International Review of Cytology, A Survey of Cell Biology, Vol. 188

VOLUME 188

SERIES EDITORS Geoffrey H. Bourne James F. Danielli Kwang W. Jeon Martin Friedlander Jonathan Jarvik

1952-1988 1952-1 984 19671984-1 992 1993-1 995

EDITORIAL ADVISORY BOARD Aimee Bakken Eve Ida Barak Rosa Beddington Howard A. Bern Robert A. Bloodgood Dean Bok Stanley Cohen Rene Couteaux Marie A. DiBerardino Laurence Etkin Hiroo Fukuda Elizabeth D. Hay P. Mark Hogarth Anthony P.Mahowald

M. Melkonian Keith E. Mostov Andreas Oksche Vladimir R. Pantic L. Evans Roth Jozef St.Schell Manfred Schliwa Robert A. Smith Wilfred D. Stein Ralph M. Steinman M. Tazawa Donald P. Weeks Robin Wright Alexander L. Yudin

Edited by

Kwang W. Jeon Department of Biochemistry University of Tennessee Knoxville, Tennessee

VOLUME 188

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Front cover photograph: Human epidermal keratinocytes grown on microcarriers. (For more details, see Chapter 1, Figure 1.)

This book is printed on acid-free paper. @ Copyright 0 1999 by ACADEMIC PRESS All Rights Reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopy, recording, or any information storage and retrieval system, without permission in writing from the Publisher. The appearance of the code at the bottom of the first page of a chapter in this book indicates the Publisher’s consent that copies of the chapter may be made for personal or internal use of specific clients. This consent is given on the condition, however, that the copier pay the stated per copy fee through the Copyright Clearance Center, Inc. (222 Rosewood Drive, Danvers, Massachusetts 01923). for copying beyond that permitted by Sections 107 or 108 of the U.S. Copyright Law. This consent does not extend to other kinds of copying, such as copying for general distribution, for advertising or promotional purposes, for creating new collective works, or for resale. Copy fees for pre-1999 chapters are as shown on the title pages. If no fee code appears on the title page, the copy fee is the same as for current chapters. 0074-7696199 $30.00

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Academic Press 24-28 Oval Road, London NWl 7DX, UK http:llwww.hbuk.co.uWapl International Standard Book Number: 0-12-364592- I PRINTED IN THE UNllED STATESOF AMERICA 99 0 0 0 1 02 03 04 EB 9 8 7 6

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Contributors .............................................................

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Microfibrils from the Arterial Subendothelium 1.

Frangoise Fauvel-Lafeve Introduction ........................................................... Structure of Subendothelial Microfibrils ...................................... Microfibrils and Elastinogenesis ........................................... The Thrombogenicity of the Subendothelium .................................

I1. 111. IV . V . The Thrombogenicity of Type VI Collagen ................................... Vl . The Thrombogenicity of Elastin-Associated Microfibrils ......................... VII. Conclusions and Perspectives ............................................ References ...........................................................

1 2 19 19 23 25 29 30

Cultivation and Transplantation of Epidermal Keratinocytes J. V . Terskikh and A . V. Vasiliev 1. II. 111 . IV. V.

Introduction ........................................................... Culture of Keratinocytes ................................................. Behavior of Keratinocytes in Culture ........................................ Transplantation of Cultured Keratinocytes ................................... Concluding Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References ...........................................................

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41 43 47 54 63

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CONTENTS

Retinoids and Maminalian Development G. M. Morriss-Kay and S. J. Ward I. Introduction ........................................................... II . The Retinoic Acid Signaling Pathway in Embryogenesis ........................ 111 . Synthesis and Catabolism of Retinoic Acid in Embryonic Tissues ................. IV. Nuclear Aspects of Retinoic Acid Signal Transduction .......................... V . Retinoic Acid in Craniofacial Development ................................... VI Retinoic Acid and Limb Development ....................................... VII. Retinoic Acid in Other Developing Systems .................................. VIII. Conclusions ........................................................... References ...........................................................

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73 77 85 97 99 106 110 116 119

Gene Expression in the Epididymis 1. II. 111. IV. V.

C. Kirchhoff Introduction ........................................................... Ontogenesis of Epididymal Gene Expression Pattem .......................... Gene Expression in the Adult Epididymis .................................... Factors Regulating Gene Expression in the Adult Epididymis .................... Concluding Remarks .................................................... References ...........................................................

133 135 153 169 184 186

Roles of Reactive Oxygen Species: Signaling and Regulation of Cellular Functions 1. A. Gamaley and 1. V . Klyobin I. II. 111 . IV. V. VI. VII. VIII.

Introduction ........................................................... Mechanisms Inducing Generation of ROS by Cells ............................ ROS Homeostasis in Animal Cells ......................................... Cytotoxic Effects of ROS ................................................ Regulation of Cell Functions by ROS ....................................... ROS as Signal Molecules ................................................ ROS in Plants ......................................................... Concluding Remarks .................................................... References ...........................................................

203 204 211 214 220 223 235 237 238

CONTENTS

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High-Density Lipoprotein: Multipotent Effects on Cells of the Vasculature 1. II. 111 . IV. V. VI. VII. VIII. IX.

