Blood Cell Biochemistry Volume 4 Basophil and Mast Cell Degranulation and Recovery Blood Cell Biochemistry Series Editor J. R. Harris, Institute for Cell and Tumor Biology, Heidelberg, Germany Volume 1 Erythroid Cells Edited by J. R. Harris Volume 2 Megakaryocytes, Platelets, Macrophages, and Eosinophils Edited by J. R. Harris Volume 3 Lymphocytes and Granulocytes Edited by J. R. Harris Volume 4 Basophil and Mast Cell Degranulation and Recovery Ann M. Dvorak A Continuation Order Plan is available for this series. A continuation order will bring delivery of each new volume immediately upon publication. Volumes are billed only upon actual shipment. For further informa tion please contact the publisher. Blood Cell Biochemistry Volume 4 Basophil and Mast Cell Degranulation and Recovery Ann M. Dvorak Beth Israel Hospital and Harvard Medical School Boston, Massachusetts Springer Science+Business Media, LLC Llbrary of Congress Cataloglng-ln-Publlcatlon Data Dvorak, Ann M. Basophl1 and mast cell degranulatlon and recovery I Ann M. Dvorak. p. cm. -- (Blood cell blochemistry ; v. 4) Includes blbllographlcal references and Index. ISBN 978-1-4757-9527-1 ISBN 978-1-4757-9525-7 (eBook) DOI 10.1007/978-1-4757-9525-7 1. Mast cells--Physlology. 2. Basophils--Physiology. 1. Title. II. Serles. [DNLM, 1. Basophi ls--physlology. 2. Cell Degranulatlon -phySlology. 3. Mast Cells--physiology. WH 200 D988bl OR185.8.M35D8S 1991 S1S.07'9--dc20 DNLM/OLC for Llbrary of Congress 91-3005 CIP ISBN 978-1-4757-9527-1 © 1991 Springer Science+Business Media New York Origina11y published by Plenum Press, New York in 1991 Softcover reprint ofthe hardcover Ist edition 1991 All rights reserved No part of this book may be reproduced, stored in a retrieval system, -or transmitted in any form or by any means, electronic, mechimica1, photocopying, microfilming, recording, or otherwise, without written permission from the Publisher Preface Basophils and mast cells are similar but unique secretory cells with a well-documented role in immediate-hypersensitivity reactions. The presence of these cells in various cell mediated hypersensitivity reactions, in tissues of multiple diseases, and as a component of the host reaction to injury and repair in numerous circumstances is well known. Release of stored and newly generated mediators of inflammation from basophils and mast cells contributes to the cascade of pathogenetic events in circumstances under which these release reactions occur. Despite insights acquired through studies of these pathologic events, the role of basophils and mast cells and their secretory products in health is not known. In this book, I review much of the structural information regarding basophils and mast cells of multiple species. Ultrastructural studies of rat mast cells historically precede and quantitatively exceed similar studies of basophils and mast cells of other species. Therefore, I first review these background studies as an entity. Then I discuss the contents of two prominent organelles-granules and lipid bodies-in basophils and mast cells of several species. The ultrastructural morphology of basophils and mast cells in three species is presented in detail to establish appropriate guidelines for their recognition and to provide general rules for analysis which are appropriate for the identification of these cells in other species as well. Discussion of secretory mechanisms for mediator release from basophils and mast cells concentrates on ultrastructural studies of these cells in several species with which we are most familiar. Similarly, I review the recovery potential, as imaged in ultrastructural studies, of guinea pig basophils and human mast cells following stimulation of anaphylactic degranulation in short-term cultures. Presentation of these structural studies as a unit will provide necessary background information for future functional studies designed to elucidate the role of basophils and mast cells in health as well as in disease. I acknowledge the stimulating discussion, helpful advice, and continuous encourage ment of my husband and collaborator, Dr. Harold F. Dvorak. Similarly, many of these studies could not have been done without the excellent assistance of my colleagues Drs. Stephen Galli, Teruko Ishizaka, and Lawrence Lichtenstein, and the numerous members of their respective laboratories. A large number of colleagues in many other distinguished laboratories have been a ready source of material for some studies. Manuscript processing and editorial assistance were provided by Peter K. Gardner, v vi Preface and expert technical assistance was provided by Ellen Morgan, Rita Monahan-Earley, Justine Osage, Kathryn Pyne, Patricia Estrella, Joanne Smith, Linda Letourneau, and Susan Kissell. All of these studies have been made possible by Public Health Service grants CA15136, CA19141, and CA28834, representing continuous support by the National In stitutes of Health of the research efforts in basophil and mast cell biology in my laboratory. Ann M. Dvorak Boston, Massachusetts Contents Chapter I Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ....................... . Chapter 2 Rat Mast Cells Overview.......................................................... 