The Pigmentary System: Physiology and Pathophysiology

Edited by

James J. Nordlund Dermatologist and Professor Emeritus, Group Health Associates, Cincinnati, OH, USA

Raymond E. Boissy Professor of Dermatology and Cell Biology, University of Cincinnati College of Medicine, Cincinnati, OH, USA

Vincent J. Hearing Chief, Pigment Cell Biology Section, Laboratory of Cell Biology, National Institutes of Health, Bethesda, MD, USA

Richard A. King Director, Genetics Division, Department of Medicine, Institute of Human Genetics, University of Minnesota, Minneapolis, MN, USA

William S. Oetting Associate Professor, Genetics Division, Department of Medicine Institute of Human Genetics, University of Minnesota, Minneapolis, MN, USA

Jean-Paul Ortonne Professor of Dermatology and Chairman, Department of Dermatology, University of Nice-Sophia Antipolis, Nice, France

SECOND EDITION

The Pigmentary System: Physiology and Pathophysiology

A

D B

C E F

H

G I A: These two individuals illustrate the wide range of skin and hair color. [Marianne Greenwood (right) graciously shared the photographs . (from her book Varför Grater Puman?) that compose this frontispiece.] B: Classical Celtic women with blue eyes and red hair. C: Typical Scandinavian with blue eyes. D: Native American. E: A Peruvian girl. F: Himalayan woman and child. G: A Venezuelan woman. H: Two teenagers from New Guinea. I: Man from the New Hebrides Islands. See Chapter 27 for further information. The Pigmentary System: Physiology and Pathophysiology

Edited by

James J. Nordlund Dermatologist and Professor Emeritus, Group Health Associates, Cincinnati, OH, USA

Raymond E. Boissy Professor of Dermatology and Cell Biology, University of Cincinnati College of Medicine, Cincinnati, OH, USA

Vincent J. Hearing Chief, Pigment Cell Biology Section, Laboratory of Cell Biology, National Institutes of Health, Bethesda, MD, USA

Richard A. King Director, Genetics Division, Department of Medicine, Institute of Human Genetics, University of Minnesota, Minneapolis, MN, USA

William S. Oetting Associate Professor, Genetics Division, Department of Medicine Institute of Human Genetics, University of Minnesota, Minneapolis, MN, USA

Jean-Paul Ortonne Professor of Dermatology and Chairman, Department of Dermatology, University of Nice-Sophia Antipolis, Nice, France

SECOND EDITION © 2006 Blackwell Publishing Ltd Blackwell Publishing, Inc., 350 Main Street, Malden, Massachusetts 02148-5020, USA Blackwell Publishing Ltd, 9600 Garsington Road, Oxford OX4 2DQ, UK Blackwell Publishing Asia Pty Ltd, 550 Swanston Street, Carlton, Victoria 3053, Australia

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First published 1998 by Oxford University Press Second edition 2006 1 2006

Library of Congress Cataloging-in-Publication Data

The pigmentary system : physiology and pathophysiology / edited by James J. Nordlund . . . [et al.]. — 2nd ed. p. ; cm. Includes bibliographical references and index. 1. Pigmentation disorders. 2. . I. Nordlund, James J. [DNLM: 1. Pigmentation Disorders–physiopathology. 2. Melanocytes. 3. Pigmentation–physiology. WR 265 P6309 2006] RL790.P53 2006 616.5¢5–dc22 2005030369

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ISBN-13: 978-1-4051-2034-0 ISBN-10: 1-4051-2034-7

