A Strategy for Research in Space Biology and Medicine in the New Century

Committee on Space Biology and Medicine

Space Studies Board

Commission on Physical Sciences, Mathematics, and Applications

National Research Council

NATIONAL ACADEMY PRESS
Washington, D.C.
1998



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--> A Strategy for Research in Space Biology and Medicine in the New Century Committee on Space Biology and Medicine Space Studies Board Commission on Physical Sciences, Mathematics, and Applications National Research Council NATIONAL ACADEMY PRESS Washington, D.C. 1998

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--> NOTICE: The project that is the subject of this report was approved by the Governing Board of the National Research Council, whose members are drawn from the councils of the National Academy of Sciences, the National Academy of Engineering, and the Institute of Medicine. The members of the committee responsible for the report were chosen for their special competences and with regard for appropriate balance. The National Academy of Sciences is a private, nonprofit, self-perpetuating society of distinguished scholars engaged in scientific and engineering research, dedicated to the furtherance of science and technology and to their use for the general welfare. Upon the authority of the charter granted to it by the Congress in 1863, the Academy has a mandate that requires it to advise the federal government on scientific and technical matters. Dr. Bruce Alberts is president of the National Academy of Sciences. The National Academy of Engineering was established in 1964, under the charter of the National Academy of Sciences, as a parallel organization of outstanding engineers. It is autonomous in its administration and in the selection of its members, sharing with the National Academy of Sciences the responsibility for advising the federal government. The National Academy of Engineering also sponsors engineering programs aimed at meeting national needs, encourages education and research, and recognizes the superior achievements of engineers. Dr. William A. Wulf is president of the National Academy of Engineering. The Institute of Medicine was established in 1970 by the National Academy of Sciences to secure the services of eminent members of appropriate professions in the examination of policy matters pertaining to the health of the public. The Institute acts under the responsibility given to the National Academy of Sciences by its congressional charter to be an adviser to the federal government and, upon its own initiative, to identify issues of medical care, research, and education. Dr. Kenneth I. Shine is president of the Institute of Medicine. The National Research Council was organized by the National Academy of Sciences in 1916 to associate the broad community of science and technology with the Academy's purposes of furthering knowledge and advising the federal government. Functioning in accordance with general policies determined by the Academy, the Council has become the principal operating agency of both the National Academy of Sciences and the National Academy of Engineering in providing services to the government, the public, and the scientific and engineering communities. The Council is administered jointly by both Academies and the Institute of Medicine. Dr. Bruce Alberts and Dr. William A. Wulf are chairman and vice chairman, respectively, of the National Research Council. Support for this project was provided by Contract NASW 96013 between the National Academy of Sciences and the National Aeronautics and Space Administration. Any opinions, findings, conclusions, or recommendations expressed in this publication are those of the author(s) and do not necessarily reflect the view of the organizations or agencies that provided support for this project. The cover was designed by Penny Margolskee. Library of Congress Catalog Card Number 98-86544 International Standard book Number 0-309-06047-8 Additional copies of this report are available from: National Academy Press 2101 Constitution Ave., NW Box 285 Washington, DC 20055 800-624-6242 202-334-3313 (in the Washington metropolitan area) http://www.nap.edu Copyright 1998 by the National Academy of Sciences. All rights reserved. Printed in the United States of America

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--> COMMITTEE ON SPACE BIOLOGY AND MEDICINE MARY J. OSBORN, University of Connecticut Health Center, Chair NORMA M. ALLEWELL, University of Minnesota ROBERT E. CLELAND, University of Washington MARY F. DALLMAN,* University of California, San Francisco FRANCIS (DREW) GAFFNEY, Vanderbilt University Medical Center JAMES LACKNER, Brandeis University ANTHONY P. MAHOWALD, University of Chicago ELLIOT MEYEROWITZ, California Institute of Technology LAWRENCE A. PALINKAS, University of California, San Diego KENNA D. PEUSNER, George Washington University Medical Center STEVEN E. PFEIFFER, University of Connecticut Health Center DANNY A. RILEY, Medical College of Wisconsin GIDEON A. RODAN, Merck Research Laboratories RICHARD SETLOW, Brookhaven National Laboratory GERALD SONNENFELD, Carolinas Medical Center T. PETER STEIN, University of Medicine and Dentistry of New Jersey SANDRA J. GRAHAM, Study Director SHOBITA PARTHASARATHY, Research Assistant (until August 1996) CATHY GRUBER, Senior Program Assistant VICTORIA P. FRIEDENSEN, Senior Program Assistant (until April 1996) *   Former member.

