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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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