National Academies Press: OpenBook

Physics of Life (2022)

Chapter:Front Matter

Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2022. Physics of Life. Washington, DC: The National Academies Press. doi: 10.17226/26403.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2022. Physics of Life. Washington, DC: The National Academies Press. doi: 10.17226/26403.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2022. Physics of Life. Washington, DC: The National Academies Press. doi: 10.17226/26403.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2022. Physics of Life. Washington, DC: The National Academies Press. doi: 10.17226/26403.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2022. Physics of Life. Washington, DC: The National Academies Press. doi: 10.17226/26403.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2022. Physics of Life. Washington, DC: The National Academies Press. doi: 10.17226/26403.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2022. Physics of Life. Washington, DC: The National Academies Press. doi: 10.17226/26403.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2022. Physics of Life. Washington, DC: The National Academies Press. doi: 10.17226/26403.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2022. Physics of Life. Washington, DC: The National Academies Press. doi: 10.17226/26403.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2022. Physics of Life. Washington, DC: The National Academies Press. doi: 10.17226/26403.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2022. Physics of Life. Washington, DC: The National Academies Press. doi: 10.17226/26403.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2022. Physics of Life. Washington, DC: The National Academies Press. doi: 10.17226/26403.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2022. Physics of Life. Washington, DC: The National Academies Press. doi: 10.17226/26403.
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Suggested Citation:"Front Matter." National Academies of Sciences, Engineering, and Medicine. 2022. Physics of Life. Washington, DC: The National Academies Press. doi: 10.17226/26403.
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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.

Prepublication Copy – Subject to Further Editorial Correction Physics of Life Decadal Committee on Biological Physics/Physics of Living Systems Board on Physics and Astronomy Division on Engineering and Physical Sciences Board on Life Sciences Division on Earth and Life Studies A Consensus Study Report of PREPUBLICATION COPY – SUBJECT TO FURTHER EDITORIAL CORRECTION

THE NATIONAL ACADEMIES PRESS 500 Fifth Street, NW Washington, DC 20001 This study is based on work supported by Contract 1760032 with the National Science Foundation. Any opinions, findings, conclusions, or recommendations expressed in this publication do not necessarily reflect the views of any agency or organization that provided support for the project. International Standard Book Number-13: 978-0-309-XXXXX-X International Standard Book Number-10: 0-309-XXXXX-X Digital Object Identifier: https://doi.org/10.17226/26403 Copies of this publication are available free of charge from Board on Physics and Astronomy National Academies of Sciences, Engineering, and Medicine 500 Fifth Street, NW Washington, DC 20001 Additional copies of this publication are available from the National Academies Press, 500 Fifth Street, NW, Keck 360, Washington, DC 20001; (800) 624-6242 or (202) 334-3313; http://www.nap.edu. Copyright 2022 by the National Academy of Sciences. All rights reserved. Printed in the United States of America Suggested citation: National Academies of Sciences, Engineering, and Medicine. 2022. Physics of Life. Washington, DC: The National Academies Press. https://doi.org/10.17226/26403. PREPUBLICATION COPY – SUBJECT TO FURTHER EDITORIAL CORRECTION

The National Academy of Sciences was established in 1863 by an Act of Congress, signed by President Lincoln, as a private, nongovernmental institution to advise the nation on issues related to science and technology. Members are elected by their peers for outstanding contributions to research. Dr. Marcia McNutt is president. The National Academy of Engineering was established in 1964 under the charter of the National Academy of Sciences to bring the practices of engineering to advising the nation. Members are elected by their peers for extraordinary contributions to engineering. Dr. John L. Anderson is president. The National Academy of Medicine (formerly the Institute of Medicine) was established in 1970 under the charter of the National Academy of Sciences to advise the nation on medical and health issues. Members are elected by their peers for distinguished contributions to medicine and health. Dr. Victor J. Dzau is president. The three Academies work together as the National Academies of Sciences, Engineering, and Medicine to provide independent, objective analysis and advice to the nation and conduct other activities to solve complex problems and inform public policy decisions. The National Academies also encourage education and research, recognize outstanding contributions to knowledge, and increase public understanding in matters of science, engineering, and medicine. Learn more about the National Academies of Sciences, Engineering, and Medicine at www.nationalacademies.org. PREPUBLICATION COPY – SUBJECT TO FURTHER EDITORIAL CORRECTION

