Dynamics of Bone and Cartilage Metabolism Principles and
Clinical Applications - 2nd Edition
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Dynamics of Bone and Cartilage Metabolism
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Dynamics of Bone
and Cartilage Metabolism
Edited by
MARKUS J. SEIBEL
Department of Medicine
Division of Endocrinology and Metabolism
University of Heidelberg Medical School
Heidelberg, Germany
SIMON P. ROBINS
Skeletal Research Unit
Rowett Research Institute
Aberdeen, United Kingdom
JOHN P. BILEZIKIAN
Departments of Medicine and Pharmacology
Division of Endocrinology
College of Physicians and Surgeons
Columbia University
New York, New York
San Diego New York Boston London Sydney Tokyo Toronto
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Library of Congress Cataloging-in-Publication Data
Dynamics of bone and cartilage metabolism / Markus J. Seibel, Simon P. Robins, and John P. Bilezikian,
editors.— 2nd ed.
p. ; cm.
ISBN-13: 978-0-12-088562-6 (hardcover : alk. paper)
ISBN-10: 0-12-088562-X (hardcover : alk. paper)
1. Bones—Metabolism. 2. Cartilage—Metabolism. 3. Extracellular matrix.
4. Bones—Metabolism—Disorders. 5. Biochemical markers.
[DNLM: 1. Bone and Bones—metabolism. 2. Biological Markers. 3. Cartilage—metabolism.
4. Extracellular Matrix—metabolism. WE 200 D997 2006] I. Seibel, M. J. II. Robins, Simon P.
III. Bilezikian, John P.
QP88.2.D96 2006
612.7’5—dc22
2006004804
British Library Cataloguing-in-Publication Data
A catalogue record for this book is available from the British Library.
ISBN 13: 978-0-12-088562-6
ISBN 10: 0-12-088562-X
For information on all Academic Press publications
visit our Web site at www.books.elsevier.com
Printed in the United States of America
06 07 08 09 10 9 8 7 6 5 4 3 2 1
Contents
Contributors xiii IV. Concluding Remarks 50
Preface xix References 50
CHAPTER 3 Vitamin K Dependent
PART I Components of the Organic Proteins of Bone and Cartilage
CAREN M. GUNDBERG AND SATORU K. NISHIMOTO
Extracellular Matrix of Bone I. Abstract 55
and Cartilage II. Introduction 55
III. Osteocalcin 56
IV. Matrix Gla Protein 59
CHAPTER 1 Structure, Biosynthesis V. Gas6 63
and Gene Regulation of Collagens in VI. Vitamin K/Warfarin 64
Cartilage and Bone References 65
KLAUS VON DER MARK
I. Introduction 3 CHAPTER 4 Noncollagenous Proteins;
II. The Collagen Families 5 Glycoproteins and Related Proteins
III. Bone Collagens 7 DICK HEINEGÅRD, PILAR LORENZO, AND TORE SAXNE
IV. Cartilage Collagens 9 I. Introduction 71
V. Collagen Biosynthesis 14 II. Cartilage Extracellular Matrix 72
VI. Collagen Genes and Transcriptional III. Bone, Extracellular Matrix 77
Regulation 17 IV. Concluding Remarks 79
VII. Factors Regulating Collagen Biosynthesis 21 References 80
VIII. Conclusions 25
References 26 CHAPTER 5 Proteoglycans and
Glycosaminoglycans
CHAPTER 2 Fibrillogenesis and TIM HARDINGHAM
Maturation of Collagens I. Introduction 85
SIMON P. ROBINS
II. Glycosaminoglycans 86
I. Introduction 41 III. Proteoglycans in Cartilage 87
II. Fibrillogenesis 42 IV. Aggrecan 88
III. Cross-linking 44
v
vi Contents
V. Leucine-rich Proteoglycans in Cartilage CHAPTER 10 Acid Phosphatases
and Bone 93 HELENA KAIJA, LILA O.T. PATRIKAINEN,
VI. Perlecan in Cartilage 96 SARI L. ALATALO, H. KALERVO VÄÄNÄNEN,
References 96 AND PIRKKO T. VIHKO
I. Acid Phosphatases 165
CHAPTER 6 Growth Factors II. Tartrate-Resistant Acid Phosphatase
PHILIPPA HULLEY, GRAHAM RUSSELL, (TRACP) 166
