ILS I ILSI Human Nutrition Reviews

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2 ILS I ILSI Human Nutrition Reviews INTERNATIONAL LIFE SCIENCES INSTITUTE Series Editor: Ian Macdonald Already published: Sweetness Edited by John Dobbing Forthcoming titles in the Series: Sucrose: Nutritional and Safety Aspects Edited by Gaston Vettorazzi and Ian Macdonald Zinc in Human Biology Edited by C. F. Mills

3 B. E. C. Nordin (Ed.) Calcium in Human Biology With 110 Figures Springer-Verlag London Berlin Heidelberg New York Paris Tokyo

4 B. E. C. Nordin Institute of Medical and Veterinary Science, Adelaide, South Australia 5000, Australia Cover design permission. The calcium triangle within the calcium hexagons showing the position ofthe phosphate groups (not to scale and a projection on a flat surface of ions arranged three-dimensionally) is reproduced by permission from Jenkins G.N. The Physiology and Biochemistry of the Mouth 4th ed (1978), Blackwell Scientific Publications, Oxford ISBN-13: e-isbn-13: DOl: / British Library Cataloguing in Publication Data Nordin, B.E.C. (Borje Edgar Christopher) Calcium in human biology. - (ILSI human nutrition reviews). 1. Man. Calcium & phosphorous. Metabolism I. Title II. Series 612'.3924 Library of Congress Cataloging-in-Publication Data Calcium in human biology / B. E. C. Nordin (ed.). p. cm. - (ILSI human nutrition reviews) Includes bibliographies and index. 1. Calcium-Physiological ~ffect. 2. Calcium-Metabolism. I. Nordin, B. E. C. (Borje Edgar Christopher) II. Series. [DNLM: 1. Calcium-metabolism. QV 276 C1436] QP535.C2C '.3924-dc 19 DNLMlDLC for Library of Congress CIP This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, re-use of illustrations, recitation, broadcasting, reproduction on microfilms or in other ways, and storage in data banks. Duplication of this publication or parts thereof is only permitted under the provisions of the German Copyright Law of September 9, 1965, in its version of June 24, 1985, and a copyright fee must always be paid. Violations fall under the prosecution act of the German Copyright Law. Springer-Verlag Berlin Heidelberg 1988 Softcover reprint of the hardcover 1st edition 1988 The use of names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. Product Liability: The publisher can give no guarantee for information about drug dosage and application thereof contained in this book. In every individual case the respective user must check its accuracy by consulting other pharmaceutical literature. Filmset by Wilmaset, Birkenhead, Wirral 2128/

5 Preface It is a sign of the times that a work designed to review the role of calcium in human biology should have devoted so much of its space to cellular calcium. Only five years ago the internal balance of a work such as this would have been quite different. However, with a realization that calcium is actively involved in virtually every biological process has come the need for all of us to understand or at least to respect its role as a messenger. No doubt it is this ubiquitous role that has necessitated its almost uniquely precise extracellular regulation, which in turn relies on the virtually unlimited stores in the skeleton to cope with intra- and inter-individual variation in calcium intake and absorption. The fact that the skeleton's humble role as a source of calcium sooner or later comes into conflict with its more obvious mechanical role is of little interest to the evolutionary process, since the final skeletal failure, when it occurs, occurs too late in life to influence the evolutionary pattern. No one author could be expected to command the expertise which this volume required, but neither can 17 be brought together to contribute the individual components without unevenness and overlap occurring between the chapters. At least, or so it seems to the Editor, there are more repetitions than gaps, and if the repetitions serve to reinforce important messages they will have served their turn. They could hardly have been avoided without injury to the smooth flow of the argument in one or other chapter and so they have been left untouched, even to the extent of the same figure appearing in almost identical guise in two places. The Editor is not responsible for all the opinions or statements in this volume; in fact, there are some with which he sharply disagrees. But editorship is not the same as censorship and views honestly held and based on respectable data must be respected, even if some would interpret the data differently from others. I am grateful to the International Life Sciences Institute for asking me to edit this volume and hope that it will prove useful to a

6 vi Preface wide readership. It is not to be equated with the proceedings of a scientific symposium. Although the material is up to date it is not, we hope, ephemeral but rests on a solid foundation of wellestablished knowledge. At least, this was the remit to the contributors which I hope they have discharged. Adelaide September 1987 B. E. C. Nordin

