inequalities of investment in introductory courses are amplified with time.

Conclusion: Equality of opportunity for students to engage with physics, including biological physics, depends on high-quality introductory courses, emphasizing the interconnectedness of education and research.

Finding: Current models for support of undergraduate research perpetuate a sharp distinction between the core curriculum (education) and the development of the scientific workforce (research). This extends to the fact that science and education are overseen by different standing committees in Congress.

Conclusion: Support for the development of the scientific workforce will require direct federal investment in the core of undergraduate education, especially at an introductory level.

These observations lead to our recommendations about the integration of teaching and research:

Specific Recommendation: Universities should provide and fund opportunities for undergraduate students to engage in biological physics research, as an integral part of their education, starting as soon as their first year.

Specific Recommendation: Funding agencies, such as the National Institutes of Health, the National Science Foundation, the Department of Energy, and the Department of Defense, as well as private foundations, should develop and expand programs to support integrated efforts in education and research at all levels, from beginning undergraduates to more senior scientists migrating across disciplinary boundaries.

As explained above, many agencies have programs that support the engagement of undergraduates in research, and so these recommendations could be read simply as a plea for expansion of these programs. While this would help, the committee feels strongly that new programs are needed for more effective integration of teaching and research. Students need to see the connection between the core of their curriculum and the advancing frontiers of science not just in summer laboratory sojourns but in the classroom as well. While not all science students will become scientists, all will benefit from making these connections. Such efforts will take different forms in different institutions and for different groups of students.

POSTDOCTORAL TRAJECTORIES

The education of professional scientists does not end with the award of the PhD. Postdoctoral positions, once a brief and luxurious pause between graduate school and the responsibilities of a university faculty position, have become the destination for half of all new physics PhDs⁵ and a near absolute requirement for advancement to faculty and independent research positions. Postdoctoral periods also have become longer, so that what was once a transitional period is becoming a substantial phase of career and life; a corollary is that postdoctoral fellows are becoming a larger part of the scientific workforce. These trends are especially strong in the biomedical sciences, where they have been identified by prominent commentators as among the “systemic flaws” in the research enterprise.⁶ The situation in the physics community is different, but might not be better. In many areas of theoretical physics, for example, postdoctoral appointments are limited to 3 years, but it is common for people to have multiple postdoctoral sojourns before arriving at a faculty position. Young biological physicists are influenced by both cultures.

Because biology is a much larger enterprise than physics, many new PhDs in biological physics will move to postdoctoral positions in biology departments or to basic science departments at medical schools. On the one hand, this is a sign

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⁵ P. Mulvey and J. Pold, 2019, Physics Doctorates: Initial Employment, American Institute of Physics, https://www.aip.org/statistics/reports/physics-doctorates-initial-employment-2016.

⁶ B. Alberts, M.W. Kirschner, S. Tilghman, and H. Varmus, 2014, “Rescuing US Biomedical Research from Its Systemic Flaws,” Proceedings of the National Academy of Sciences U.S.A. 111(16):5773–5777.

of success for the field, and creates opportunities for productive exchange of ideas and expertise. On the other hand, the dispersal of young biological physicists adds to the problems of maintaining coherence in the field, discussed at many points in this report. For the individuals involved, the differences in culture surrounding postdoctoral positions in physics and biology can be a source of anxiety and can also make it difficult to return to a physics environment, if desired, at the next step in their careers.

Physics has a tradition of treating postdocs as budding independent investigators rather than merely as skilled labor. To maintain this tradition, fellowships that provide competitive salaries and some degree of independence are essential. These issues bridge the challenges of education and those of supporting the field more generally, addressed in Chapter 9.

Conclusion: Accelerating young researchers to independence is critical to empowering the next generation of biological physicists. As in other fields of physics, independent, individual fellowships are an effective mechanism.

To maintain coherence as postdoctoral fellows move to a wide variety of research environments requires support for their attendance at events that bring them into contact with the broad biological physics community. In this respect, an important role is played by institutions such as the Aspen Center for Physics and the Kavli Institute for Theoretical Physics, which have hosted many programs on topics in biological physics alongside those in better established subfields of physics. Such gatherings promote the exchange of ideas and formation of new independent collaborations, as well bringing promising postdoctoral fellows into contact with senior colleagues from outside their immediate circle of mentors.

