Biological Effects and Physics of Solar and Galactic Cosmic Radiation: Part A PDF
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Biological Effects and Physics of Solar and Galactic Cosmic Radiation Part A NATO ASI Series Advanced Science Institutes Series A series presenting the results of activities sponsored by the NATO Science Committee, which aims at the dissemination of advanced scientific and technological knowledge, with a view to strengthening links between scientific communities. The series is published by an international board of publishers in conjunction with the NATO Scientific Affairs Division A Life Sciences Plenum Publishing Corporation B Physics New York and London C Mathematical and Physical Sciences Kluwer Academic Publishers D Behavioral and Social Sciences Dordrecht, Boston, and London E Applied Sciences F Computer and Systems Sciences Springer-Verlag G Ecological Sciences Berlin, Heidelberg, New York, London, H Cell Biology Paris, Tokyo, Hong Kong, and Barcelona I Global Environmental Change Recent Volumes in this Series Volume 243A —Biological Effects and Physics of Solar and Galactic Cosmic Radiation, Part A edited by Charles E. Swenberg, Gerda Horneck, and E. G. Stassinopoulos Volume 243B —Biological Effects and Physics of Solar and Galactic Cosmic Radiation, Part B edited by Charles E. Swenberg, Gerda Horneck, and E. G. Stassinopoulos Volume 244 —Forest Development in Cold Climates edited by John Alden, J. Louise Mastrantonio, and Soren 0dum Volume 245 —Biology of Salmonella edited by Felipe Cabelol Volume 246 —New Developments in Lipid-Proteni Interactions and Receptor Function edited by K. W. A. Wirtz, L. Packer, J. A. Gustafsson, A. E. Evangelopoulos, and J. P. Changeux Volume 247—Bone Circulation and Vascularization in Normal and Pathological Conditions edited by A. Schoutens, J. Arlet, J. W. M. Gardeniers, and S. P. F. Hughes Series A: Life Sciences Biological Effects and Physics of Solar and Galactic Cosmic Radiation Part A Edited by Charles E. Swenberg Armed Forces Radiobiology Research Institute Bethesda, Maryland and Complexity Incorporated Potomac, Maryland Gerda Horneck DLR Institute of Aerospace Medicine Cologne, Germany and E. G. Stassinopoulos NASA-Goddadr Space Flight Center Greenbelt, Maryland Springer Science+Business Media, LLC Proceedings of a NATO Advanced Study Institute on Biological Effects and Physics of Solar and Galactic Cosmic Radiation, held October 13-23, 1991, in Algarve, Portugal NATO-PCO-DAA TBASE The electronic index to the NATO ASI Series provides full bibliographical references (with keywords and/or abstracts) to more than 30,000 contributions from international scientists published in all sections of the NATO ASI Series. Access to the NATO-PCO-DATA BASE is possible in two ways: —via online FILE 128 (NATO-PCO-DATA BASE) hosted by ESRIN, Via Galileo Galilei, I-00044 Frascati, Italy —via CD-ROM "NATO-PCO-DATA BASE" with user-friendly retrieval software in English, French, and German ( ©WTV GmbH and DATAWARE Technologies, Inc. 1989) The CD-ROM can be ordered through any member of the Board of Publishers or through NATO-PCO, Overijse, Belgium. Library of Congress Cataloglng-ln-PublIcat1on Data NATO Advanced Study Institute on Biological Effects and Physics of Solar and Galactic Cosmic Radiation (1991 : Algarve, Portugal) Biological effects and physics of solar and galactic cosmic radiation. Part A / edited by Charles E. Swenberg, Gerda Horneck, and E.G. StassInopoulos. k p. cm. — (NATO advanced science institutes series. Series A, Life sciences ; v. 243A) "Proceedings of a NATO Advanced Study Institute on Biological Effects and Physics of Solar and Galactic Cosmic Radiation, held October 13-23, 1991 in Algarve, Portugal"—T.p. verso. Includes bibliographical references and index. ISBN 978-1-4613-6266-1 ISBN 978-1-4615-2918-7 (eBook) DOI 10.1007/978-1-4615-2918-7 1. Radiation—Toxicology—Animal models—Congresses. 2. Radiation—Toxicology—Mathematical models—Congresses. 