Fibrocytes New Insights Into Tissue Repair And Systemic

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 FIBROCYTES
New Insights into
TissueRepair and Systemic Fibrosis

Editor

Richard Bucala
Yale University, USA

World Scientific
NEW JERSEY • LONDON • SINGAPORE • BEIJING • SHANGHAI • HONG KONG • TA I P E I • CHENNAI
 Published by
World Scientific Publishing Co. Pte. Ltd.
5 Toh Tuck Link, Singapore 596224
USA office: 27 Warren Street, Suite 401-402, Hackensack, NJ 07601
UK office: 57 Shelton Street, Covent Garden, London WC2H 9HE

Library of Congress Cataloging-in-Publication Data

Fibrocytes : new insights into tissue repair and systemic fibroses / editor,
Richard Bucala.
p. cm.
Includes bibliographical references and index.
ISBN-13 978-981-256-869-4 -- ISBN-10 981-256-869-7
1. Fibroblasts. 2. Fibroblasts--pathology. 3. Fibroblasts--physiology.
4. Fibroses--physiopathology. 5. Wound Healing--physiology. I. Bucala, Richard.

QP88.23 .F53 2006

612.7'5--dc22 2006051165

British Library Cataloguing-in-Publication Data

A catalogue record for this book is available from the British Library.

Copyright © 2007 by World Scientific Publishing Co. Pte. Ltd.

All rights reserved. This book, or parts thereof, may not be reproduced in any form or by any means,
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SC - Fibrocytes.pmd 1 2/27/2007, 9:40 AM

November 21, 2006 20:22 WSPC/SPI-B408: Fibrocytes: New Insights into Tissue Repair and Systemic Fibroses fm FA1

CONTRIBUTORS

Peter J. Barth, MD
Institute of Pathology
University Hospital Giessen and Marburg GmbH
Location Marburg
Medical Faculty of Philipps — University Malburg
Baldingerstraße, 35043 Malburg
Germany

Rick Bucala, MD, PhD

Yale University School of Medicine
The Anlyan Center, S525
PO Box 208031
300 Cedar Street
New Haven, CT 06520-8031
USA

Jason A. Chesney, MD, PhD

J.G. Brown Cancer Center
University of Louisville
Delia Baxter Research Bldg., Rm. 204E
508 So. Preston Street
Louisville, KY 40202
USA

v
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vi Contributors

Shawn E. Cowper, MD
Department of Dermatology and Pathology
Yale University School of Medicine
P.O. Box 208059
15 York St. LMP 5032
New Haven, CT 06520-8059
USA

Richard Gomer, PhD

Department of Biochemistry and Cell Biology
MS-140, Rice University
Houston, TX 77005-1892
USA

Brigitte N. Gomperts, MD
Department of Pediatrics
Division of Hematology and Oncology
University of Washington
School of Medicine in St. Louis
660 S. Euclid Avenue, Campus Box 8116
St. Louis, MO 63110
USA

Shuichi Kaneko, MD, PhD

Department of Gastroenterology and Nephrology
Kanazawa University
Graduate School of Medical Science
Takara-Machi 13-1
Kanazawa 920-8641,
Japan

Cynthia L. Kucher, MD
Greenwich Hospital — Pathology Department
5 Perryridge Road
Greenwich, CT 06830
USA
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Contributors vii

Amanda C. LaRue, PhD

Department of Veterans Affairs Medical Center
Division of Experimental Hematology
Department of Medicine
Medical University of South Carolina
Charleston, SC 29401
USA

Kouji Matsushima, MD, PhD

Department of Molecular Preventive Medicine
Graduate School of Medicine and
Faculty of Medicine
University of Tokyo
7-3-1, Hongo, Bunkyo-ku,
Tokyo, 113-0033
Japan

Sabrina Mattoli, MD, PhD

Avail Biomedical Research Institute, Basel, Switzerland
Avail GmbH
PO Box 110, CH-4003 Basel
Switzerland

Heather Medbury, PhD

Senior Scientist
Department of Surgery
University of Sydney
Westmead Hospital
Westmead 2145
Australia

Abelardo Medina
UBC Experimental Medicine
344A Jack Bell Research Centre
2660 Oak Street
Vancouver, BC
Canada
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viii Contributors

Makio Ogawa, MD, PhD

Department of Veterans Affairs Medical Center
Division of Experimental Hematology
Department of Medicine
Medical University of South Carolina
Charleston, SC 29401
USA

Darrell Pilling, PhD

Department of Biochemistry and Cell Biology
MS-140
Rice University
Houston, TX 77005-1892
USA

Arnold Postlethwaite, MD
Division of Rhematology
Department of Medicine
University of Tennessee Health Science Center
956 Court Avenue
Coleman Building, G326
Memphis, Tennessee 38163
USA

Norihiko Sakai, MD, PhD

Department of Gastroenterology and Nephrology
Kanazawa University Graduate School of Medical Science
Ishikawa
Japan