AND PETER CROUCHER III. Prostatic Acid Phosphatase (PAP) 173
I. Introduction 99 References 176
II. Insulin-like Growth Factors 99
III. The Transforming Growth Factor-Beta/Bone CHAPTER 11 Matrix Proteinases
Morphogenetic Protein Superfamily 100 IAN M. CLARK AND GILLIAN MURPHY
IV. Fibroblast Growth Factors 103 I. Introduction 181
V. Wnts 104 II. Aspartic Proteinases 181
VI. Additional Growth Factors 105 III. Cysteine Proteinases 182
VII. Summary 106 IV. Serine Proteinases 184
References 107 V. Metalloproteinases 186
References 192
CHAPTER 7 Prostaglandins and
Proinflammatory Cytokines
LAWRENCE G. RAISZ AND JOSEPH A. LORENZO PART II Structure and Metabolism
I. Introduction 115
II. Prostaglandins 115 of the Extracellular Matrix of
III. The Role that Cytokines have in Osteoclast Bone and Cartilage
Formation and Function 117
IV. The Role that Proinflammatory Cytokines have
in Bone and Cartilage Metabolism 118 CHAPTER 12 Mineralization, Structure
References 121 and Function of Bone
ADELE L. BOSKEY
CHAPTER 8 Integrins and Other I. Abstract 201
Adhesion Molecules II. The Structure and Function of Bone 201
M. H. HELFRICH AND M. A. HORTON III. Bone Mineralization 204
I. Abstract 129 IV. Bone Modeling and Remodeling 209
II. Introduction 129 References 209
III. Molecular Structure of Adhesion
Molecules 130 CHAPTER 13 Bone Structure
IV. Adhesion Molecules in Cells of the Osteoblast and Strength
Lineage 134 EGO SEEMAN
V. Adhesion Molecules in Osteoclasts 139 I. Introduction 213
VI. Adhesion Molecules in Chondrocytes 142 II. Gravity and the Need for Stiffness, Flexibility,
VII. Conclusion 144 Lightness and Speed 213
References 144 III. The Material Composition and Structural Design
of Bone 214
CHAPTER 9 Alkaline Phosphatases IV. Bone Modeling and Remodeling – The
JOSÉ LUIS MILLÁN Mechanism of Bone’s Construction during
I. Introduction 153 Growth and Decay with Advancing Age 216
II. Structure And Regulation of the TNAP V. Strength Maintenance 217
Gene 153 VI. Conclusion 219
III. Protein Structure 155 References 219
IV. Function of TNAP 157
V. Clinical Use 160
References 161
Contents vii
CHAPTER 14 The Cells of Bone IX. Activation of the Cyclic Adenosine
JANE B. LIAN AND GARY S. STEIN Monophosphate Second-Messenger System by
I. Abstract 221 Parathyroid Hormone 281
II. Introduction 221 X. Identification of a Second PTH Receptor 282
III. Developmental Signals for Cartilage and Bone XI. Physiological Actions of PTH 282
Tissue Formation 222 XII. Cell-to-Cell Communication: Osteoblasts and
IV. Osteogenic Lineage Cells 227 Osteoclasts 284
V. The Osteoclast: A Functionally Unique Cell XIII. Preferential Actions of PTH at Selected Skeletal
for Physiologically Regulated Resorption Sites 285
of Bone Mineral 236 References 287
VI. Perspectives 242
References 242 CHAPTER 17 Interaction of Parathyroid
Hormone-related Peptide with the
CHAPTER 15 Signaling in Bone Skeleton
T. JOHN MARTIN AND NATALIE A. SIMS DAVID GOLTZMAN
I. Abstract 259 I. Abstract 293
II. Introduction 259 II. Introduction 293
III. The Control of Osteoclasts 260 III. Molecular Biology and Mechanism
IV. Signaling in the Control of Osteoclast of Action 294
Activity 261 IV. The Skeletal Actions of PTHrP 298
V. Signals from the Osteoblast Lineage that V. Summary 301
Control Osteoclast Formation 261 References 302
VI. Hormone and Cytokine Influences on
the Contact-dependent Regulation CHAPTER 18 The Vitamin D Hormone
of Osteoclasts 261 and its Nuclear Receptor: Mechanisms