7 Contents Chapter 1. Chemistry and Biochemistry of Calcium W. G. Robertson... 1 Introduction Evolutionary Aspects of Calcium... 2 Chemical Properties Extracellular Calcium Intracellular Calcium Calcium as a Second (or Third?) Messenger Calcium Ionophores Calcium Antagonists Calcium in Other Living Organisms Chapter 2. Calcium-Regulating Hormones: Vitamin D D. R. Fraser Introduction Structure and Synthesis Vitamin D Formation in Skin Metabolism of Vitamin D Regulation of Vitamin D Metabolism Vitamin D Function Vitamin D Deficiency Chapter 3. Calcium-Regulating Hormones: Parathyroid Hormone and Calcitonin G. D. Aurbach Introduction Parathyroid Hormone Parathyroid Hormone and the Kidney Actions of Parathyroid Hormone on Bone Effects in Other Tissues Mechanism of Action of Parathyroid Hormone Calcitonin Physiology of Calcitonin

8 viii Contents Chapter 4. Calcium-Regulating Hormones: General P. H. Adams Introduction Adrenal Corticosteroids Thyroid Hormone Hormones of the Anterior Pituitary Gland Insulin Sex Steroids Chapter 5. Gastrointestinal Absorption of Calcium F. Bronner Introduction Calcium Absorption Active Calcium Transport Development Aspects Analysis of Whole Animal Absorption Studies Altering Calcium Absorption Therapeutic Considerations Chapter 6. Renal Excretion of Calcium M. Peacock Introduction Calcium Excretion Form and Measurement of Calcium in Urine Expression of Urinary Calcium Variation in Daily Calcium Excretion Renal Excretion of Calcium Regulation of Plasma Calcium by the Kidney Hypercalciuria Diseases Affecting Urinary Calcium Chapter 7. Calcified Tissues: Chemistry and Biochemistry A. L. Boskey Introduction: Composition ofthe Calcified Tissue Comparative Histogenesis of the Calcified Tissues... " 173 Calcification Mechanisms Remodelling Conclusions Chapter 8. Calcified Tissues: Cellular Dynamics F. Melsen and L. Mosekilde Introduction Primary Bone Formation Postnatal Development of Bone Internal Reorganization Metabolic States Mainly Characterized by Altered Bone Turnover Metabolic States Mainly Characterized by Disturbed Osteoid Mineralization Metabolic States Mainly Characterized by a Reduction in Bone Mass

9 Contents ix Chapter 9. Calcified Tissues: Structure-Function Relationships J. Dequeker Introduction Macroanatomy Age-Related Changes in Bone Mass Age-Related Fracture and Osteoporosis Bone Mass and Risk of Fractures Identification of Those at Risk for Fractures Treatment of Osteoporosis Chapter 10. Calcium in ExtraceUular Fluid: Homeostasis G. Schaafsma Introduction Plasma Calcium Homeostasis Effects of Non-nutritional Factors on Plasma Calcium Homeostasis Effects of Nutritional Factors on Plasma Calcium Homeostasis Chapter 11. Calcium as an Intracellular Regulator A. K. Campbell Intracellular Calcium and Cell Behaviour The Cell Biology of Intracellular Calcium The Evidence for Calcium as an Intracellular Regulator How to Measure Free Calcium in Living Cells The Source of Intracellular Calcium for Cell Activation How Calcium Acts The Pathology of Intracellular Calcium Pharmacology of Intracellular Ca Perspectives Chapter 12. Cellular Calcium: Muscle S. Ebashi Historical Survey of Calcium Research in Muscle Troponin Myosin-Linked Regulation Ca 2 + Regulation in Smooth Muscle Actomyosin in Non-muscle Tissue Excitation-Contraction Coupling Ca 2 + Control of Metabolic Processes in Muscle Calcium Antagonists Chapter 13. Cellular Calcium: Nervous System M. P. Blaustein Introduction Regulation of Intracellular Calcium in Nerve Cells Physiological Role of Intracellular Calcium in Nerve Cells Summary and Conclusions