The growth in biological physics also has attracted numerous scientists transitioning from other subfields of physics at the postdoctoral level. This again is a testament to the excitement and promise of the field. However, unlike students who enter biological physics as undergraduate or graduate students, postdoctoral scholars have limited time to learn, and often have more competing demands on their time. Intensive summer schools play an important role as “crash courses” that help these young scientists learn essential background quickly, and sample the frontiers of the subject. There are successful examples of this at the locations of well-known theoretical physics schools, such as École de Physique des Houches, Intitut d’Études Scientifiques de Cargèse, and the University of Colorado Boulder. There is an independent tradition of summer schools in different parts of experimental biology, and several of these have evolved in response to the influx of ideas, methods, and young people from physics; examples include the Marine Biological Laboratory, Cold Spring Harbor Laboratory, and Friday Harbor. These different courses span the range from “pure” physics courses to fully interdisciplin-

ary courses to biology courses where physicists are welcome. In some cases, these courses have run for decades, and their alumni have grown into leaders in both physics and biology. Continued financial support and community engagement with these courses is important.

Biological physics also is remarkable for the diversity of potential career opportunities, in both academic and nonacademic settings. While postdoctoral fellows in some areas of physics largely confine their searches for academic jobs to physics departments, biological physicists often face a bewildering array of academic job options. There are opportunities in physics departments, but these are not proportional to the size of the field. Parts of what this report describes as the broad field of biological physics have their natural home in chemistry departments, and at several institutions topics outside the more traditional areas of physics are found in applied physics departments; theorists may find homes in applied mathematics departments. Many departments in engineering schools (not just bioengineering) are more and more deeply connected to problems in the life sciences, in ways that often resonate with the biological physics community. Finally, there are opportunities in the many different kinds of biology departments that one finds at universities, and in the even wider range of departments and research centers found at medical schools.

There are practical challenges for postdoctoral fellows in preparing for this broad range of opportunities. Different departments give different weight to publications in different journals, for example, and still have varying attitudes about e-print archives (although these views are converging). Problems that physicists find closely connected might be the focus of different departments or programs in the biological sciences, or different groups in the same department, and dividing research effort among these problems could weaken the case in the eyes of each specialized group. Senior scientists have a responsibility to attend to these issues, often case by case.

Career opportunities for postdoctoral scholars outside of academia are varied and broad, and include positions in biotech and pharmaceutical industries as well as quantitative analysis and data science positions in an increasingly wide range of industries. As in academia more generally, there is considerable room for improvement in how young biological physicists are introduced to these opportunities.

SUMMARY

Bringing the physicists’ style of inquiry to bear on the phenomena of life has a unique appeal. Beyond the intellectual opportunities of the field itself, biological physics provides a path for communicating the excitement of physics more broadly, for attracting talented scientists from the widest cross-section of our society, and for contributing to the scientific literacy of the society at large. But realizing these

opportunities requires rethinking how we teach physics, biology, and science more generally. Today, physics students interested in biological problems can retain their identity as physicists, and, in less than a generation, the number of such students has become comparable to those in well-established subfields of physics. On the other hand, even though biological physics has emerged as a coherent subfield of physics, the phenomena of life are largely absent from the core undergraduate physics curriculum. Similarly, while methods and concepts from physics have played a central role our modern view of life, the principles of biological physics are largely absent from the core biology curriculum. These are symptoms of the large gap that has developed between the practice of science and the education of undergraduates. This chapter has examined these issues, leading to a series of interlocking findings, conclusions, and recommendations about:

• the integration of biological physics into the core physics curriculum;
• the need for modernization of the physics curriculum;
• courses on biological physics for advanced physics students;
• the special role of biological physics in building a more quantitative biology; and
• integration of education and research, and support for this integration.

Uniting these issues is a concern for the communication of general physics culture—an emphasis on general principles, on the power of both mathematical sophistication and simple arguments, and on the benefits of holding high standards both for quantitative measurements and for the interaction between experiment and theory. Beyond their formal education, young biological physicists face a remarkable diversity of potential career opportunities, but also must navigate the cultural differences they will encounter along these paths. Attention of senior scientists to these differences, as well as the availability of postdoctoral opportunities that provide some degree of independence, are essential for helping young biological physicists realize the promise of the field.