Consensus Study Reports published by the National Academies of Sciences, Engineering, and Medicine document the evidence-based consensus on the study’s statement of task by an authoring committee of experts. Reports typically include findings, conclusions, and recommendations based on information gathered by the committee and the committee’s deliberations. Each report has been subjected to a rigorous and independent peer-review process and it represents the position of the National Academies on the statement of task. Proceedings published by the National Academies of Sciences, Engineering, and Medicine chronicle the presentations and discussions at a workshop, symposium, or other event convened by the National Academies. The statements and opinions contained in proceedings are those of the participants and are not endorsed by other participants, the planning committee, or the National Academies. For information about other products and activities of the National Academies, please visit www.nationalacademies.org/about/whatwedo. PREPUBLICATION COPY – SUBJECT TO FURTHER EDITORIAL CORRECTION

COMMITTEE ON BIOLOGICAL PHYSICS/PHYSICS OF LIVING SYSTEMS: A DECADAL SURVEY WILLIAM BIALEK, NAS, Princeton University, Chair BRIDGET CARRAGHER. Columbia University IBRAHIM CISSÉ, Max Planck Institute of Immunobiology and Epigenetics MICHAEL DESAI, Harvard University OLGA DUDKO, University of California, San Diego DANIEL GOLDMAN, Georgia Institute of Technology JANE KONDEV, Brandeis University PETER B. LITTLEWOOD, University of Chicago ANDREA J. LIU, NAS, University of Pennsylvania MARY E. MAXON, Lawrence Berkeley National Laboratory JOSÉ N. ONUCHIC, NAS, Rice University MARK SCHNITZER, Stanford University CLARE M. WATERMAN, NAS, National Institutes of Health Staff CHRISTOPHER J. JONES, Senior Program Officer, Study Director STEVEN MOSS, Program Officer NEERAJ P. GORKHALY, Associate Program Officer AMISHA JINANDRA, Associate Program Officer RADAKA LIGHTFOOT, Senior Financial Assistant LINDA WALKER, Program Coordinator COLLEEN N. HARTMAN, Director, Board on Physics and Astronomy JAMES C. LANCASTER, Director, Board on Physics and Astronomy (until April 2021) PREPUBLICATION COPY – SUBJECT TO FURTHER EDITORIAL CORRECTION v

BOARD ON PHYSICS AND ASTRONOMY ANDREW LANKFORD, University of California at Irvine, Chair MEIGAN ARONSON, University of British Columbia WILLIAM BIALEK, NAS, Princeton University JILL DAHLBURG, Naval Research Laboratory (retired) SALLY DAWSON, Brookhaven National Laboratory LOUIS F. DIMAURO, Ohio State University TIM HECKMAN, NAS, Johns Hopkins University WENDELL HILL III, University of Maryland ALAN HURD, Los Alamos National Laboratory CHUNG-PEI MA, University of California, Berkeley ANGELA VILLELA OLINTO, University of Chicago DAVID H. REITZE, California Institute of Technology SUNIL SINHA, University of California, San Diego RISA H. WECHSLER, Stanford University WILLIAM A. ZAJC, Columbia University Staff COLLEEN N. HARTMAN, Director, Board on Physics and Astronomy JAMES C. LANCASTER, Director, Board on Physics and Astronomy (until April 2021) CHRISTOPHER J. JONES, Senior Program Officer GREGORY MACK, Senior Program Officer NEERAJ P. GORKHALY, Associate Program Officer AMISHA JINANDRA, Associate Program Officer MEG KNEMEYER, Financial Officer RADAKA LIGHTFOOT, Senior Financial Assistant LINDA WALKER, Program Coordinator PREPUBLICATION COPY – SUBJECT TO FURTHER EDITORIAL CORRECTION vi