VII. Discovery of the Physiological Signaling
Mechanisms in Osteoclast Control 262
Involved in Bone Biology
GEERT CARMELIET, ANNEMIEKE VERSTUYF,
VIII. Rank Signaling 263
CHRISTA MAES, GUY EELEN, AND ROGER BOUILLON
IX. Coupling of Bone Formation to
Resorption – Release of Growth Factors I. Introduction 307
from bone Matrix 264 II. Metabolism of Vitamin D 308
X. Coupling of Bone Formation to III. Nuclear Vitamin D Receptor 309
Resorption – Autocrine/Paracrine Regulation IV. Vitamin D and Bone Cells 312
by Differentiating Osteoblasts 266 V. Pathology and Therapy Related to Vitamin D
XI. Coupling of Bone Formation to Resorption – Are Availability, Metabolism, and Function 314
Osteoclasts a Source of Coupling Activity? 266 VI. Conclusions 319
References 267 References 319
CHAPTER 16 Parathyroid Hormone: CHAPTER 19 Sex Steroid Effects
Structure, Function and Dynamic Actions on Bone Metabolism
LORRAINE A. FITZPATRICK AND JOHN P. BILEZIKIAN DAVID G. MONROE, THOMAS C. SPELSBERG,
AND S. KHOSLA
I. Introduction 273
II. Structure of the PTH Gene 273 I. Abstract 327
III. Chromosome Location 274 II. Introduction 327
IV. Control of Gene Expression 275 III. Molecular Structures, Synthesis, Mechanism of
V. Biosynthesis of Parathyroid Hormone 278 Action of Major Sex Steroids, and Transcriptional
VI. Metabolism of Parathyroid Hormone 278 Coregulator Function 328
VII. Receptor Interactions of Parathyroid Hormone IV. Effects of Sex Steroids on Bone Cells and Bone
and Parathyroid Hormone-Related Protein 279 Turnover 331
VIII. Structure of the PTH/PTHrP (PTH1R) V. Effects of Estrogens and Androgens on Bone
Receptor 279 Metabolism in Men Versus Women 335
viii Contents
VI. Effects of Sex Steroids on Extraskeletal Calcium CHAPTER 23 Products of Bone Collagen
Homeostasis 336 Metabolism
VII. Summary 337 JUHA RISTELI AND LEILA RISTELI
References 338
I. Introduction 391
II. Products of Bone Collagen Synthesis,
CHAPTER 20 Physiology of Calcium and the Procollagen Propeptides 392
Phosphate Homeostases III. Degradation Products of Type I Collagen 397
RENÉ RIZZOLI AND JEAN-PHILIPPE BONJOUR IV. Closing Remarks 402
I. Abstract 345 References 403
II. Introduction 345
III. Body Distribution of Calcium 346 CHAPTER 24 Supramolecular Structure
IV. Determinants of Extracellular Calcium of Cartilage Matrix
Concentration 346 PETER BRUCKNER
V. Relative Importance of the Various Calcium
I. Summary 407
Fluxes in Controlling Extracellular Calcium
II. Introduction 407
Homeostasis 349
III. Light and Electron Micrography 408
VI. Homeostatic Responses to Hypocalcemia 349
IV. Biochemistry of Cartilage 409
VII. Calcium and Bone Growth 352
V. Studies of Fibril Structures by X-ray
VIII. Body Distribution of Phosphorus 353
Diffraction 411
IX. Determinants of Extracellular Phosphate
VI. Structure of Fibril Fragments Obtained by
Concentration 353
Mechanical Disruption of Tissue 411
X. Homeostatic Responses to Changes in Phosphate
VII. Studies of Collagen Cross-linking in Cartilage
Supply or Demand 356
Fibrils 412
XI. Conclusions 357
VIII. Reconstitution of Aggregates from Soluble
References 357
Collagens and other Macromolecules 412
IX. Studies of Transgenic Mice and of Human
CHAPTER 21 The Central Control Genetic Matrix Diseases 414
of Bone Remodeling X. Correlating Structure With the Biomechanical
PAUL A. BALDOCK, SUSAN J. ALLISON, HERBERT HERZOG, Role of Articular Cartilage 415
AND EDITH M. GARDINER XI. Models of Cartilage Fibril Structure 416
I. Introduction 361 XII. Future Perspectives 417