3 2.1. Ultrastructural Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2.2. Development In Vivo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 2.3. Development In Vitro . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 2.4. Granules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 2.5. Degranulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 2.6. Recovery from Degranulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Chapter 3 Biochemical Contents of Granules and Lipid Bodies-Two Distinctive OrgaPelles Found in Basophils and Mast Cells Overview . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 3.1. Granules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 3.2. Lipid Bodies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52 Chapter 4 Ultrastructural Morphology of Basophils and Mast Cells Overview . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 4.1. Guinea Pig Basophils . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 4.2. Guinea Pig Mast Cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71 4.3. Mouse Basophils . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78 4.4. Mouse Mast Cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79 4.5. Human Basophils . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90 4.6. Human Mast Cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92 vii viii Contents Chapter 5 Degranulation of Basophils and Mast Cells Overview . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 1 5 .1. Anaphylactic Degranulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101 5. 2. Piecemeal Degranulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 202 Chapter 6 Recovery of Basophils and Mast Cells from Degranulation Overview . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 277 6.1. Guinea Pig Basophils . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 279 6.2. Human Mast Cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 312 Chapter 7 Concluding Remarks 369 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 1 Index............................................................. 393 Chapter 1 Introduction Basophils and mast cells, two unique but closely related cells, have fascinated investiga tors since their original descriptions by Ehrlich (1877, 1879). We also succumbed to this fascination when we identified large numbers of basophils in skin lesions of guinea pigs that had been sensitized to antigen emulsified in incomplete Freund's adjuvant and sampled for ultrastructural analysis within 1 to 2 weeks after skin testing-reactions we called cutaneous basophil hypersensitivity (CBH) to distinguish them from classical de layed hypersensitivity (DH) skin reactions (H. F. Dvorak, 1974; H. F. Dvorak et al., 1970). The study of basophil and mast cell biology has been a major part of our ultrastruc tural research efforts during the past 20 years. In this review, I will draw most heavily on my knowledge of basophils and mast cells from the three species with which I have the most experience (guinea pig basophils, mouse mast cells, and human basophils and mast cells), recognizing that rat mast cells have served as research models for others for many years (Anderson and Uvnas, 1975, Anderson et al., 1973; Benditt, 1958; Benditt and Arase, 1959; Benfey et al., 1987; Bloom and Chakravarty, 1970; Burwen and Satir, 1977a,b; Chi et al., 1975, 1976, 1982; Chock and Schmauder-Chock, 1985; Combs, 1966; Combs et al., 1965; Enerback and Lundin, 1974; Enerback and Rundquist, 1981; Fawcett, 1954; Foreman, 1977; Gibson and Miller, 1986; Henderson et al., 1980a,b; Ishizaka et al., 1976, 1977a; Kriiger and Lagunoff, 1981; Kriiger et al., 1974; Kurosawa and Parker, 1986; Lagunoff, 1972b; Lagunoff, 1973; Lagunoff and Benditt, 1960, 1963; Lagunoff and Chi, 1976; Lagunoff and Pritzl, 1976; Lawson et al., 1975, 1977, 1978; Levi-Schaffer et al., 1985; Mann, 1969; Nemeth and Rolich, 1982; Nielsen and Clausen, 1982; Nielsen et al., 1981a,b; Padawer, 1963, 1969, 1970, 1971, 1974; Padawer and Gordan, 1955; Rolich et al., 1971a,b; Spicer, 1960; Stevens et al., 1983; Sullivan and Parker, 1979; Thon and Uvnas, 1966; Uvnas, 1974a,b,c, 1978; Yong, 1981). Our experience with rat mast cells is limited (Login et al., 1987). For this reason and for historical ones, we will initially review rat mast cell studies (including ultrastructural morphology, development in vivo and in vitro, granule contents, anaphylactic degranulation, other secretory mechanisms, and recovery from degranulation) as a unit, prior to review of similar areas of research with guinea pig and human basophils and mouse and human mast cells. Basophils and mast cells were initially identified by Ehrlich ( 1877, 1879) because of 1