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HANDBOOK OFPLATELET PHYSIOLOGY ANDPHARMACOLOGYEdited byGundu H. R. RaoUniversity of MinnesotaKLUWER ACADEMIC PUBLISHERSBoston / Dordrecht / LondonDistributors for North, Central and South America:Kluwer Academic Publishers101 Philip DriveAssinippi ParkNorwell, Massachusetts 02061 USATelephone (781) 871-6600Fax (781) 871-6528E-Mail <kluwer@wkap.com>Distributors for all other countries:Kluwer Academic Publishers GroupDistribution CentrePost Office Box 3223300 AH Dordrecht, THE NETHERLANDSTelephone 3178 6392 392Fax 3178 6546 474E-Mail <services@wkap.nl>Electronic Services <http://www.wkap.nl>Library of Congress Cataloging-in-Publication DataHandbook of platelet physiology and pharmacology / edited by GunduH.R. Rao.p. cm.Includes bibliographical references and index.ISBN 0-7923-8538-1 (alk. paper)1. Blood platelets Handbooks, manuals, etc. I. Rao, Gundu H.R 1938- .QP97.H36 1999612.ri7-dc21 99-27962CIPCopyright © 1999 by Kluwer Academic PublishersAll rights reserved. No part of this publication may be reproduced, stored in aretrieval system or transmitted in any form or by any means, mechanical, photo-copying, recording, or otherwise, without the prior written permission of thepublisher, Kluwer Academic Publishers, 101 Philip Drive, Assinippi Park, Norwell,Massachusetts 02061Printed on acid-free paper.Printed in the United States of AmericaList of Contributors1 . Kailash C. Agarwal, Ph. D.Department ofMolecular Pharmacology,Brown UniversityProvidence, RI02912,USA2. Colin N. Baigent, Ph. D.ATT CollaborationCTSU Harkness BuildingRadcliffe InfirmaryWoodstock RoadOxford 0X2 6HEUnited Kingdom3. Rodger L. Bick, M. D., Ph. D.Departments ofPathology & PharmacologyLoyola University Med. Center21260 South First Ave.Maywood, IL60153,USA4. David C. Calverley, M. D.USC School of Medicine1441EastlakeAveNOR MS 34Los Angeles, CA90033, USA5. Thomas Chandy, Ph. D.Chemical Engineering andMaterial SciencesUniversity of MinnesotaMinneapolis, MN55455, USA6. Kenneth J. Clemetson, Ph. D.Theodor Kocher InstitutDer Universitat BernFreiestrasse 1, Ch-3012 BerneSwitzerland7. Robert W. Colman, M. D.Thrombosis Research CenterTemple Univ. Sch. of Medicine3400 N. Broad StreetPhiladelphia, PA19140,USA8. Maribel Diaz-Ricart, Ph. D.Servicio de HemoterapiaHospital Clinico ProvincialVillarroel 170, Barcelona08036, Spain9. Gines Escolar, M. D. Ph. D.Servicio de HemoterapiaHospital Clinico ProvincialVillarroel 170, Barcelona08036, Spain.10. Daniel Fareed, B.Sc.Departments ofPathology & PharmacologyLoyola University Med. Center2 1260 South First Ave.Maywood, IL60153,USA1 1 . Jawed Fareed, Ph. D.Departments ofPathology & PharmacologyLoyola University Med. Center2 1 260 South First Ave.Maywood, IL60153,USA12. Deborah French, M. D.Department of MedicineMount Sinai Hospital &Medical SchoolOne Gustave L. Levy PlaceNew York, NY10029-6574,USA1 3. Mony M. Frojmovic, Ph. D.Mclntyre MedicalScience BuildingMcGiIl University3655 Drummond StreetMontreal, QBCanada, H3GIY614. Nicholas J.Greco, Ph. D.Platelet Biology LaboratoryAmerican Red cross1 5601 Crabbs Branch WayRockville, MD20855,USA15. HolmHolmsen,Ph.D.Department of Biochemistry andMolecular BiologyUniversity of BergenAstradveien 19, BergenN5009, Norway1 6. Debra Hoppensteadt, Ph. D.Departments of Pathology &PharmacologyLoyola University Med. Center2 1260 South First Ave.Maywood, IL60153,USA17. Huzoor-Akbar, Ph. D.Molecular and Cellular BiologyDepartment of BiologicalSciences, Irvine HallAthens, OH45701,USA1 8. G. A. Jamieson, Ph. D, D. Sc.Platelet Biology LaboratoryAmerican Red Cross15601 Crabbs Branch WayRockville, MD20855,USA1 9. Gerhard J. Johnson, M. D.Veterans