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Contributors, x 15 Chemistry of , 282 Foreword, xv Shosuke Ito & Kazumasa Wakamatsu Preface, xvii 16 The Physical Properties of Melanins, 311 Acknowledgements, xviii Tadeusz Sarna & Harold A. Swartz Frontispiece can be found between pages ii and iii 17 Photobiology of Melanins, 342 Antony R. Young Part I: The Physiology of the Pigmentary System 18 Toxicological Aspects of and Section 1: Historical and Comparative Perspectives of Melanogenesis, 354 the Pigmentary System, 3 Edward J. Land, Christopher A. Ramsden, 1 A History of the Science of Pigmentation, 5 & Patrick A. Riley Sidney N. Klaus 19 Regulation of Pigment Type Switching by Agouti, 2 Comparative Anatomy and Physiology of Pigment Cells Melanocortin Signaling, Attractin, and Mahoganoid, in Nonmammalian Tissues, 11 395 Joseph T. Bagnara & Jiro Matsumoto Gregory S. Barsh 20 Human Pigmentation: Its Regulation by Ultraviolet Section 2: The Science of Pigmentation, 61 Light and by Endocrine, Paracrine, and Autocrine 3 General Biology of Mammalian Pigmentation, 63 Factors, 410 Walter C. Quevedo Jr. & Thomas J. Holstein Zalfa Abdel-Malek & Ana Luisa Kadekaro 4 Extracutaneous Melanocytes, 91 21 Paracrine Interactions of Melanocytes in Pigmentary Raymond E. Boissy & Thomas J. Hornyak Disorders, 421 5 Regulation of Melanoblast Migration and Genji Imokawa Differentiation, 108 22 Growth Factor Receptors and Signal Transduction David M. Parichy, Mark V. Reedy, Regulating the Proliferation and Differentiation of & Carol A. Erickson Melanocytes, 445 6 Melanoblast Development and Associated Ruth Halaban & Gisela Moellmann Disorders, 140 23 Aging and Senescence of Melanocytes, 464 Richard A. Spritz Debdutta Bandyopadhyay & Estela E. Medrano 7 Biogenesis of , 155 24 The Genetics of Melanoma, 472 Raymond E. Boissy, Marjan Huizing, Vanessa C. Gray-Schopfer & Dorothy C. Bennett & William A. Gahl 25 The Transformed Phenotype of Melanocytes, 489 8 Trafficking and Transfer, 171 Dong Fang & Meenhard Herlyn Glynis A. Scott 9 Melanosome Processing in Keratinocytes, 181 Part II: The Pathophysiology of Pigmentary Disorders H. Randolph Byers Section 3: An Overview of and 10 The Regulation of Melanin Formation, 191 its Disorders, 497 Vincent J. Hearing 26 A More Precise Lexicon for Pigmentation, 11 The Tyrosinase Gene Family, 213 Pigmentary Disorders, and “Chromatic” Abnormalities, William S. Oetting & Vijayasaradhi Setaluri 499 12 Molecular Regulation of Melanin Formation: James J. Nordlund, Tania Cestari, Pearl Grimes, Melanosome Transporter Proteins, 230 Henry Chan, & Jean-Paul Ortonne Murray H. Brilliant 27 The Normal Color of Human Skin, 504 13 Transcriptional Regulation of Function, 242 James J. Nordlund & Jean-Paul Ortonne Kazuhisa Takeda & Shigeki Shibahara 28 Mechanisms that Cause Abnormal Skin 14 Enzymology of Melanin Formation, 261 Color, 521 Francisco Solano & José C. García-Borrón Jean-Paul Ortonne & James J. Nordlund