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--> COMMITTEE ON SPACE BIOLOGY AND MEDICINE DISCIPLINE PANELS Task Group on the Biological Effects of Space Radiation RICHARD SETLOW, Brookhaven National Laboratory, Chair JOHN F. DICELLO, Johns Hopkins University School of Medicine R.J. MICHAEL FRY, Oak Ridge National Laboratory JOHN B. LITTLE, Harvard University School of Public Health R. JULIAN PRESTON, Chemical Industry Institute of Toxicology JAMES B. SMATHERS, University of California, Los Angeles ROBERT L. ULLRICH, University of Texas Medical Branch at Galveston Panel on Human Behavior LAWRENCE A. PALINKAS, University of California, San Diego, Chair EARL B. HUNT, University of Washington NICK KANAS, University of California, San Francisco, Veterans Affairs Medical Center PETER J. LANG, University of Florida PATRICIA A. SANTY, University of Texas Medical Branch at Galveston PETER SUEDFELD, University of British Columbia

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--> SPACE STUDIES BOARD CLAUDE R. CANIZARES, Massachusetts Institute of Technology, Chair MARK R. ABBOTT, Oregon State University JAMES P. BAGIAN,* Environmental Protection Agency DANIEL N. BAKER, University of Colorado LAWRENCE BOGORAD, Harvard University DONALD E. BROWNLEE, University of Washington GERARD W. ELVERUM, JR., TRW Space and Technology Group ANTHONY W. ENGLAND, University of Michigan MARILYN L. FOGEL, Carnegie Institution of Washington MARTIN E. GLICKSMAN,* Rensselaer Polytechnic Institute RONALD GREELEY, Arizona State University WILLIAM GREEN, former member, U.S. House of Representatives ANDREW H. KNOLL, Harvard University JANET G. LUHMANN,* University of California, Berkeley ROBERTA BALSTAD MILLER, CIESIN BERRIEN MOORE III, University of New Hampshire KENNETH H. NEALSON,* University of Wisconsin MARY JANE OSBORN, University of Connecticut Health Center SIMON OSTRACH, Case Western Reserve University MORTON B. PANISH, AT&T Bell Laboratories (retired) CARLÉ M. PIETERS, Brown University THOMAS A. PRINCE, California Institute of Technology MARCIA J. RIEKE,* University of Arizona PEDRO L. RUSTAN, JR., U.S. Air Force (retired) JOHN A. SIMPSON, Enrico Fermi Institute GEORGE L. SISCOE, Boston University EDWARD M. STOLPER, California Institute of Technology RAYMOND VISKANTA, Purdue University ROBERT E. WILLIAMS, Space Telescope Science Institute MARC S. ALLEN, Director (through December 12, 1997) JOSEPH K. ALEXANDER, Director (as of February 17, 1998) *   Former member.

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--> COMMISSION ON PHYSICAL SCIENCES, MATHEMATICS, AND APPLICATIONS ROBERT J. HERMANN, United Technologies Corporation, Co-chair W. CARL LINEBERGER, University of Colorado, Co-chair PETER M. BANKS, Environmental Research Institute of Michigan WILLIAM BROWDER, Princeton University LAWRENCE D. BROWN, University of Pennsylvania RONALD G. DOUGLAS, Texas A&M University JOHN E. ESTES, University of California, Santa Barbara MARTHA P. HAYNES, Cornell University L. LOUIS HEGEDUS, Elf Atochem North America, Inc. JOHN E. HOPCROFT, Cornell University CAROL M. JANTZEN, Westinghouse Savannah River Company PAUL G. KAMINSKI, Technovation, Inc. KENNETH H. KELLER, University of Minnesota KENNETH I. KELLERMANN, National Radio Astronomy Observatory MARGARET G. KIVELSON, University of California, Los Angeles DANIEL KLEPPNER, Massachusetts Institute of Technology JOHN KREICK, Sanders, a Lockheed Martin Company MARSHA I. LESTER, University of Pennsylvania NICHOLAS P. SAMIOS, Brookhaven National Laboratory CHANG-LIN TIEN, University of California, Berkeley NORMAN METZGER, Executive Director