BOARD ON LIFE SCIENCES BARBARA A. SCHALL, NAS, Chair, Washington University in St. Louis A. ALONSO AGUIRRE, George Mason University DENISE N. BAKEN, Shield Analysis Technologies, LLC VALERIE H. BONHAM, Ropes & Gray, LLP PATRICK M. BOYLE, Ginko Bioworks DOMINIQUE BROSSARD, University of Wisconsin, Madison SCOTT V. EDWARDS, NAS, Harvard University GERALD L. EPSTEIN, National Defense University ROBERT J. FULL, University of California, Berkeley BERONDA MONTGOMERY, Michigan State University LOUIS J. MUGILA, Burroughs Wellcome Fund ROBERT NEWMAN, The Aspen Institute LUCILA OHNO-MACHADO, NAM, University of California, San Diego SUDIP S. PARIKH, American Association for the Advancement of Science NATHAN D. PRICE, Onegevity SUSAN RUNDELL SINGER, Rollins College DAVID R. WALT, NAE/NAM, Harvard Medical School PHYLLIS M. WISE, NAM, University of Colorado Staff ANDREW BREMER, Program Officer NANCY D. CONNELL, Senior Scientist JESSICA DE MOUY, Research Associate CYNTHIA GETNER, Financial Business Partner LYLY LUHACHACK, Associate Program Officer STEVEN M. MOSS, Program Officer FRAN SHARPLES, Advisor AUDREY THEVENON, Senior Program Officer DAISHA WALSTON, Program Assistant KAVITA M. BERGER, Director, Board on Life Sciences PREPUBLICATION COPY – SUBJECT TO FURTHER EDITORIAL CORRECTION vii

Preface Every 10 years, the National Academies of Sciences, Engineering, and Medicine (the National Academies) survey the state of physics research in the United States. To make the task manageable, it is broken down by field, so that there are separate reports on astronomy and astrophysics; atomic, molecular, and optical physics; condensed matter and materials physics; elementary particle physics; nuclear physics; and plasma physics. In this cycle, for the first time, biological physics, or the physics of living systems, stands alongside these fields as part of physics. For many of us, this is a moment to celebrate. It is nearly 20 years since the National Science Foundation (NSF) launched a small program to support biological physics within its Physics Division. That effort has evolved into the Physics of Living Systems program. NSF is the sponsor of this survey, charging the committee to take a broad look at the field, its connections to other fields, and the challenges that the community faces in realizing the field’s potential (see Appendix A). Our work began at a meeting on February 6–7, 2020, with presentations from NSF staff, followed by a lively discussion. It is a pleasure to thank Krastan Blagoev, Denise Caldwell, and their colleagues for their support of the project and for their candor. We could not know, of course, that this would also be our last gathering, as the COVID-19 pandemic soon swept through the country and around the world. The February meeting was followed by a second meeting online, April 1–3, 2020. At these two events we heard from a number of colleagues—chosen to complement the expertise of the committee— about the state of science and education in the field: Cliff Brangwynne, Lucy Colwell, Catherine Crouch, Jeff Hasty, Ted Hodapp, Sarah Keller, Chandralekha Singh, Xiaoliang Sunney Xie, and Xiaowei Zhuang. In addition we heard from representatives of other organizations and federal funding agencies beyond NSF, and about the federal budget process more generally, from Linda Horton (Department of Energy Office of Science), Matt Hourihan (American Association for the Advancement of Science), Peter Preusch (National Institute for General Medical Sciences, National Institutes of Health), and Elizabeth Strychalski (National Institute for Standards and Technology). Further insights into an important segment of our audience were provided by Mary Guenther and Alexis Rudd (Senate Committee on Commerce, Science, and Transportation) and by Dahlia Sokolov (House Committee on Science, Space, and Technology). All of these presentations led to significant discussion in the committee, and we appreciate the time and care taken by all of our speakers. A crucial part of the decadal survey process is community input. We solicited written input through the survey website, and held two town hall meetings—one in person at the Biophysical Society Meeting in San Diego (February 16, 2020) and one online through the Division of Biological Physics of the American Physical Society (April 16, 2020). It was wonderful to hear from so many members of the community, speaking from many different perspectives—from undergraduates attending their first scientific conferences and from senior faculty; from researchers in industry, research institutes, and medical schools; from faculty at community colleges, primarily undergraduate institutions, and research universities. As described in the report, it was exciting both to hear such a wide range of voices and to see the emergence of common themes. The work of the committee was done in the two meetings described above and a third (July 27– 29, 2020, also online), along with a series of 28 video conferences; there were numerous meetings of subgroups involved in generating first drafts of different chapters or addressing gaps in subsequent drafts. PREPUBLICATION COPY – SUBJECT TO FURTHER EDITORIAL CORRECTION ix