3. Outer space—Exploration—Health aspects—Congresses. I. Swenberg, Charles E. II. Horneck, G. (Gerda) III. Stassinopou1os, E. G. IV. Title. V. Series. RC1151.R33N38 1991 616.9'897—dc20 93-8450 CIP Additional material to this book can be downloaded from http://extra.springer.com. ©1993 Springer Science+Business Media New York Originally published by plenum Press, New York in 1993 Softcover reprint of the hardcover 1st edition 1993 All rights reserved No part of this book may be reproduced, stored in retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording, or otherwise, without written permission from the Publisher PREFACE Space missions subject human beings or any other target of a spacecraft to a radiation environment of an intensity and composition not available on earth. Whereas for missions in low earth orbit (LEO), such as those using the Space Shuttle or Space Station scenario, radiation exposure guidelines have been developed and have been adopted by spacefaring agencies, for exploratory class missions that will take the space travellers outside the protective confines of the geomagnetic field sufficient guidelines for radiation protection are still outstanding. For a piloted Mars mission, the whole concept of radiation protection needs to be reconsidered. Since there is an increasing interest ci many nations and space agencies in establishing a lunar base and lor exploring Mars by manned missions, it is both, timely and important to develop appropriate risk estimates and radiation protection guidelines which will have an influence on the design and structure of space vehicles and habitation areas of the extraterrestrial settlements. This book is the result of a multidisciplinary effort to assess the state of art in our knowledge on the radiation situation during deep space missions and on the impact of this complex radiation environment on the space traveller. ]t comprises the lectures by the faculty members as well as short contributions by the students given at the NATO Advanced Study Institute "Biological Effects and Physics of Solar and Galactic Cosmic Radiation" held in Armacao de Pera, Portugal, 12-23 October, 1991. The following scientists served on the Organizing Committee: C. E. Swenberg, Armed Forces Radiobiology Research Institute, Bethesda, Maryland, USA G. Horneck, Deutsche Forschungsanstalt fUr Luft-und Raumfahrt, Koln, Germany E.G. Stassinopoulos, NASA Goddard Space Right Center, Greenbelt, Maryland, USA P.D. McCormack, US Naval Medical Center, Washington D.C., USA The participants, coming from various countries including Russia, Ukraine, Czechoslovakia, Bulgaria are listed at the end of this book. The event is in many respects a sequel to the NATO Advanced Study Institute "Terrestrial Space Radiation and its Biological Effects", Corfu, Greece, October 1987, which was mainly concerned with radiation problems for manned missions in Low Earth Orbit (LEO). During this meeting, it was emphasised that in order to safeguard future human enterprises in space, especially those of very long duration or beyond our geomagnetic shield, an intense research program has to be initiated. The program objectives should include: (1) with respect to the radiation environment in deep space missions: development of a better physical model of the galactic cosmic radiation modulation as a function of solar cycle observables; development of models of HZE particles propagation in the interplanetary medium; a better understanding on the periodicity (magnitude, duration) of a solar cycle, in order to make a prediction several years in advance; a standardized approach on an international scale for predicting/forecasting solar flares that give rise to proton events and an v efficient warning system; microdosimetric approaches in future development of space dosimetry; determination of the dose contributing products as a function of shielding; testing of new space technologies with respect to their vulnerability to space radiation; (2) with respect to biological responses to the radiation in space: selection of appropriate biological test systems for radiobiological