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--> Foreword The space life sciences occupy a unique niche in the nation's extensive biomedical research enterprise. Only in space is it possible to explore fully the role of gravity on biological systems. In the case of the most complex systems, namely humans, the possible effects of long-term exposure to zero gravity is of more than academic interest. Astronauts have been spending increasing amounts of time in low Earth orbit, extended sojourns in the International Space Station will become routine, and someday humans will likely return to the moon and venture farther. In studies of fundamental biological processes at the cellular or organismic level, the ability to fully manipulate the gravity vector enables a range of studies that cannot be performed in terrestrial laboratories. The cost and complexity of doing any experiment in space demand that careful priorities be set for research. This was done by the National Research Council for space biology and medicine over a decade ago. The present strategy is a complete reformulation of research agendas in the context of current scientific understanding and current or projected opportunities for conducting investigations in space. It is particularly timely given the nation's decision to make a large investment in an orbiting laboratory on the space station. Biological research is a relative newcomer to NASA and still occupies a relatively modest portion of the agency's resources. But there is a growing appreciation of the importance of life sciences within NASA. Outside NASA, space research has often been seen by bench biologists as far from the mainstream of their discipline. However, successful life sciences missions on the space shuttle, joint programs with the National Institutes of Health, and effective peer review have enhanced perceptions about the program. In preparing this report, the Space Studies Board's Committee on Space Biology and Medicine, which itself includes many biologists with little or no connection to space research, convened workshops involving participants drawn widely from the relevant disciplines. The product should help to reinforce the positive trends in both the reality and perceptions about space biology and medicine by providing a science-based assessment of the most important topics to pursue for the decade to come. Claude R. Canizares, Chair Space Studies Board

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--> Preface In 1987, the Committee on Space Biology and Medicine (CSBM) produced a research strategy, A Strategy for Space Biology and Medical Science for the 1980s and 1990s.1 In 1991, the committee's Assessment of Programs in Space Biology and Medicine 19912 examined the National Aeronautics and Space Administration's (NASA's) progress in implementing the 1987 strategy. Since publication of these reports there have been major changes in the direction and status of NASA's life sciences program. The unprecedented amount of biological and medical data gathered from Spacelab missions since 1987 has allowed NASA investigators to move from experiments of an exploratory nature to those that address more fundamental questions. This development has been accompanied by a program shift away from human physiology, the area of major emphasis in the 1987 CSBM report, to more diverse plant and animal studies. As a consequence of these and numerous programmatic changes at NASA, the committee believed that a new strategy, which builds on the current scientific understanding of space biology questions and issues, was needed. After a series of discussions with NASA's Life Sciences Division, the committee agreed to undertake a comprehensive review of the status of research in the various fields of space life sciences and to develop a science strategy that could guide NASA in its long-term research and mission planning. This study was carried out over a 3-year period, and its objectives remained the same as those outlined in the 1987 report: "(1) to identify and describe those areas of fundamental scientific investigation in space biology and medicine that are both exciting and important to pursue and (2) to develop 1   Space Science Board, National Research Council. 1987. A Strategy for Space Biology and Medical Science for the 1980s and 1990s. National Academy Press, Washington, DC. 2   Space Studies Board, National Research Council. 1991. Assessment of Programs in Space Biology and Medicine 1991. National Academy Press, Washington, D.C.

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--> the foundation of knowledge and understanding that will make long-term manned space habitation and/or exploration feasible."3 Specifically, the committee attempted to provide the following in this report: A review of the disciplines of biology and medicine that can usefully be studied in the space environment, including sciences that study plant, animal, and human systems at the molecular, cellular, system, and whole-organism levels; Discussion of the fundamental research issues and questions within these disciplines; Identification of the most promising experimental challenges and opportunities within each discipline; Evaluation of the potential for space research to provide advances within each discipline; and Prioritization of research topics to the extent feasible. In addition to numerous expert speakers from NASA and academia, who were invited to give presentations at regular committee meetings, the CSBM used a variety of approaches to gather information for its task. Three workshops were organized by the committee, each focusing on a broad life sciences discipline, and both NASA and non-NASA investigators were invited to participate. The committee also sent delegates to several international life sciences workshops organized by NASA and its international partners. Each workshop was directed at reviewing progress in a specific discipline and included participation by space life sciences investigators from around the world. Of course, the committee also reviewed both NASA source materials and the relevant literature, published and online, on flight- and ground-based research. Separate discipline panels, each chaired by a member of the CSBM, were developed to review and discuss the areas of space radiation and human behavioral studies. These two groups were given responsibility for drafting the sections of this report representing their disciplines, although the final report is the responsibility of the committee as a whole. As originally planned, the recommendations and analysis developed by the Task Group on the Biological Effects of Space Radiation and published separately in 19964 form the basis of Chapter 11, "Radiation Hazards," in CSBM's new strategy for research. 3   Space Science Board, 1987, A Strategy for Space Biology and Medical Science for the 1980s and 1990s, p. XI. 4   Space Studies Board. 1996. Radiation Hazards to Crews of Interplanetary Missions: Biological Issues and Research Strategies. National Academy Press, Washington, D.C.