II. Actions of Leptin 362 References 418
III. Sympathetic Nervous System 365
IV. Neuropeptide Y and the Y Receptors 369 CHAPTER 25 Products of Cartilage
V. Interaction between Leptin and Y2-Regulated Metabolism
Bone Antiosteogenic Pathways 371 DANIEL-HENRI MANICOURT, JEAN-PIERRE DEVOGELAER,
VI. Concluding Remarks 373 AND EUGENE J.-M. A. THONAR
References 373
I. Introduction 421
II. The Chondrocyte and its Extracellular
CHAPTER 22 New Concepts Matrix 422
in Bone Remodeling III. Products of Collagen Metabolism 424
DAVID W. DEMPSTER AND HUA ZHOU IV. Products of Aggrecan Metabolism 429
I. Introduction 377 V. Products of the Metabolism of other
II. An Overview of the Remodeling Cycle 377 Proteoglycans 434
III. Functions of Bone Remodeling 380 VI. Products of the Metabolism of Link Protein
IV. The Role of Apoptosis in Regulating Bone and Hyaluronan 435
Balance 381 VII. Other Products of Chondrocyte
V. Possible Mechanisms whereby a Reduction Metabolism 437
in Activation Frequency may Protect against VIII. Concluding Statement 438
Fracture 383 References 439
References 386
Contents ix
CHAPTER 26 Fluid Dynamics of the Joint CHAPTER 30 Measurement of
Space and Trafficking of Matrix Products Parathyroid Hormone
PETER A. SIMKIN HARALD JÜPPNER AND GHADA EL-HAJJ FULEIHAN
I. Introduction 451 I. Abstract 507
II. Interpretation of Marker Data and Strategies for II. Background 507
Dealing with Them 451 III. Different Immunometric Assays for
III. Conclusion 456 the Detection of PTH 509
References 456 IV. Primary Hyperparathyroidism 510
V. Hyperparathyroidism in Renal
CHAPTER 27 Transgenic Models of Osteodystrophy 510
Bone Disease VI. Pseudohypoparathyroidism (PHP) 510
BARBARA E. KREAM AND JOHN R. HARRISON VII. Conclusion 511
I. Introduction 457 References 511
II. Generation of Mouse Models 457
III. Transgenic Models in Bone Biology 460 CHAPTER 31 New Horizons for
IV. Perspectives and Future Directions 465 Assessment of Vitamin D Status in Man
References 465 GARY L. LENSMEYER, NEIL BINKLEY,
AND MARC K. DREZNER
I. Measurement of Vitamin D3 (Cholecalciferol)
PART III Markers of Bone and and Vitamin D2 (Ergocalciferol) 513
II. Measurement of 25-Hydroxyvitamin D 513
Cartilage Metabolism III. Measurement of 1,25-Dihydroxyvitamin D 524
References 525
CHAPTER 28 The Role of Genetic
CHAPTER 32 Measurement of
Variation in Osteoporosis
ANDRÉ G. UITTERLINDEN, JOYCE B.J. VAN MEURS,
Biochemical Markers of Bone
FERNANDO RIVADENEIRA, JOHANNES P.T.M.VAN LEEUWEN, Formation
AND HUIBERT A.P. POLS KIM E. NAYLOR AND RICHARD EASTELL
I. Abstract 471 I. Abstract 529
II. Osteoporosis has Genetic Influences 471 II. Introduction 529
III. Genome-wide Approaches to Find the III. Propeptides of Type I Procollagen 530
Genes 473 IV. Total Alkaline Phosphatase 533
IV. Association Analysis of Candidate Gene V. Bone Alkaline Phosphatase 533
Polymorphisms 476 VI. Osteocalcin 535
V. Haplotypes 478 VII. Discussion 537
VI. Meta-analyses 480 References 537
VII. Osteoporosis Candidate Genes: Collagen Type Iα1
and the Vitamin D Receptor 482 CHAPTER 33 Measurement of
VIII. Summary 483 Biochemical Markers of Bone
References 484 Resorption
MARIUS E. KRAENZLIN AND MARKUS J. SEIBEL
CHAPTER 29 Measurement of Calcium, I. Introduction 541
Phosphate and Magnesium II. Collagen Related Markers 543
HEINRICH SCHMIDT-GAYK III. Non-Collagenous Proteins of the Bone
I. Measurement of Calcium 487 Matrix 553
II. Measurement of Phosphate 495 IV. Osteclast Enzymes 555