None of this would have been possible, especially in this challenging time, without the support of the NASEM staff, including Neeraj Gorkhaly, Amisha Jinandra, Steven Moss, Fran Sharples, and Linda Walker. Kim DeRose and Anne Johnson provided guidance on writing style and process. Throughout the project, Radaka Lightfoot managed the budget as plans were continually revised in response to the pandemic. At crucial moments James Lancaster came with excellent advice in his role as Director of the Board on Physics and Astronomy; late in the project he was succeeded by Colleen N. Hartman, who brought fresh eyes, insights, and enthusiasm. Very special thanks to Christopher Jones, who led this effort and provided both wise counsel and gentle reminders of the passing months. This report argues that breadth is an essential part of the excitement in biological physics. The physicist’s approach to understanding the phenomena of life is yielding fascinating results in contexts ranging from the folding of proteins to the flocking of birds, from the internal mechanics of cells to the collective dynamics of neurons in the brain, and more. We see glimpses of the sorts of unifying ideas that we hope for in physics, cutting across this huge range of scales. At the same time, each of these problems also connects to a larger community of biologists, sometimes reaching as far as applications in medicine and technology. Surveying all this required assembling a committee that represents a broad range of interests and expertise, and even so each of us had to stretch to be sure that we could, together, provide our readers with a sense for all of what is exciting in the field. As expected, it has been wonderful fun for all of us on the committee to talk about all these scientific developments. Less expected, perhaps, has been the pleasure of participating in the emergence of consensus on how our excitement about the science translates into recommendations about policy on matters of great concern to the community. The collective effort involved in producing this report has been substantial, especially in the context of the disruptions that all of us have experienced during the pandemic. I think I can speak for the whole committee in expressing thanks to the numerous colleagues, coauthors, friends, and family members who were patient and understanding with us during this time. Finally, let me exercise the chair’s prerogative and add a personal perspective. When I was a student, physicists who became interested in the phenomena of life were perceived as becoming biologists. Physicists and biologists agreed that there were productive applications of physics to biology, but the idea that living systems posed real challenges to our understanding of physics itself was not popular. I don’t think that these views were fair to the history of the field, but they were widely held. Today, much has changed, both in the substance of what has been accomplished and in the perception of these accomplishments, especially by the physics community. The search for the physics of life now is a research program rather than a fantasy, and biological physics has emerged as a branch of physics. This happened not in one dramatic moment, but through decades of progress and gradual realizations. The result is nothing less than a redrawing of the intellectual landscape, the consequences of which continue to unfold in beautiful and sometimes surprising ways. I hope that we have done justice to these remarkable developments. William Bialek, Chair Committee on Biological Physics/Physics of Living Systems: A Decadal Survey PREPUBLICATION COPY – SUBJECT TO FURTHER EDITORIAL CORRECTION x