Affairs Medical CenterOne Veterans WayMinneapolis, MN55417,USA20. BeateKehrel,Ph.D.Experimental and ClinicalHaemostaseologyDepartment of Anaesthesiologyand Intensive Care MedicineUniversity of MuensterD-48149 Muenster, Germany2 1 . Bruce R.Lester, Ph. D,.Knowledge Frontiers3989 Central Ave, N. E., # 625Minneapolis, MN55421,USA22. Mahadev Murthy, Ph. D.Division Endocrinolgy,Metabolism & NutritionDepartment of MedicineHennepin County Medical Center914 South Eighth Street, D-3Minneapolis, MN55404. USA23. Ellinor I. Peerschke, Ph. D.Cornell Medical CenterNew York University525 E 68th Street, Rm F51 1 JNew York, NY10021,USA24. Anna S. RadomskiDivision of R and DLacer, S.A.08025 BarcelonaSpain25. Marek W. Radomski, M.D,D.Sc.Division of R and DLacer, S.A., 08025 BarcelonaSpain26. Gundu H. R. Rao, Ph. D.Departments of Lab. Med. &Pathol. and Biomed. EngineeringP.B. 609 UMHCAcademic Health CenterUniversity of MinnesotaMinneapolis, MN55455,USA27. A. Koneti Rao, M. D.Department of MedicineTemple University School ofMedicine3400 N. Broad St Rm 300-OMSPhiladelphia, PA19140,USA28. Gerald J. Roth, M. D.Division of HematologyV.A. Medical Center1660 South Columbian WaySeattle, WA98108,USA29. Anita Ryningen, Ph. D.Department of Biochemistryand Molecular BiologyUniversity of BergenAstradveien 19Bergen N-5009Norway30. Shivendra D. Shukla, Ph. D.University of Missouri-Columbia517B Medical Science BuildingOne Hospital DriveColumbia, MO65212,USA3 1 . Cathie Sudlow, Ph. D.ATT CollaborationCTSU Harkness BuildingRadcliffe InfirmaryWoodstock RoadOxford OX2 6HEUnited Kingdom32. Narendra N. Tandon, Ph. D.Thrombosis & Vascular BiologyOtsuka America Pharmaceutical9900 Medical Center DriveRockville, Maryland20850, USA33. Jeanine M. Walenga, Ph. D.Departments of Pathology &PharmacologyLoyola UniversityMedical Center2 1260 South First Ave.Maywood, IL60153,USA34. Douglas J. Weiss, D. V.M., Ph. D.Department of Pathobiologyand Veterinary SciencesUniversity of MinnesotaSt. Paul, MN55108,USA35. Helmut Wolf, M. D, Ph. D.Departments of Pathology &PharmacologyLoyola UniversityMedical Center2 1260 South First Ave.Maywood, IL60153,USAPREFACEDespite my many years of research and teaching in platelet physiology andpharmacology at the University of Minnesota, I am often confronted with conflictingopinions as to the relevance of nonnucleated platelets in human health and disease. Itis fascinating to think that how cells with no apparent nucleus, have such a toweringimpact on concepts, dealing with often overlapping physiological (i.e. hemostasis,wound healing, etc.) and pathophysiological (i.e. thrombosis, stroke, atherosclerosis,wound healing, diabetes, inflammation and cancer) components. Although the idea ofcompiling new frontiers of platelet research in the form of a book was quite simple atthe beginning, the project turned out to be a major undertaking from my part. At theend, I am elated that the contributors to this book were gracious enough to write chaptersin their area of research expertise despite their pressing and highly valuable time. Forme, it has been an humbling experience as the chapters that I have compiled, are writtenby people with incredible recognition for their relentless contributions over the years tostrengthen the understanding of platelet physiology and pharmacology. In my opinion,this has added an immense value to the book. I am proud to have been involved in thisundertaking despite several unexpected problems and delays during this project. I amconfident that this book would be highly useful to the community of scientists, includinggraduate students, researchers, academicians, physicians and other health careprofessionals, and