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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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