space experiments in order to quantify and qualify various long-term biological radiation effects; analysis of the biological responses to single particle traversals; development of biological dosimeters; a better understanding of the radiobiological chain of events as a function of radiation characteristics from the initial interactions altering the essential chemical processes, such as DNA strand breaks to the biological response, e.g. cellular lethality, mutagenesis, transformation; investigation of transeffects, i. e. radiation lesions in the DNA that may lead to changes at remote sites; development of radiobiological models appropriate for determining biological effects of HZE particles; international cooperation in the analysis of radiation effects in higher organisms and humans including data obtained in space; (3) with respect to risk estimates for deep space missions: development of new and more appropriate concepts to quantitate the radiation risk in space missions; development of shielding concepts; development of countermeasures including radioprotectants, nutritional supplements; determination of the radiation tolerances of different individuals. This list is by no means complete. It reflects the need for a long-term program where ground based studies will be augmented by flight experiments, especially in high inclination orbits or on precursor missions to Moon and Mars. It was considered to be extremely important to reach a standardisation on an international level with respect to data collection, protocol comparison and formulation of guidelines for future exploratory class missions. The committee is most grateful to the North Atlantic Treaty Organization for the outstanding support provided for this meeting and for the production of this monograph. It also acknowledges substantial financial support provided by German Aerospace Research Establishment DLR, The US Armed Forces Radiobiology Research Institute, Bethesda MD, the US Department of Energy, and the Committee on Interagency Radiation Research and Policy Coordination in cooperation with Oak Ridge Associated Universities. We thank Lisa Steimel for her valuable and efficient assistance in assuring that the meeting was truly successful. The editors acknowledge the advice and guidance of Mr. Gregory Safford of Plenum Press and the assistance of Dr. Mei-Lie Swenberg in typing and reorganizing the manuscripts to the final version. Charles E. Swenberg Gerda Horneck E. G. Stassinopoulos vi CONTENTS RADIATION EFFECTS ON BIOLOGICAL SYSTEMS Radiation-Induced DNA Lesions in Eukaryotic Cells, Their Repair and Biological Relevance .......... . 1 M. Frankenberg-Schwager Assessment of Heavy Ion Induced DNA Strand Breaks in Mammalian Cells 33 J. Heilmann and H. Rink Does the Topology of Closed Supercoiled DNA Affect Its Radiation Sensitivity? 37 C. E. Swenberg, J. M. Speicher and J. H. Miller Plasmids as Testsystem for the Detection of DNA Strand Breaks 49 J. Wehner, G. Homeck and H. BUcker High Resolution Gel Electrophoresis Methods for Studying Sequence-Dependence of Radiation Damage and Effects of Radioprotectants in Deoxyoligonucleotides ................ . 53 B. Mao, C. E. Swenberg, Y. Vaishnav and N. E. Geacintov Thymine Dimer Formation Mediated by the Photosensitizing Properties of Pharmaceutical Constituents . . . . . . . . . . . . . . . . 63 L. F. Salter, B. S. Martincigh, K. Bolton, S. R. Aliwell and S. J. Clemmett The Assay of 2'Deoxyuridine-3' -monophosphate (dUMP) by 3' Phosphorylation and Two Dimensional Thin Layer Chromatography: A Potential Radiation Marker of DNA Damage 71 A. Cajigas and J.J. Steinberg Free-Radical Yields in Proton Irradiation of Oriented DNA: Relationship to Energy Transfer along DNA Chains 85 J. H. Miller, D. L. Frasco, M. Ye, C. E. Swenberg, L. S. Myers, Jr., and A. Rupprecht Free Radical Formation in Amino Acids Induced by Heavy Ions 93 T. Heck, J. HUttermann and G. Kraft The Biostack Concept and Its Application in Space and at Accelerators: Studies on Bacillus subtilis Spores ....... . 