III. Measurement of Magnesium 499 References 557
References 502
x Contents
CHAPTER 34 Variability in the CHAPTER 38 Monitoring
Measurement of Biochemical Markers Anabolic Treatment
of Bone Turnover JOHN P. BILEZIKIAN AND MISHAELA R. RUBIN
TUAN V. NGUYEN, CHRISTIAN MEIER, I. Introduction 629
AND MARKUS J. SEIBEL II. Cellular and Regulatory Mechanisms of the
I. Introduction 565 Anabolic Actions of Parathyroid Hormone 630
II. Sources of Pre-Analytical Variability in III. Pharmacokinetics of Teriparatide in Human
the Measurement of Biochemical Markers Subjects 631
of Bone Turnover 566 IV. Actions of Parathyroid Hormone to Improve
III. Statistical Consideration of Variability 571 Bone Quality 632
IV. Summary 577 V. Conclusions 642
References 578 References 642
CHAPTER 35 Validation of Biochemical CHAPTER 39 Monitoring of
Markers of Bone Turnover Antiresorptive Therapy
KIM BRIXEN AND ERIK FINK ERIKSEN CHRISTIAN MEIER, TUAN V. NGUYEN,
AND MARKUS J. SEIBEL
I. Introduction 583
II. Validation of Biochemical Markers by Calcium I. Introduction 649
Kinetics 584 II. Effects of Pretreatment Bone Turnover and
III. Validation of Biochemical Markers by Bone Mineral Density on Therapeutic Outcomes 650
Histomorphometry 588 III. The Role of Markers of Bone Turnover in
IV. Conclusions 592 Monitoring Antiresorptive Osteoporosis
References 592 Therapy 651
IV. Interpretation of Changes in Bone Turnover
CHAPTER 36 Genetic Markers Markers 661
of Joint Disease References 665
MICHEL NEIDHART, RENATE E. GAY, AND STEFFEN GAY
CHAPTER 40 Age-related Osteoporosis
I. Introduction 595
II. Ankylosing Spondylitis 595 and Skeletal Markers of Bone
III. Reactive Arthritis 597 Turnover
IV. Rheumatoid Arthritis 598 CLIFFORD J. ROSEN
V. Juvenile Rheumatoid Arthritis 605 I. Introduction 671
VI. Conclusion 606 II. Epidemiology and Pathogenesis of
References 606 Age-Related Osteoporosis-Relationship to
Bone Turnover 672
CHAPTER 37 Laboratory III. Markers of Bone Turnover and Age-Related
Assessment of Postmenopausal Osteoporosis – Clinical Implications 679
Osteoporosis IV. Summary 683
PATRICK GARNERO AND PIERRE D. DELMAS References 683
I. Introduction 611
CHAPTER 41 Steroid-Induced
II. Postmenopausal Bone Loss 612
III. Management of Postmenopausal Osteoporosis
Osteoporosis 616 IAN R. REID
IV. Conclusion 624 I. Summary 689
References 624 II. Introduction 689
III. Epidemiology 689
IV. Pathogenesis 690
V. Effects of Glucocorticoids on Histological Indices
of Bone Turnover 691
Contents xi
VI. Effect of Glucocorticoids on Markers of IV. Summary 762
Bone Turnover 692 References 762
VII. Evaluation of Steroid-Treated Patients 694
VIII. Treatment and Follow-up 694 CHAPTER 46 Primary
References 695 Hyperparathyroidism
SHONNI J. SILVERBERG AND JOHN P. BILEZIKIAN
CHAPTER 42 Transplantation I. Introduction 767
Osteoporosis: Biochemical Correlates II. Etiology 767
of Pathogenesis and Treatment III. Clinical Presentation 768
CAROLINA A. MOREIRA KULAK AND ELIZABETH SHANE IV. Bone Markers in Primary
I. Introduction 701 Hyperparathyroidism 770
II. Kidney Transplantation 702 V. Cytokines in Primary Hyperparathyroidism 772
III. Cardiac Transplantation 705 VI. Treatment of Primary Hyperparathyroidism 772
IV. Liver Transplantation 706 VII. Summary 775
V. Lung Transplantation 706 References 775
VI. Bone Marrow Transplantation 707
VII. Mechanisms of Bone Loss After CHAPTER 47 Paget’s Disease of Bone
Transplantation 707 ANDREAS GRAUER, ETHEL SIRIS, AND STUART RALSTON