Matthhias Schmidt
Avail Biomedical Research Institute, Basel, Switzerland
Avail GmbH
PO Box 110, CH-4003 Basel
Switzerland
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Contributors ix

Paul G. Scott
Department of Surgery
2D3. 81 WMSHC
8440-112 Street
University of Alberta
Edmonton, Alberta
Canada

Robert M. Strieter, MD
Department of Internal Medicine
University of Virginia
School of Medicine
P.O. Box 800466, VA 22908-0466
USA

Edward E. Tredget, MD
Department of Surgery
2D3. 81 WMSHC
8440-112 Street
University of Alberta
Edmonton, Alberta
Canada

Takashi Wada, MD, PhD

Department of Gastroenterology and Nephrology
Kanazawa University Graduate School of Medical Science
13-1 Takara-machi
Kanazawa 920-8641
Japan
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x Contributors

JianFei Wang
Department of Surgery
2D3.81 WMSHC
8440-112 Street
University of Alberta
Edmonton, Alberta
Canada

Yaojiong Wu
Department of Surgery
2D3. 81 WMSHC
8440-112 Street
University of Alberta
Edmonton, Alberta
Canada
November 21, 2006 20:22 WSPC/SPI-B408: Fibrocytes: New Insights into Tissue Repair and Systemic Fibroses fm FA1

CONTENTS

Contributors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v

Chapter 1
Fibrocytes: Discovery of a Circulating Connective Tissue
Cell Progenitor 1
Richard Bucala
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Phenotypic Properties . . . . . . . . . . . . . . . . . . . . . . . 4
Functional Roles in Wound Repair . . . . . . . . . . . . . . . . 6
Role in Health and Disease . . . . . . . . . . . . . . . . . . . . . 7
Wound Repair . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Tumor Biology . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Immunostimulatory Properties . . . . . . . . . . . . . . . . 9
Infectious Diseases . . . . . . . . . . . . . . . . . . . . . . . 10
Scleroderma . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Nephrogenic Systemic Fibrosis (NSF) . . . . . . . . . . . . . 11
Asthma, Acute Lung Injury, and Pulmonary Fibrosis . . . . 12
Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . 14
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

Chapter 2
Fibrocytes: Immunologic Features 19
Jason Chesney
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Requirement of T Cells for the Development of Fibrosis . . . . 20

xi
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xii Contents

T Cell-mediated Fibrosis in Human Disease . . . . . . . . . . . 21

Fibrocytes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
Fibrocytes are Potent Antigen Presenting Cells . . . . . . . . . 24
Fibrocytes Secrete Type I Collagen and
Inflammatory Cytokines . . . . . . . . . . . . . . . . . . . . . 28
Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

Chapter 3
Regulatory Pathways for Fibrocyte Differentiation 37
Darrell Pilling and Richard H. Gomer
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Biology of the Monocyte-Macrophage System . . . . . . . . . . 39
Differentiation of Monocytes into Cell Types Other than
Macrophages . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Peripheral Blood Mononuclear Cells can also Differentiate into
Fibroblast/Stromal Cells . . . . . . . . . . . . . . . . . . . . . 41
Soluble Factors that Regulate Fibrocyte Differentiation . . . . . 42
Regulation of Fibrocytes by Glucose and Insulin . . . . . . . . 44
Regulation of Fibrocyte Differentiation by SAP and
Aggregated IgG . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Regulation of Fibrocyte Differentiation by T cells and
Extracellular Matrix . . . . . . . . . . . . . . . . . . . . . . . 50
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . 52
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52

Chapter 4
Hematopoietic Origin of Fibrocytes 61
Amanda C. LaRue and Makio Ogawa
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Fibroblast Precursors . . . . . . . . . . . . . . . . . . . . . . . . 62
Clonal Transplantation . . . . . . . . . . . . . . . . . . . . . . . 63
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Contents xiii

HSC Origin of Fibroblasts . . . . . . . . . . . . . . . . . . . . . 65

Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . 69
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69