Acknowledgment of Reviewers This Consensus Study Report was reviewed in draft form by individuals chosen for their diverse perspectives and technical expertise. The purpose of this independent review is to provide candid and critical comments that will assist the National Academies of Sciences, Engineering, and Medicine in making each published report as sound as possible and to ensure that it meets the institutional standards for quality, objectivity, evidence, and responsiveness to the study charge. The review comments and draft manuscript remain confidential to protect the integrity of the deliberative process. We thank the following individuals for their review of this report: Catherine Crouch, Swarthmore College Robert Full, University of California, Berkeley Margaret Gardel, University of Chicago James C. Gumbart, Georgia Institute of Technology Judith Klinman, NAS, University of California, Berkeley Robert Phillips, California Institute of Technology Stephen Quake, NAS/NAE/NAM, Stanford University Elizabeth Strychalski, National Institute of Standards and Technology Kandice Tanner, National Cancer Institute Yuhai Tu, IBM T. J. Watson Research Center Although the reviewers listed above provided many constructive comments and suggestions, they were not asked to endorse the conclusions or recommendations of this report nor did they see the final draft before its release. The review of this report was overseen by Thomas Budinger, Lawrence Berkeley National Laboratory, and Herbert Levine, Northeastern University. They were responsible for making certain that an independent examination of this report was carried out in accordance with the standards of the National Academies and that all review comments were carefully considered. Responsibility for the final content rests entirely with the authoring committee and the National Academies. PREPUBLICATION COPY – SUBJECT TO FURTHER EDITORIAL CORRECTION xi

Contents EXECUTIVE SUMMARY ES-1 INTRODUCTION I-1 Defining the field Connections Challenges A Decadal Survey in Context Findings, Conclusions, and Recommendations Part I: Exploring Big Questions 1 WHAT PHYSICS PROBLEMS DO ORGANISMS NEED TO SOLVE? 1-1 Energy conversion Mechanics, Movement, and the Physics of Behavior Sensing the Environment Structures in Space and Time 2 HOW DO LIVING SYSTEMS REPRESENT AND PROCESS INFORMATION? 2-1 Information Encoded in DNA Sequence Information in Molecular Concentrations Information in the Brain Communication and Language 3 HOW DO MACROSCOPIC FUNCTIONS OF LIFE EMERGE FROM INTERACTIONS AMONG MANY MICROSCOPIC CONSTITUENTS? 3-1 Protein Structure, Folding, and Function Chromosome Architecture and Dynamics Phases and Phase Separation Cellular Mechanics and Active Matter Networks of Neurons Collective Behavior 4 HOW DO LIVING SYSTEMS NAVIGATE PARAMETER SPACE? 4-1 Adaptation Learning Evolution Part II Connections 5 RELATION TO OTHER FIELDS OF PHYSICS 5-1 PREPUBLICATION COPY – SUBJECT TO FURTHER EDITORIAL CORRECTION xiii

6 BIOLOGY AND CHEMISTRY 6-1 Tools for Discovery Molecular and Structural Biology Genes, Genomes, and Evolution Cell and Developmental Biology From Neuroscience to Psychology 7 HEALTH, MEDICINE, AND TECHNOLOGY 7-1 Imaging, Diagnostics, and Treatment Molecular Design Synthetic Biology Predicting and Controlling Evolution Biomechanics and Robotics Neural Networks and AI Part III Realizing the Promise 8 EDUCATION 8-1 Current State of Education in Biological Physics Strengthening Biological Physics Education Postdoctoral Trajectories Summary 9 FUNDING, COLLABORATION, AND COORDINATION 9-1 Current Funding for Research and Education Responding to Challenges and Opportunities User Facilities 10 BUILDING AN INCLUSIVE COMMUNITY 10-1 APPENDIXES A Statement of Task A-1 B Recommendations B-1 C Queries to Funding Agencies C-1 D Agency Missions D-1 E Details Regarding NSF and NIH Grants E-1 F Minimal Support Levels F-1 G Committee Biographies G-1 PREPUBLICATION COPY – SUBJECT TO FURTHER EDITORIAL CORRECTION xiv

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Biological physics, or the physics of living systems, 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. This new field brings the physicist's style of inquiry to bear on the beautiful phenomena of life. The enormous range of phenomena encountered in living systems - phenomena that often have no analog or precedent in the inanimate world - means that the intellectual agenda of biological physics is exceptionally broad, even by the ambitious standards of physics.

Physics of Life is the first decadal survey of this field, as part of a broader decadal survey of physics. This report communicates the importance of biological physics research; addresses what must be done to realize the promise of this new field; and provides guidance for informed decisions about funding, workforce, and research directions.

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