VIII. Prevention and Management of Transplantation I. Introduction 779
Osteoporosis 709 II. Etiology and Pathogenesis 779
IX. Conclusions 711 III. Treatment 784
References 711 References 788
CHAPTER 43 Secondary Osteoporosis CHAPTER 48 Metastatic Bone Disease
JEAN E. MULDER, CAROLINA A. MOREIRA KULAK, JEAN-JACQUES BODY
AND ELIZABETH SHANE I. Abstract 793
I. Introduction 717 II. Introduction 794
II. Hyperthyroidism and Osteoporosis 717 III. Use of Markers of Bone Turnover for the
III. Osteoporosis Secondary to Hypogonadism 724 Diagnosis of Bone Metastases 796
IV. Anticonvulsant Drugs and Osteoporosis 730 IV. Use of Markers of Bone Turnover for the
References 732 Monitoring of Tumor Bone Disease 802
V. Prediction of the Development of Bone
CHAPTER 44 Osteomalacia and Rickets Metastases 806
MARC K. DREZNER References 806
I. Definition 739
II. Etiology 741 CHAPTER 49 Rare Bone Diseases
III. Incidence and Epidemiology 742 MICHAEL P. WHYTE
IV. Calciopenic Rickets and Osteomalacia 743 I. Introduction 811
V. Phosphopenic Rickets and Osteomalacia 747 II. Osteopenia 812
VI. Normal Mineral Rickets and Osteomalacia 750 III. Osteosclerosis and Hyperostosis 817
References 751 IV. Ectopic Calcification 823
V. Other Disorders 825
CHAPTER 45 Assessment of Bone References 826
and Joint Diseases:
Renal Osteodystrophy CHAPTER 50 Osteogenesis Imperfecta
FRANCIS H. GLORIEUX AND FRANK RAUCH
ESTHER A. GONZÁLEZ, ZIYAD AL ALY,
AND KEVIN J. MARTIN I. Introduction 831
II. Classification 832
I. Introduction 755
III. Diagnosis 833
II. Biochemical Assessment of Renal
IV. Differential Diagnosis 834
Osteodystrophy 757
V. Pathogenesis 834
III. Skeletal Imaging in Renal Osteodystrophy 761
VI. Bisphosphonate Therapy in OI 835
xii Contents
VII. Medical Therapies Other than CHAPTER 52 Osteoarthritis and
Bisphosphonates 838 Degenerative Spine Pathologies
VIII. Potential Future Therapies 839 KRISTINA ÅKESSON
IX. Conclusions 839
I. Introduction 871
References 839
II. Characteristics of OA 872
III. Etiology 872
CHAPTER 51 Rheumatoid Arthritis and IV. Treatment Options 873
other Inflammatory Joint Pathologies V. Current Diagnostic Procedures 873
STEVEN R. GOLDRING AND MARY B. GOLDRING VI. Biochemical Aspects of Osteoarthritis 873
I. Abstract 843 VII. Markers of Bone Turnover 874
II. Introduction 843 VIII. Markers of Cartilage Metabolism 878
III. Effects of Joint Inflammation on Skeletal IX. Spine Degeneration and Markers 882
Remodeling 844 X. Summary 883
IV. Effects of Joint Inflammation on Cartilage XI. Conclusions 884
Remodeling 852 References 884
V. Conclusion 858
References 858
Contributors
Kristina Åkesson John P. Bilezikian
Associate Professor, Department of Orthopedics, Department of Medicine, College of Physicians
Malmö University Hospital, 205 02 Malmö, and Surgeons, Columbia University, New York,
Sweden NY, USA
Departments of Medicine and Pharmacology,
Sari L. Alatalo College of Physicians and Surgeons, Columbia
Finnish Red Cross Blood Service, Helsinki, Finland University, New York, USA
Susan J. Allison Neil Binkley
Postgraduate Scholar, Bone Research Program, Garvan University of Wisconsin, Madison, Wisconsin, USA
Institute of Medical Research, 384 Victoria Street,
Sydney NSW 2010, Australia Jean-Jacques Body
Dept of Internal Medicine and Endocrinology/
Ziyad Al Aly Bone Diseases Clinic, Institut J. Bordet, Univ. Libre de
Division of Nephrology, Saint Louis University School Bruxelles, Brussels, Belgium
of Medicine, St. Louis, Missouri, USA