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--> Acknowledgment of Reviewers This report has been reviewed by individuals chosen for their diverse perspectives and technical expertise, in accordance with procedures approved by the National Research Council's (NRC's) Report Review Committee. The purpose of this independent review is to provide candid and critical comments that will assist the authors and the NRC in making the published report as sound as possible and to ensure that the report meets institutional standards for objectivity, evidence, and responsiveness to the study charge. The contents of the review comments and draft manuscript remain confidential to protect the integrity of the deliberative process. We wish to thank the following individuals for their participation in the review of this report: S. James Adelstein, Harvard Medical School, Robert M. Berne, University of Virginia, Joseph V. Brady, Johns Hopkins University, Robert R. Burris, University of Wisconsin-Madison, Robert A. Frosch, Harvard University, Sally K. Frost-Mason, University of Kansas, Ursula W. Goodenough, Washington University, J. Richard Hackman, Harvard University, Jack P. Landolt, Defence and Civil Institute of Environmental Medicine, Ontario, Canada, Philip Osdoby, Washington University, Robert O. Scow, National Institute of Diabetes and Digestive and Kidney Diseases, and Frank A. Witzman, Indiana University Purdue University-Columbus. Although the individuals listed above have provided many constructive comments and suggestions, responsibility for the final content of this report rests solely with the authoring committee and the NRC.

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--> Contents EXECUTIVE SUMMARY   1 PART I—OVERVIEW     1   INTRODUCTION   21     History   22     Gravity and Low Gravity   23     Gravity, Microgravity, and Weightlessness   23     Direct and Indirect Effects of Microgravity   24     References   25 PART II—PHYSIOLOGY, GRAVITY, AND SPACE     2   CELL BIOLOGY   29     Introduction   29     Previous Cell Biological Research in Space   31     Opportunities for NASA-supported Research in Cell Biology   33     Mechanisms of Cellular Response to Mechanical Force   33     Cellular Response to Environmental Stress   34     Development of Advanced Instrumentation and Methodologies   34     References   35

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--> 3   DEVELOPMENTAL BIOLOGY   37     Introduction   37     Progress in Developmental Biology   38     Developmental Genetics   38     Molecular Conservation   38     Genome Sequencing Project   39     Major Issues in Space Developmental Biology   40     Complete Life Cycles in Microgravity   40     Development of the Vestibular System   41     Neural Space Maps   42     Neuroplasticity   43     References   45 4   PLANTS, GRAVITY, AND SPACE   49     Introduction   49     Space Horticulture   49     Reasons for Studies on Space Horticulture   49     Accomplishments   50     Future Directions   51     Role of Gravity in Plant Processes   52     Scientific Problems   52     Accomplishments   53     Future Directions   54     Response of Plants to a Change in the Direction of the Gravity Vector   55     Known Responses   55     Gravitropism   55     Gravitaxis   60     Effects of Gravity-induced Tissue Stresses on Plant Development   60     References   60 5   SENSORIMOTOR INTEGRATION   63     Introduction   63     Spatial Orientation   63     Posture and Locomotion   66     Vestibulo-Ocular Reflexes and Oculomotor Control   68     Vestibular Processing During Microgravity   70     Space Motion Sickness   71     Central Nervous System Reorganization   73     Teleoperation and Telepresence   73     General Strategic Issues   74     References   74 6   BONE PHYSIOLOGY   80     Introduction   80     Bone Functions, Growth and Development, and Remodeling   80     Functions of Bone   80