v CONTENTS

Section 4: Disorders of , Bird-Headed Dwarfism (Seckel Syndrome), 657 and Hypochromia, 539 Stan P. Hill 29 Genetic Hypomelanoses: Disorders Characterized by Down Syndrome, 658 Congenital White Spotting — , Waardenburg Rosemary Geary Syndrome, and Related Genetic Disorders of Fisch Syndrome, 659 Melanocyte Development — Clinical Aspects, 541 Stan P. Hill Richard A. Spritz Premature Canities, 659 30 Genetic Hypomelanoses: Acquired Depigmentation, 551 James J. Nordlund Rozycki Syndrome, 551 Mandibulofacial Dysostosis (Treacher Collins Jean L. Bolognia Syndrome), 660 Vulgaris, 551 Rosemary Geary James J. Nordlund, Jean-Paul Ortonne, , 660 & I. Caroline Le Poole Peggy Tong 31 Genetic Hypomelanoses: Generalized PHC Syndrome (Böök Syndrome), 661 Hypopigmentation, 599 Stan P. Hill Oculocutaneous , 599 Pierre Robin Syndrome, 661 Richard A. King & William S. Oetting James J. Nordlund Albinoid Disorders, 613 Prolidase Deficiency, 662 Philippe Bahadoran & Jean-Paul Ortonne Pranav B. Sheth Ataxia Telangiectasia, 621 34 Metabolic, Nutritional, and Endocrine Disorders, 664 Anne-Sophie Gadenne Metabolic and Nutritional Hypomelanoses, 664 Hallerman–Streiff Syndrome, 623 Peter S. Friedmann James J. Nordlund Hypomelanosis Associated with Endocrine Disorders, Histidinemia, 623 667 Marnie D. Titsch Peter S. Friedmann Homocystinuria, 625 35 Chemical, Pharmacologic, and Physical Agents Causing Allan D. Mineroff Hypomelanoses, 669 Oculocerebral Syndrome with Chemical and Pharmacologic Agents Causing Hypopigmentation, 626 Hypomelanoses, 669 Jean L. Bolognia Stefania Briganti, Monica Ottaviani, , 630 & Mauro Picardo Jean-Paul Ortonne Physical Agents Causing Hypomelanoses, 683 Kappa-Chain Deficiency, 631 Jean-Philippe Lacour Jean-Paul Ortonne 36 Infectious Hypomelanoses, 686 Menkes’ Kinky Hair Syndrome, 631 Jean-Philippe Lacour Tanusin Ploysangam 37 Inflammatory Hypomelanoses, 699 Phenylketonuria, 634 Jean-Philippe Lacour Allan D. Mineroff 38 Hypomelanoses Associated with Melanocytic 32 Genetic Hypomelanoses: Localized Neoplasia, 705 Hypopigmentation, 636 Lieve Brochez, Barbara Boone, “Hypomelanosis of Ito” and Mosaicism, 636 & Jean-Marie Naeyaert Wolfgang Küster & Rudolf Happle 39 Miscellaneous Hypomelanoses: Depigmentation, 725 Focal Dermal Hypoplasia, 645 Alezzandrini Syndrome, 725 James J. Nordlund Wiete Westerhof, David Njoo, & Henk E. Menke Hypomelanosis with Punctate Keratosis of the Palms Idiopathic Guttate Hypomelanosis, 726 and Soles, 646 Wiete Westerhof, David Njoo, & Henk E. Menke Jean L. Bolognia Leukoderma Punctata, 729 Darier–White Disease (Keratosis Follicularis; Wiete Westerhof, David Njoo, & Henk E. Menke 124200), 647 Lichen Sclerosus et Atrophicus, 731 Jean L. Bolognia Philippe Bahadoran Depigmentosus, 649 Vagabond Leukomelanoderma, 732 Stella D. Calobrisi Wiete Westerhof, David Njoo, & Henk E. Menke Tuberous Sclerosis Complex, 652 Vogt–Koyanagi–Harada Syndrome, 734 Pranav B. Sheth Wiete Westerhof, David Njoo, & Henk E. Menke 33 Genetic Hypomelanoses: Disorders Characterized by Westerhof Syndrome, 741 Hypopigmentation of Hair, 657 Wiete Westerhof, David Njoo, & Henk E. Menke vi CONTENTS