99 G. Homeck Biological Action of Single Heavy Ions on Individual Yeast Cells 117 M. Kost and J. Kiefer vii Repair of Radiation Induced Genetic Damage under Microgravity: An Experimental Proposal 125 0 0 0 0 0 0 0 0 0 0 0 H.-Do Pross, Mo Kost and Jo Kiefer DNA Damage Repair and Mutagenesis in Mutants of Escherichia coli 129 Sol. Ahmad Neoplastic Transformation Induced by Heavy Ions: Studies on Transformation Efficiencies and Molecular Mechanisms 135 0 0 0 0 0 0 0 0 L. Hieber, Jo Smida and Ao Mo Kellerer On the Mechanism and Consequences of Radiation-Induced Gene Amplification in Mammalian Cells 143 0 0 0 0 Co Lticke-Huhle Alteration in Lipid Peroxidation in Plant Cells after Accelerated Ion Irradiation 155 Ao Vasilenko, So Zhadko and Po Go Sidorenko Tumor Induction in Animals and the Radiation Risk for Man 161 1. Jo Broerse, 1. Davelaar and C. Zurcher Progress in the Extrapolation of Radiation Cataractogenesis Data across Longer-Lived Mammalian Species 177 0 0 0 0 0 Ao Bo Cox, Yo L. Salmon, Ao Co Lee, 1. To Lett, Go R. Williams, Jo Jo Broerse, Go Wagemaker and Do Do Leavitt Effects of Linear Energy Transfer on the Formation and Fate of Radiation Damage to the Photoreceptor Cell Complement of the Rabbit Retina: Implications for the Projected Manned Mission to Mars 185 0 0 0 0 0 0 0 Jo To Lett and Go R. Williams Effects of Neon Ions, Neutrons and X Rays on Small Intestine 203 K. Eo Carr, Jo So McCullough, R. St. Co Gilmore, Bo Abbas, So Po Hume, Ao Co Nelson, To L. Hayes and Eo Jo Ainsworth Mucopolysaccharide Vascular Coating Relationship to Environmental Factors 217 Do Eo Philpott and K. Kato An Evaluation of the Relative Behavioral Toxicity of Heavy Particles 227 0 0 0 Bo Mo Rabin, Wo Ao Hunt, Jo Ao Joseph, To K. Dalton and So Bo Kandasamy THEORETICAL MODELS OF BIOLOGICAL RADIATION ACTION A Model of Cell Damage in Space Right 235 0 0 0 0 0 0 0 0 0 0 0 0 0 R. Katz, F. Ao Cucinotta, Jo Wo Wilson, Jo L. Shinn and Do Mo Ngo Concepts of Microdosimetry and Their Applicability to Radiation Protection Problems in Manned Space Missions 269 0 0 0 0 0 0 0 0 0 0 0 Jo Breckow Theoretical Analysis of Heavy Ion Action on Cells: Model-Free Approaches, Consequences for Radiation Protections 283 0 0 0 0 0 0 0 0 0 Jo Kiefer Mathematical Models of Lesion Induction and Repair in Irradiated Cells 291 So Kozubek and Go Homeck Cell Kinetics and Track Structure 295 0 0 0 0 0 0 Jo Wo Wilson, F. Ao Cucinotta and Jo L. Shinn Index 339 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 viii RADIATION-INDUCED DNA LESIONS IN EUKARYOTIC CELLS, THEIR REPAIR AND BIOLOGICAL RELEVANCE M. Frankenberg-Schwager GSF: Forschungszentrum fUr Umwelt und Gesundheit, Institut fUr Biophysikalische Strahlenforschung, Paul-Ehrlich-Str. 20 D-6OOO Frankfurt 70, FRG INTRODUCTION A variety of lesions can be detected in the DNA of eukaryotic cells irradiated with ioniring radiation. These include DNA- protein crosslinks (DPCs), base alterations and base detachments, sugar alterations, bulky lesions, i.e. clusters of base damage, DNA single and double-strand breaks. For most of these lesions, i. e. DNA-protein crosslinks, base damage, single-and double-strand breaks and bulky lesions, the kinetics of enzymatic repair has been studied. The influence of various factors, such as oxia/anoxia,linear energy transfer (LET) of the radiation used, incubation medium, cell-cycle stage, cell age, thiol content, hyperthermia,on the induction and repair of these lesions is described and their biological relevance is outlined. Radiation-sensitive cell lines are also included. This paper is an extended and updated version of two earlier publications (Frankenberg-Schwager, 1989 and 1990) with special emphasis on the effect of LEf. RADIATION-INDUCED DNA-PROTEIN CROSSLINKS Induction of DNA- Protein Crosslinks DNA-protein crosslinks are formed by covalent linkage between DNA and proteins of the nuclear matrix (e.g. Oleinick et al., 1986). Mainly those DNA regions which contain actively transcribed, and presumably also replicated, sequences are involved in the linkage to