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-->     Bone Growth and Development   81     Bone Remodeling: Hormonal Effects   82     Mechanical Effects on Bone Remodeling   84     Clinical Observations and Human Experimentation   84     Animal Studies   85     Putative Mechanisms   86     Microgravity Effects on the Skeleton   87     Caveats   87     Human Studies   88     Animal Studies   90     Equipment Needs   92     References   93 7   SKELETAL MUSCLE   97     Introduction   97     Background   98     Research Done on Muscle Biology   98     Previous Space- and Ground-based Research   99     Primary In-flight Changes   100     Simple Deconditioning and Adaptation   100     Pathological Alteration and Metabolic Adaptation   101     Contractile Physiology, Contractile Proteins, and Myofilaments   101     Preservation of Function During Atrophy   102     Reentry- and Reloading-induced Secondary Changes   103     Movements in Space and Upon Return to Earth   103     Compromised Microcirculation   103     Increased Susceptibility to Structural Damage   104     Cellular and Molecular Mechanisms   106     References   108 8   CARDIOVASCULAR AND PULMONARY SYSTEMS   118     Introduction   118     Cardiovascular Physiology in Microgravity   119     Pulmonary Physiology in Microgravity   121     Postflight Cardiovascular Physiology   123     In-flight Countermeasures   124     Future Directions   125     Cardiopulmonary Equipment   126     Research   127     References   128 9   ENDOCRINOLOGY   132     Introduction   132     Current Status of Research   133     Effects of Spaceflight on Humans   133     Hypothalmic-Pituitary-Adrenal Axis   133

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-->     Models   135     Energy Metabolism and Balance   136     Reproduction   137     Fluid and Electrolyte Balance   138     Hormonal Systems and Changes   138     Models   139     Hematology   140     In-flight Observations   140     Models   141     Endocrine Aspects of Muscle Loss   141     Hormones Involved   142     Models   144     Bone   144     Circadian Rhythms   145     Gender   145     References   146 10   IMMUNOLOGY   156     Introduction   156     Spaceflight Experiments   157     Animal Studies   157     Human Studies   160     Cell Culture Studies   162     Ground-based Models of the Effects of Spaceflight on Immune Responses   162     References   163 PART III—ADDITIONAL SPACE ENVIRONMENT ISSUES     11   RADIATION HAZARDS   171     Introduction   171     Statement of the Problem   172     Current Understanding of Biological Effects of Radiation   174     Types of Effects   174     Effects Induced by Protons and Heavy Ions   175     Priority Research Recommendations and Strategies   177     Higher-Priority Research Recommendations   177     Lower-Priority Research Recommendations   182     Time Scale of Research   186     Need for Animal Use   186     Experimental Techniques and New Data Required   186     Ground- versus Space-based Research   190     References   190

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--> 12   BEHAVIORAL ISSUES   194     Introduction   194     Program History   194     Statement of Goals   195     Definition and Assessment of Behavior and Performance in Space   195     Research in Analogue Environments   196     Integration of Research and Operations   197     Organizational Support of Research   197     Environmental Factors   198     Environmental Conditions Unique to Spaceflight   198     Environmental Conditions Common to Isolated, Confined Environments   199     Psychophysiological Issues   201     Circadian Rhythms and Sleep   201     The Psychophysiology of Emotion and Stress   203     Psychophysiological Measurement in Space   204     Individual Issues   206     Psychological Issues   206     Psychiatric Issues   210     Countermeasures   211     Interpersonal Issues   213     Crew Tension and Conflict   213     Crew Cohesion   214     Ground-Crew Interaction   215     Leadership Role   216     Countermeasures   216     Organizational Issues   219     Organizational Culture   219     Mission Duration   220     Management   220     References   222 PART IV—RESEARCH PRIORITIES AND PROGRAMMATIC ISSUES     13   SETTING PRIORITIES IN RESEARCH   231     Physiological and Psychological Effects of Spaceflight   232     Loss of Weight-bearing Bone and Muscle   232     Vestibular Function, the Vestibular Ocular Reflex, and Sensorimotor Integration   232     Orthostatic Intolerance Upon Return to Earth Gravity   233     Radiation Hazards   233     Physiological Effects of Stress   234     Psychological and Social Issues   234     Fundamental Gravitational Biology   235     Mechanisms of Graviperception and Gravitropism in Plants   235     Mechanisms of Graviperception in Animals   235     Effects of Spaceflight on Reproduction and Development   236

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--> 14   PROGRAMMATIC AND POLICY ISSUES   237     Space-based Research   238     Criteria for Space Research   238     Development of Advanced Instrumentation and Methodologies   239     Utilization of the International Space Station for Life Sciences Research   241     Science Policy Issues   242     Peer Review   242     Integration of Research Activities   242     Human Flight Data: Collection and Access   244     Publication and Outreach   245     Professional Education   246     References   247 APPENDIXES     A   Acronyms and Abbreviations   251 B   Glossary   253 C   Workshops   266 D   Committee Biographies   272

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A Strategy for Research in Space Biology and Medicine in the New Century

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