pharmaceutical industry scientists.Circulating platelets which lack nucleus neither adhere to the vessel wall nor aggregateunless they encounter a zone of injury. Upon encountering such a zone of injury, theybecome almost instantly activated, which leads to their adhesion and aggregation, bothreactions are of fundamental importance to hemostasis and thrombosis. Because of thisreason, platelet research has clearly led the way in the continuing development of newstrategies and drugs that can help prevent and treat arterial thrombosis, stroke andatherosclerosis. Unquestionably, platelet research has also impacted concepts dealingwith many other diseases. Nevertheless, considerable progress has been made in thedevelopment of new antiplatelet agents in recent years. These newer agents are aimedat interrupting specific sites and pathways of platelet activation. Inhibitors of specificplatelet agonist-receptor interactions include antithrombins, thromboxane A2 receptorantagonists, and adenosine diphosphate receptor blockers (i.e. ticlopidine, clopidogrel).In addition, inhibitors of arachidonic acid metabolism and thromboxane A2 includeaspirin, newer COX-2 inhibitors, other NSAIDs, thromboxane A2 synthase inhibitorsand o>-3 fatty acids. Moreover, long awaiting drugs that block ligand binding to theplatelet glycoprotein Ilb/IIIa complex (i.e. tirofiban) have now entered the market.In this book, the chapters are organized into six major sections, including Introduction,Receptor Biology, Platelet Biochemistry, Experimental Physiology, Platelet Pathologyand Platelet Pharmacology. Authoritative chapters in each section have provided acollective strength to our initial philosophy of accomplishing a comprehensive reviewof current concepts in each discipline. Although every attempt has been made to providean interdisciplinary discussion on the subject of platelets in this book, there may still besome gaps and lapses for which readers are urged to consult other articles and reviews.I have deliberately avoided going into any specific comments on reviews in order to letthe imagination of the readers flow freely. I believe that the readers are intelligentenough to judge and form their own critical opinion.I must humbly express my deep gratitude to thirty five scientists in the field for theirinvaluable contributions. I now honestly believe that this publication would not havebeen possible without their meritorious contributions.I am deeply indebted to my dear friend and close research collaborator, MahadevMurthy, Ph. D., Director of Research, Division of Endocrinology, Metabolism andNutrition, Department of Medicine, Hennepin County Medical Center, Minneapolis,MN, USA, for his commitment and contribution to this project. He has spent countlesshours during this project in reviewing and preparing camera ready manuscripts for finalsubmission to the Kluwer Academic Publisher. In addition, he has written two excellentchapters for the book. I must confess that this publication would not have beencompleted without his generous and truly dedicated efforts.I would like to take this opportunity to thank Charles W. Schmieg, Jr., AcquisitionsEditor, Kluwer Academic Publishers, 101 Philip Drive, Assinippi Park, Norwell, MA,02061, USA, for facilitating the publication of this book. I am specially thankful for hiscooperation and patience even though this project was delayed by about four months.Finally, I would not be in this field today without my mentor, James G. White, M. D.,Regents' Professor & Associate Dean, Academic Health Center, University ofMinnesota, Minneapolis, MN, USA. I humbly dedicate this publication to James