proteins, (e.g. Chiu et al., 1982, Oleinicket al., 1986}.There is evidence that these DNA sequences are associated with the nuclear matrix ( Ciejek et al., 1983, Robinson et al., 1983, Mc Cready et al., 1982). A relatively high amount of about 6000 DPCs is observed per normal, unirradiated cell of Chinese hamster V79 lung fibroblasts in exponential growth (Oleinick et al' , 1986). DPCs are induced linearly with radiation dose in the range of 10 to 100 Gyas shown for exponentially growing leukemia mouse cells (Cress and Bowden ,1983)and Chinese hamsterV79 cells (Fig. I) (Chiu et al. ,1984 , Oleinicket al., 1986 }.y-rays increase the number of new linkages between DNA and protein at a frequency of 150 DPCs per Gy per V79 cell (Ramakrishnan et al' ,1987). Thus DPCs are induced at about a 4 time higher frequency than DNA double-strand breaks (Ramakrishnan et al" 1987). The same yield of radiation-induced DPCs is found in normal and radiation-sensitive cell lines (like Ataxia telangiectasia (AT) cells and the Chinese hamster CHO mutant XTS )(Oleinick et aI., 1990). At higher doses the number of DPCs approaches a plateau value which corresponds to the Biological Effects and Physics of Solar and Galactic Cosmic Radiation, Part A. Edited by C.B. Swenberg et al. • Plenum Press. New York, 1993 • ~ § ~ !! , ·s ) ·5 .!! ~ 2 ~ 15 30 45 60 75 90 1a; Dose/Gy Figure 1. Induction of DNA-protein Crosslinks in Exponentially Growing Chinese Hamster V79 cells Exposed to 6OCo-y rays (Redrawn from Chiu et al. 1984). number of DNA attachment sites to the nuclear matrix (Ramakrishnan et al. ,1987). The yield of DPCs in irradiated V79 cells is reduced in the presence of cysteamine (Radford ,1986a). Based on the protective effect of the OH radical scavenger dimethylsulphicoxide (DMSO), which is more pronounced in active DNA than in bulk DNA(Chiu et al., 1986), it appears that OH radicals are responsible for the production of DPCs in V79 cells (Oleinick et al., 1986). The yield of DPCs is enhanced when cells are irradiated under hypoxic conditions as shown fora human fibroblast cell line (Fomace and Little, 1977), for CHO cells (Meyn and Jenkins, 1984) and for mouse L and Chinese hamster V79 cells (Radford, 1986a). Interestingly, DPCs are detected in cells from tissues irradiated in situ in acutely hypoxic (N2 - asphyxiated) mice and even in air-breathing animals (Meyn et al., 1986), indicating the presence of radio biological hypoxic cells in tissues of air-breathing animals. Preirradiation hyperthermia (430C for 15 min or longer) has no effect on the yield of radiation-induced DPCs in exponentially growing mouse leukemia cells (Cress and Bowden,1983)and in V79 cells (Radford, 1986a), suggesting that the increased amount of nuclear protein after hyperthermia (Roti-Roti and Winward, 1981) is not available for covalent bonding to the DNA damaged by ionizing radiation. In contrast, pre-irradiation hyperthermia increased the yield of DPCs in mouse L cells (Radford, 1986a). Repair of DNA-Protein Crosslinks Mammalian cells are capable of repairing radiation-induced DPCs. DNA is removed from protein with biphasic kinetics during postirradiation incubation of cells at 370 C under growth conditions. Human fibroblasts in stationary phase from a confluent monolayer irradiated with 50 Gy under oxic or anoxic conditions exhibit a rapid component of repair of DPCs with a half-time constant t 112 of about 2 h and a slow component with at 1I2-value of 12-13 h (Fomace and Little ,1977). In actively transcribed DNA sequences. DPCs are not only preferentially induced but also the removal of DPCs in these regions is faster as compared to nontranscribed DNA (Oleinick et al.,I986). In agreement with these findings is the fast repair of DPCs in exponentially growing monolayer cultures of V79 cells irradiated with 60 Gy. The t1l2- value reported for the rapid component is 1 h which is followed by a slower \component of repair (Chiu et al.,l984). Likewise, for growing mouse leukemia cells 2
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