40 Miscellaneous Hypomelanoses: Hypopigmentation, 745 Hereditary Sclerosing , 795 Disseminated Hypopigmented Keratoses, 745 Eulalia Baselga & Nancy Burton Esterly Wiete Westerhof, David Njoo, & Henk E. Menke Mendes Da Costa Disease, 796 Hypermelanocytic Punctata et Guttata Eulalia Baselga & Nancy Burton Esterly Hypomelanosis, 746 Naegeli–Franceschetti–Jadassohn Syndrome, 798 Wiete Westerhof, David Njoo, & Henk E. Menke Eulalia Baselga & Nancy Burton Esterly Progressive Macular Hypomelanosis, 748 Reticulated Acropigmentation of Dohi Henk E. Menke, Germaine Relyveld, David Njoo, (Dyschromatosis Symmetrica Hereditaria), 799 & Wiete Westerhof Eulalia Baselga & Nancy Burton Esterly Sarcoidosis, 751 Reticulate Acropigmentation of Kitamura, 802 Wiete Westerhof, David Njoo, & Henk E. Menke Eulalia Baselga & Nancy Burton Esterly 41 Miscellaneous Hypomelanoses: Extracutaneous Loss of Rothmund–Thomson Syndrome, 804 Pigmentation, 754 Eulalia Baselga & Nancy Burton Esterly Alopecia Areata, 754 45 Genetic Epidermal Syndromes with Café-au-lait Wiete Westerhof Macules, 809 Heterochromia Irides, 756 Familial Multiple Café-au-lait Spots, 809 Wiete Westerhof, David Njoo, & Henk E. Menke Nancy Burton Esterly Senile Canities, 760 Neurofibromatosis, 809 Wiete Westerhof, David Njoo, & Henk E. Menke Nancy Burton Esterly, Eulalia Baselga, Sudden Whitening of Hair, 764 & Sheila S. Galbraith Wiete Westerhof, David Njoo, & Henk E. Menke Neurofibromatosis 1 with Noonan 42 Hypochromia without Hypomelanosis, 767 Syndrome, 816 Jean-Philippe Lacour Nancy Burton Esterly McCune–Albright Syndrome, 817 Section 5: Disorders of and Sheila S. Galbraith & Nancy Burton Esterly Hyperchromia, 769 Segmental Neurofibromatosis, 819 43 Genetic Epidermal Syndromes: Disorders Characterized Nancy Burton Esterly & Eulalia Baselga by Generalized Hyperpigmentation, 771 Silver–Russell Syndrome, 820 Adrenoleukodystrophy, 771 Nancy Burton Esterly, Eulalia Baselga, Sheila S. Galbraith & Nancy Burton Esterly & Sheila S. Galbraith Familial Progressive Hyperpigmentation, 774 Watson Syndrome, 823 Nancy Burton Esterly, Eulalia Baselga, Beth A. Drolet, Nancy Burton Esterly Susan Bayliss Mallory, & Sharon A. Foley 46 Genetic Epidermal Pigmentation with Lentigines, Fanconi Anemia, 776 824 Sheila S. Galbraith & Nancy Burton Esterly Simplex, 824 Gaucher Disease, 778 Mary K. Cullen Sheila S. Galbraith & Nancy Burton Esterly Lentigo Senilis et Actinicus, 829 44 Genetic Epidermal Syndromes: Disorders Characterized Mary K. Cullen by Reticulated Hyperpigmentation, 780 Centrofacial , 837 Berlin Syndrome, 780 Mary K. Cullen Eulalia Baselga & Nancy Burton Esterly LEOPARD Syndrome, 842 Cantu Syndrome, 781 Mary K. Cullen Eulalia Baselga & Nancy Burton Esterly Carney Complex, 851 Kindler Syndrome, 781 Mary K. Cullen Eulalia Baselga & Nancy Burton Esterly Other Lentiginoses, 863 Dermatopathia Pigmentosa Reticularis, 784 Mary K. Cullen Eulalia Baselga & Nancy Burton Esterly 47 Genetic Epidermal Syndromes: Localized Dyschromatosis Universalis Hereditaria, 786 Hyperpigmentation, 873 Eulalia Baselga & Nancy Burton Esterly Anonychia with Flexural Pigmentation, 873 Epidermolysis Bullosa with Mottled Nancy Burton Esterly, Eulalia Baselga, Pigmentation, 788 & Beth A. Drolet Eulalia Baselga & Nancy Burton Esterly , 873 Familial Mandibuloacral Dysplasia, 790 Sheila S. Galbraith & Nancy Burton Esterly Eulalia Baselga & Nancy Burton Esterly Periorbital Hyperpigmentation, 879 Hereditary Acrokeratotic Poikiloderma, 792 Nancy Burton Esterly, Eulalia Baselga, Eulalia Baselga & Nancy Burton Esterly & Beth A. Drolet