Gillian W . Cockerill and Stephen Reed Introduction ........................................................... In Wtro Effects of HDL on Endothelial Cells .................................. Effects of HDL on Leukocyte Activation ..................................... Effects of HDL on Cell Transmigration ...................................... HDL. Thrombosis. and Platelet Function .................................... Effect of HDL on Vasoactive Molecules ..................................... Acute-Phase Response and HDL .......................................... HDL Receptors and Cell Signal Transduction ................................ Concluding Remarks .................................................... References ...........................................................

Index .....................................................................

257 262 267 271 273 276 279 281 286 287 299

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CONTRIBUTORS

Numbers in parentheses indicate the pages on which the author's contributions begin.

Gillian W. Cockerill (257),Department of Cardiovascular Medicine, National Heart and Lung Institute, Imperial College School of Medicine, HammersmithHospital Campus, London W12 ONN, United Kingdom FranGoise Fauvel-Lefeve(1), Unite353 INSERM, lnstitut d'Hematologie, H6pital SaintLouis, 75475 Paris Cedex 10, France I. A. Gamaley (203),Institute of Cytology, Russian Academy of Sciences, 194064 St. Petersburg, Russia Christiane Kirchhoff (133),IHF Institute forHormone and Fertili!y Research, D-22529 Hamburg, Germany

I. V. Klyubin (203),Institute of Cytology, Russian Academy of Sciences, 194064 St. Petersburg, Russia Gillian M. Morriss-Kay (73),Department of Human Anatomy and Genetics, University of Oxford, Oxford OX1 3QX, United Kingdom Stephen C. Reed (257),School of Biosciences, University of Westminstec London WIM US, United Kingdom V. V. Terskikh (41), Instituteof DevelopmentalBiology, RussianAcademy of Sciences, Moscow 117334, Russia A. V. Vasiliev (41), Institute of Developmental Biology, Russian Academy of Sciences, Moscow 117334, Russia Simon J. Ward (73),Department of Biomedical Science, University of Shefield, Sheffield, United Kingdom

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Microfibrils from the Arterial Subendothelium Frangoise Fauvel-Lafeve Unit6 353 INSERM, Institut d'HCmatologie, HGpital Saint-Louis,75475 Pans Cedex 10, France

Microfibrillar structures of the subendothelium are represented by either type VI collagen or elastin-associated microfibrils which are also referred to as fibrillin-containing microfibrils. These structures are present throughout the subendothelium irrespective of the presence of elastin. The localization, structure, and protein composition of microfibrils are reviewed. The arterial subendothelium is thrombogenic despite its very low content in fibrillar collagens. This thrombogenicity is linked to the microfibrillar structures, essentially to type VI collagen and to thrombospondin-containing microfibrils. Their respective ability to bind the von Willebrand factor and to activate blood platelets is discussed. KEY WORDS: Microfibrils,Thrombospondin, Fibrillins, Type VI collagen, Platelets, von Willebrand factor. o 1999 Academic Press

1. Introduction The arterial subendothelium is formed by the endothelial cell basement membrane and the extracellular matrix located between the basement membrane and the internal elastic lamina. The thickness of the subendothelium is variable according to its localization within the vascular tree. In capillaries, endothelial cells are in close contact with the fenestrated elastin lamina, while in aorta, the subendothelium could reach a size of 1 pm (Wight, 1996). In man, the subendothelium thickens with increasing age, a process known as diffuse intimal thickening, and an accelerated intimal thickening indicates a predisposition for the development of atherosclerotic lesions (Mayne, 1987;Stary er al., 1992).Due to its location just beneath endothelial cells which form a barrier between the blood and vascular tissues, components of the subendothelium play an important role in all physiological and Inrernational Review of Cytology,VoL

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Copyright 8 1999 by Academic Press. All rights of reproduction in any form reserved.

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FRANCOISE FAUVEL-IAFEVE

pathological phenomena taking place after an endothelial cell injury or during the circulating cell diapedesis. This vascular wall layer is thus implicated in hemostasis and thrombosis (Hawiger, 1994), arterosclerosis (Ross, 1995),inflammation (Gimbrone, 1995),and tumor cell metastatic dissemination (Crissman et al., 1988). The subendothelium is composed of numerous macromolecules (Kefalides, 1994). In addition to the basement membrane components (type IV collagen, laminin, nidogen, and heparan sulfate proteoglycans), the subendothelium contains large amounts of proteoglycans, fibronectin, and fibrillar structures formed by fibrillar type I11 collagen (Gay et al., 1975) and microfibrils. Microfibrils were first described as a histological entity (Low, 1961). These structures are particularly insoluble and thus much less is known about them than collagens. Since the discovery of type VI collagen (Engval et al., 1986) and fibrillin (Sakai et al., 1986), microfibrillar structures of the subendothelium have been separated into two groups: type VI collagen and elastin-linkedmicrofibrils. In fact, the term microfibrils is now restricted to 10-nm fibrils regardless of whether they are associated with elastin or not. Our results (Fauvel-Lafkve et aZ., 1996) and studies from Zhang et al. (1995) show that noncollagenous microfibrils are also heterogeneous. In this review, the structure and physiological roles of subendothelial microfibrillar structures (microfibrilsand type VI collagen) will be discussed with a special regard to their role in the thrombogenicity of the subendothelium.

II. Structure of Subendothelial Microfibrils A. Ultrastructure In early electron microscopic studies, fine extracellular filaments were described in many tissues. In stained tissue sections they were seen free in the ground substance or linked to collagen fibers or to basement membrane and associated to elastin. Microfibrilswere first defined by Low (1961) as thin filaments with diameters of 50 years) compared to younger men (