Jean-Philippe Bonjour
Paul A. Baldock Division of Bone Diseases, WHO Collaborating Center
Senior Research Officer, Bone Research Program, for Osteoporosis Prevention, Department of
Garvan Institute of Medical Research,384 Victoria Rehabilitation and Geriatrics, University Hospitals,
Street, Sydney NSW 2010, Australia CH - 1211 Geneva 14 (Switzerland)
John P. Bilezikian Adele L. Boskey
Departments of Medicine and Pharmacology, Starr Chair in Mineralized Tissue Research, Hospital for
College of Physicians and Surgeons, Columbia Special Surgery, New York, NY 10021 and Weill
University, New York, NY Medical College and Graduate School of Medical
Sciences of Cornell University, New York, NY 10021
xiii
xiv Contributors
Roger Bouillon Erik Fink Eriksen
Laboratorium for Experimental Medicine and Novartis Pharma, Basel, Switzerland
Endocrinology, K. U. Leuven, Gasthuisberg,
Herestraat 49, 3000 Leuven, Belgium Lorraine A. Fitzpatrick
Global Development, Amgen, Thousand Oaks, CA
Kim Brixen
Department of Endocrinology, Odense University Ghada El-Hajj Fuleihan
Hospital, DK-5000 Odense C, Denmark Calcium Metabolism and Osteoporosis Program,
American University of Beirut-Medical Center,
Peter Bruckner Beirut, Lebanon
Department of Physiological Chemistry and
Pathobiochemistry, University of Münster, Münster, Edith M. Gardiner
Germany Associate Professor, School of Medicine,
The University of Queensland, Head, Skeletal Biology
Geert Carmeliet Unit, Centre for Diabetes & Endocrine Research,
Laboratorium for Experimental Medicine and Ground Floor, C Wing, Bldg 1, Princess Alexandra
Endocrinology, K. U. Leuven, Gasthuisberg, Hospital, Ipswich Road, Brisbane QLD 4102,
Herestraat 49, 3000 Leuven, Belgium Australia
Ian M Clark Patrick Garnero
School of Biological Sciences, University of East Research Scientist, INSERM research unit 403 and
Anglia, Norwich NR4 7TJ, UK Vice-President Synarc Molecular Marker Division,
Lyon, France
Peter Croucher
Academic Unit of Bone Biology, University of Sheffield Renate E. Gay
Medical School, Sheffield S10 2RX, United Kingdom Research Scientist, INSERM research unit 403 and
Vice-President Synarc Molecular Marker Division,
Pierre D. Delmas Lyon, France
Professor of Medicine, Université Claude Bernard,
Lyon, France and Director INSERM Research Unit 403, Steffen Gay
Lyon, France Research Scientist, INSERM research unit 403 and
Vice-President Synarc Molecular Marker Division,
David W. Dempster Lyon, France
Regional Bone Center, Helen Hayes Hospital,
West Haverstraw, New York, USA Francis H. Glorieux
Genetics Unit, Shriners Hospital for Children, 1529
Jean-Pierre Devogelaer Cedar Avenue, Montréal, Québec, Canada H3G 1A6
Department of Rheumatology, Saint Luc University
Hospital, Université Catholique de Louvain, 1200 Mary B. Goldring
Brussels, Belgium Medical Center, Harvard Medical School, Boston, MA;
New England Baptist Bone and Joint Institute, Boston,
Marc K. Drezner MA 02215
Professor of Medicine, University of Wisconsin,
Madison, Wisconsin, USA Steven R. Goldring
Department of Medicine, Rheumatology Division,
Richard Eastell Beth Israel Deaconess
From the Academic Unit of Bone Metabolism,
University of Sheffield, Sheffield, UK David Goltzman
Calcium Research Laboratory, Department of Medicine,
Guy Eelen Royal Victoria Hospital, McGill University, Montreal,
Laboratorium for Experimental Medicine and H3A 1A1, Canada
Endocrinology, K. U. Leuven, Gasthuisberg, Herestraat
49, 3000 Leuven, Belgium