10 x Contents Chapter 14. Cellular Calcium: Secretion of Hormones W. 1. Malaisse Introduction Evidence that Calcium Participates in Secretion Calcium Fluxes in Endocrine Cells Targets for Cytosolic Ca Concluding Remarks Chapter IS. Cellular Calcium: Action of Hormones R. H. Wasserman Introduction Control of Intracellular Calcium Modulation of Intracellular Free Ca 2 + Concentrations Calmodulin and Other Calcium-Binding Proteins The Phosphoinositide Cycle G Proteins Linking Receptors to Enzyme Activation Phospholipase C Activation and Phosphoinositide Second Messengers Hormone-Stimulated Ca 2 + Fluxes Diacylglycerol and Protein Kinase C Physiological and Biochemical Responses to Ca 2 +- Mobilizing Agonists in Selected Systems Epilogue Chapter 16. Cellular Calcium: Cell Growth and Differentiation T. Fujita Introduction Extracellular Calcium and Cellular Calcium Requirement Intracellular Calcium and Calcium Influx Calmodulin and Other Calcium-Binding Proteins Calcium and Cyclic Nucleotides Phospholipid and Protein Phosphorylation Calcium and Immune Function Oncogenes and Growth Factors Bone Cell Differentiation Calcium-Regulating Hormones Calcium Deficiency and Neoplasia Summary Chapter 17. Dietary Requirements for Calcium B. E. C. Nordin and D. H. Marshall Introduction Calcium Absorption Calcium Excretion Effects of Calcium Deficiency Calcium Requirements and Recommended Allowances Comparison with Current Intakes and Recommended Allowances Worldwide Subject Index

11 Contributors Prof. P. H. Adams Department of Medicine, The Royal Infirmary, Manchester M139WL, UK Prof. G. D. Aurbach Department of Health and Human Services, National Institutes of Health, Bethesda, Maryland 20205, USA Prof. M. P. Blaustein Department of Physiology, University of Maryland, School of Medicine, 655 West Baltimore Street, Baltimore, Maryland 21201, USA Dr. A. L. Boskey Ultrastructural Biochemistry Laboratory, The Hospital for Special Surgery, Cornell University Medical College, 535 East 70th Street, New York, New York 10021, USA Dr. F. Bronner School of Dental Medicine, University of Connecticut Health Center, Farmington, Connecticut 06032, USA Dr. A. K. Campbell Department of Medical Biochemistry, University of Wales College of Medicine, Heath Park, Cardiff CF4 4XN, UK Dr. J. Dequeker Arthritis and Metabolic Bone Disease Research Unit, Universitaire Ziekenhuizen, Weligerveld 1,3041 Pellenberg, Belgium Prof. S. Ebashi National Institute for Physiological Sciences, Myodaiji, Okazaki 444,Japan

12 xii Contributors Prof. D. R. Fraser Department of Animal Husbandry, University of Sydney, Sydney 2006, New South Wales, Australia Prof. T. Fujita Department of Medicine, Kobe University School of Medicine, 5-1 Kusunoki-cho 7-Chome, Chuo-ku, Kobe, Japan Prof. W. J. Malaisse Laboratory of Experimental Medicine, Boulevard de Waterloo 115, B-1000 Brussels, Belgium Dr. D. H. Marshall MRC Unit for Hearing Research, University of Nottingham, Nottingham, UK Dr. F. Melsen University Institute of Pathology, Aarhus Amtssygehus, DK-8000 Aarhus C, Denmark Prof. L. Mosekilde University Department of Medicine, Aarhus Amtssygehus, DK-800 Aarhus C, Denmark Dr. Y. Nakao Department of Medicine, Kobe University School of Medicine, 5-1 Kusunoki-cho 7-Chome, Chuo-ku, Kobe, Japan Prof. B. E. C. Nordin Institute of Medical and Veterinary Science, Box 14 Rundle Mall Post Office, Adelaide 5000, South Australia, Australia Dr. M. Peacock Department of Medicine, Emerson Hall-421, 545 Barnhill Drive, Indiana University Medical Center, Indianapolis, Indiana 46223, USA Dr. W. G. Robertson King Faisal Specialist Hospital, Research Centre, Biological and Medical Research, PO Box 3354, Riyadh, Kingdom of Saudi Arabia Dr. G. Schaafsma Department of Nutrition, CIVO-Institutes TNO, PO Box 360, 3700 AJ Zeist, The Netherlands Dr. R. H. Wasserman Department of Physiology, 717 Veterinary Research Tower, New York State College of Veterinary Medicine, Cornell University, Ithaca, New York, 14853, USA