Chapter 5
The Role of Fibrocytes in Post-burn Hypertrophic Scarring 75
JianFei Wang, Yaujiong Wu, Abelardo Medina,
Paul. G. Scott and Edward E. Tredget
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
Altered Structure and Composition of the Extracellular Matrix
of Hypertrophic Scars . . . . . . . . . . . . . . . . . . . . . . 77
A Th2 Polarized Immune Response in Hypertrophic Scar . . . 79
Dysregulated Apoptosis in Hypertrophic Scar . . . . . . . . . . 81
Apoptosis in the Resolution of Inflammation . . . . . . . . 81
Delayed Fibroblast and Myofibroblast Apoptosis
in Hypertrophic Scar . . . . . . . . . . . . . . . . . . . . . . . 82
Increased Levels of the Profibrotic Growth Factors TGF-β
and CTGF in Hypertrophic Scar . . . . . . . . . . . . . . . . 84
Hypertrophic Scarring is Associated with
Blood Borne Fibrocytes . . . . . . . . . . . . . . . . . . . . . 86
Increased Numbers of Fibrocytes can be Cultured from
the Blood of Burn Patients . . . . . . . . . . . . . . . . . . . . 87
Establishment of LSP-1 as a Fibrocyte Marker . . . . . . . . . . 88
Increased Numbers of Fibrocytes in Post-burn Hypertrophic
Scar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
Potential Interaction of Fibrocytes and Endothelial Cells . . . . 92
Possible Interactions of Fibrocytes and Fibroblasts . . . . . . . 93
Elevated TGF-β and CTGF mRNA Levels in Burn Patient
Fibrocytes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
Fibrocytes may Contribute to the Myofibroblast Population . . 94
Possible Role of Fibrocytes in the Polarized Th2 Immune
Response . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
Proposed Role of Fibrocytes in Hypertrophic Scar Formation . 96
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xiv Contents

Summary and Prospects for Future Work . . . . . . . . . . . . 97

Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . 97
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97

Chapter 6
Role in Asthmatic Lung Disease 105
Sabrina Mattoli and Matthias Schmidt
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
Phenotypic and Functional Characteristics of Fibrocytes . . . . 107
Differentiation of Fibrocytes at the Tissue Sites . . . . . . . . . 109
Fibrocytes in Asthma . . . . . . . . . . . . . . . . . . . . . . . . 110
Fibrocytes in Asthma Models . . . . . . . . . . . . . . . . . . . 113
Potential Fibrocyte Chemoattractants in Asthma . . . . . . . . 115
Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119

Chapter 7
Fibrocytes and Other Fibroblast/Myofibroblast Progenitors
in Systemic Sclerosis 125
Arnold E. Postlethwaite
Systemic Sclerosis Clinical Characteristics . . . . . . . . . . . . 126
The Vasculature in SSc . . . . . . . . . . . . . . . . . . . . . 126
The Immune System in SSc . . . . . . . . . . . . . . . . . . 127
The Fibroblast Phenotype in SSc . . . . . . . . . . . . . . . . 128
Accumulation of T cells, Monocytes and Mast Cells in
Clinically Involved Skin in SSc . . . . . . . . . . . . . . . 129
Relationship of Autoimmunity, Vascular Abnormalities and
Fibrosis in SSc (the Old Paradigm) . . . . . . . . . . . . . . . 129
Possible Alternative Sources of Fibroblasts in SSc . . . . . . . . 130
Resident Fibroblast Progenitors . . . . . . . . . . . . . . . . 130
Fibroblast Progenitors from the Circulation in Patients
with SSc and Related Fibrotic Conditions . . . . . . . . . 131
Circulating Fibrocytes and other Progenitors
of Fibroblast-like Cells (FLC) . . . . . . . . . . . . . . . . 132
Overall Hypothetical Scheme for Pathogenesis of SSc . . . . . 134
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Contents xv

New Treatment Strategies for SSc based on Circulating

Fibroblast Progenitors . . . . . . . . . . . . . . . . . . . . . . 136
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137

Chapter 8
Fibrocytes in Interstitial Lung Disease 143
Brigitte N. Gomperts and Robert M. Strieter
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
The Fibrocyte is a Unique Cell Population that has been
Implicated in Wound Repair . . . . . . . . . . . . . . . . 144
Fibrocyte Trafficking . . . . . . . . . . . . . . . . . . . . . . 145
The Fibrocyte Demonstrates Plasticity Compatible with
the Concept of an Adult Stem Cell/Progenitor Cell . . . 148
Fibrocytes in Pulmonary Fibrosis . . . . . . . . . . . . . . . . . 149
Pulmonary Fibrosis . . . . . . . . . . . . . . . . . . . . . . . 149
The Origin of the Fibroblast/Myofibroblast: A Pivotal Cell
in Mediating Fibroproliferation in Pulmonary Fibrosis . 150
Fibrocytes in Asthma . . . . . . . . . . . . . . . . . . . . . . . . 157
Repair and Remodeling of the Airway in Asthma . . . . . . 157
Fibrocytes in Airway Remodeling in Asthma . . . . . . . . 158
Fibrocytes in Pulmonary Vascular Remodeling . . . . . . . . . 159
Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160

Chapter 9
Role of Fibrocytes in Renal Fibrosis 163
Norihiko Sakai, Takashi Wada, Kouji Matsushima
and Shuichi Kaneko
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 164
Fibrocytes in an Experimental Renal Fibrosis Model . . . . . . 165
1) Presence of fibrocytes in fibrotic kidneys . . . . . . . . . 165
2) CCL21/CCR7 signaling regulates fibrocyte infiltration
and renal fibrosis . . . . . . . . . . . . . . . . . . . . . . . 165
3) Infiltration routes of fibrocytes to fibrotic kidneys . . . . 167