G.White, M. D., who has been my mentor, teacher, associate and dear friend, during mylong career in platelet research. In the end, my academic success and accomplishmentsover the years, would not have been possible without the support of my wife Yashoda,my daughter Aupama and my son Prashanth. I sincerely acknowledge and appreciatetheir patience and support throughout my career.Gundu H. R. Rao University of MinnesotaProfessor Minneapolis, MN55455 v This page has been reformatted by Knovel to provide easier navigation. Contents Contributors ix Preface xiii Introduction 1 1. Platelet Physiology & Pharmacology: an Overview 1 1.1 Introduction 1 1.2 Role of Platelets in Hemostasis and Thrombosis 2 1.3 Platelet Morphology and Biochemistry 2 1.4 Platelet Physiology 5 1.5 Altered Physiology and Function 6 1.6 Platelet Pharmacology 8 1.7 Platelet Function Inhibitory Drugs 9 1.8 Acknowledgements 14 References 15 Receptor Biology 21 2. Human Platelet Thrombin Receptors and the Two Receptor Model for Platelet Activation 21 2.1 Introduction 21 2.2 Binding Studies 22 2.3 Membrane Microviscosity 24 2.4 Candidate Receptors 26 vi Contents This page has been reformatted by Knovel to provide easier navigation. 2.5 The GPIb-IX-V Complex 27 2.6 Two Receptor Model 31 References 33 3. Platelet Thromboxane Receptors: Biology and Function 38 3.1 Introduction 38 3.2 Biological Effects of TP Receptor Activation 39 3.3 Smooth Muscle Contraction 39 3.4 TP Receptor Structure 41 3.5 TP Receptor Function 49 3.6 Altered TP Receptor Function 58 References 66 4. Collagen Receptors: Biology and Functions 80 4.1 Introduction 80 4.2 Collagens 82 4.3 Von-Willebrand-Factor 83 4.4 P65 84 4.5 CD36 84 4.6 a2b1-Integrin (GPIa/IIa, VLA2, ECMRII) 87 4.7 GPVI/FcRg 89 4.8 Collagen-Induced Signal Transduction 90 References 92 5. Adenosine Receptors: Biology and Function 102 5.1 Introduction 102 5.2 Adenosine Receptors 103 5.3 Antiplatelet Action of Adenosine 104 5.4 Adenosine Production and Platelet Inactivation 106 5.5 Agents Affecting Adenosine Actions 109 Contents vii This page has been reformatted by Knovel to provide easier navigation. 5.6 Adenosine Effects on Intracellular Ca2+ Mobilization 113 5.7 Conclusions 114 References 115 6. Platelet Activating Factor and Platelets 120 6.1 PAF Discovery, Structure and Heterogeneity 120 6.2 PAF Biosynthesis in Platelets 121 6.3 Responses of Platelets to PAF 122 6.4 PAF Receptor and Signal Transduction Pathways in Platelets 123 6.5 Antagonist 124 6.6 PAF Receptor 125 6.7 Phospholipases 126 6.8 Platelet and PAF in Pathophysiological and Disease States 129 6.9 Acknowledgement 133 References 133 7. Platelet Glycoprotein Ib-V-IX: Biology and Function 142 7.1 Introduction 142 7.2 Structure 143 7.3 Post-Translational Modification of GPIb-V-IX 145 7.4 Basic Functions 146 7.5 Signal Transduction 148 7.6 GPIb-V-IX as a Target for Pharmacological Inhibition 149 7.7 Genetic Disorders Affecting GPIb-V-IX 151 7.8 Tissue Specific Expression of GPIb-V-IX Subunits 153 7.9 Future Developments 154 References 155 [...]... both human and animal models Furthermore, a careful correlation between signal-driven responses and platelet function will lead to a better understanding of platelet pathophysiology (27,31, also discussed in other chapters) 1.6 Platelet pharmacology Earlier studies on platelet biochemistry, physiology and function suggest that release of granule contents is essential for recruitment of platelets, irreversible... provide a comprehensive review on various aspects of platelet physiology and pharmacology 1.2 Role of platelets in hemostasis and thrombosis Platelets contribute significantly to the normal hemostatic process (1-6) They play a critical role in the recognition of injured vasculature, formation of hemostatic plugs, prevention of bleeding, retraction of clots and wound healing (6-9) When they develop severe... of Platelet Function 460 21.5 Mechanisms of NO Action on Platelets 461 21.6 Peroxynitrite 462 21.7 Nitric Oxide and Vascular Disorders 463 21.8 Nitric Oxide, Platelets and Septicemia 464 21.9 Pharmacology of NO Generation and Action in the Platelet Microenvironment 465 Acknowledgements 469 References 469 22 Aspirin, Prostaglandins and Platelet Function: Pharmacology. .. aggregation and secretion of granule contents Platelet activation is a prerequisite for the formation of a hemostatic plug and arrest of bleeding at the injured site Although formation of a hemostatic plug is a natural response to injury, the role of platelets in atherosclerosis, thrombosis and stroke are pathological manifestations (1-19) 1.3 Platelet morphology and biochemistry Blood platelets have... discussed in other chapters of this book) Binding results in the activation and stimulation of various effector enzymes and formation of second messengers, leading to aggregation and secretion of granule contents Specific mechanisms involved in the process of centralization of granules and release of their contents are poorly understood(26,27, 31) 1.5 Altered physiology and function Many investigators... easier navigation 1 PLATELET PHYSIOLOGY AND PHARMACOLOGY: AN OVERVIEW Gundu H R Rao, Ph D Departments of Laboratory Medicine & Pathology and Biomedical Engineering Institute Academic Health Center University of Minnesota, Minneapolis, MN 55455, USA 1.1 INTRODUCTION Dr Gundu Rao has spent the last 25 years studying and teaching platelet physiology and pharmacology at the University of Minnesota Over the... antiplatelet and antithrombotic drugs have provided a great deal of excitement and have led to the discovery of many specific antagonists Although the biochemistry of platelets remains highly complex, we now have well defined pathways that could impact platelet functional responses It is very likely that further understanding of the role of these pathways and the specific mechanisms that underlie platelet. .. Activation and Aggregation in Flow 316 14.3 Range of Shear Rates in Normal and Pathological Settings 317 14.4 Flow Regimes and Corresponding Devices Used to Study in Vitro Platelet Aggregation 318 14.2 Ligands and Receptors Involved in Platelet Aggregation 319 14.3 Quantitation of Aggregation: Theoretical and Experimental Approaches 322 14.4 Platelet Aggregation in Non-Stirred Platelet. .. Acid and Platelets 302 13.8 PAF and Phospholipase A2 303 13.9 LDL and Platelet Function 304 Concluding Remarks 304 Acknowledgements 305 References 305 Experimental Physiology 315 14 Platelet Biorheology: Adhesive Interactions in Flow 315 14.1 Introduction: General Overview for Flow Studies of Platelet Aggregation 315 14.2 General Physiology of Platelet. .. Medicine & Pathology and Biomedical Engineering Institute References 1 Marcus AJ, Zucker MB The physiology of blood platelets New York: Crime and Scrutton, 1965 2 Kowalski E, Niewiarowski S Biochemistry of blood platelets New York: Academic Press, 1966 3 Caen J Platelet aggregation Paris: Masson and Cie, 1971 4 Brinkhous KM, Sherman RW, Mostof FK The Platelet Baltimore: Williams and Wilkson Co, 1971 . HANDBOOK OF PLATELET PHYSIOLOGY AND PHARMACOLOGY Edited byGundu H. R. RaoUniversity of MinnesotaKLUWER ACADEMIC. <http://www.wkap.nl>Library of Congress Cataloging-in-Publication Data Handbook of platelet physiology and pharmacology / edited by GunduH.R.
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