vii CONTENTS

Pigmentary Demarcation Lines, 880 Erythema Dyschromicum Perstans, 933 Sheila S. Galbraith & Nancy Burton Esterly Norman Levine & Cynthia Burk Dowling–Degos Disease, 882 Erythromelanosis Follicularis Faciei et Colli, 935 Sheila S. Galbraith & Nancy Burton Esterly Norman Levine & Cynthia Burk 48 Genetic Epidermal Syndromes: Disorders of Aging, 884 Erythrose Péribuccale Pigmentaire of Brocq, 937 Acrogeria, 884 Norman Levine & Cynthia Burk Nancy Burton Esterly, Eulalia Baselga, Extracutaneous Neuroendocrine Melanoderma, 938 Peter M. H. Chan, & Beth A. Drolet Norman Levine & Cynthia Burk Metageria, 886 Felty Syndrome and Rheumatoid Arthritis, 939 Nancy Burton Esterly, Eulalia Baselga, Norman Levine & Cynthia Burk Peter M. H. Chan, & Beth A. Drolet Hyperpigmentation Associated with Human Progeria, 886 Immunodeficiency Virus (HIV) Infection, 941 Nancy Burton Esterly, Eulalia Baselga, Philippe Bahadoran Peter M. H. Chan, & Beth A. Drolet Melanoacanthoma, 946 Xeroderma Pigmentosum, 889 Norman Levine & Cynthia Burk Anita P. Sheth, Nancy Burton Esterly, Phytophotodermatitis, 948 Eulalia Baselga, Peter M. H. Chan, Norman Levine & Cynthia Burk & Beth A. Drolet Polyneuropathy, Organomegaly, Endocrinopathy, Werner Syndrome, 894 M Protein, and Skin Changes: POEMS Syndrome, 951 Nancy Burton Esterly, Eulalia Baselga, James J. Nordlund Peter M. H. Chan, Beth A. Drolet, Urticaria Pigmentosum and Mastocytosis, 954 & Cindy L. Lamerson James J. Nordlund 49 Congenital Epidermal Hypermelanoses, 898 , 959 , 898 Vlada Groysman & Norman Levine Susan Bayliss Mallory, Peggy L. Chern, Riehl’s , 961 & Sharon A. Foley Scott Bangert & Norman Levine Ectodermal Dysplasias, 901 Atrophoderma of Pasini et Pierini, 963 Susan Bayliss Mallory, Peggy L. Chern, James J. Nordlund, Norman Levine, & Sharon A. Foley Charles S. Fulk, & Randi Rubenzik Transient Neonatal Pustular Melanosis, 905 Hyperpigmentation Associated with Scleromyxedema Susan Bayliss Mallory, Peggy L. Chern, and Gammopathy, 965 & Sharon A. Foley Kazunori Urabe, Juichiro Nakayama, & Yoshiaki Hori Universal Acquired Melanosis, 906 Ichthyosis Nigricans, Keratoses, and Epidermal Susan Bayliss Mallory, Peggy L. Chern, Hyperplasia, 965 & Sharon A. Foley James J. Nordlund 50 Acquired Epidermal Hypermelanoses, 907 Morphea and Scleroderma, 967 , 907 James J. Nordlund Norman Levine & Cynthia Burk Pigmentary Changes Associated with Addison Acromelanosis Progressiva, 914 Disease, 969 Norman Levine & Cynthia Burk Cindy L. Lamerson & James J. Nordlund Becker Nevus, 915 Pigmentary Changes Associated with Cutaneous Norman Levine & Cynthia Burk Lymphomas, 972 Café-au-lait Spots, 917 Debra L. Breneman Norman Levine & Cynthia Burk 51 Hypermelanosis Associated with Gastrointestinal Carcinoid Syndrome, 919 Disorders, 979 Norman Levine & Cynthia Burk Porphyria Cutanea Tarda, 979 Confluent and Reticulated Papillomatosis, 922 Eun Ji Kwon & Victoria P. Werth Norman Levine & Cynthia Burk Cronkhite–Canada Syndrome, 983 Cutaneous Amyloidosis, 924 Eun Ji Kwon, James J. Nordlund, Norman Levine & Cynthia Burk & Victoria P. Werth Dermatosis Papulosa Nigra, 928 Hemochromatosis and Hemosiderosis, 986 Norman Levine & Cynthia Burk Joerg Albrecht & Victoria P. Werth Ephelides (), 929 Primary Biliary Cirrhosis, 992 Norman Levine & Cynthia Burk Joerg Albrecht & Victoria P. Werth Erythema ab Igne, 931 Inflammatory Bowel Disease and Pigmentation, 995 Norman Levine & Cynthia Burk Joerg Albrecht & Victoria P. Werth viii CONTENTS

Pellagra, 995 56 Pigmentary Abnormalities and Discolorations of the Eun Ji Kwon & Victoria P. Werth Mucous Membranes, 1069 Peutz–Jeghers Syndrome, 999 John C. Maize, Jr. & John C. Maize, Sr. Nancy Burton Esterly, Eulalia Baselga, & Beth A. Drolet Section 7: Benign Neoplasms of Melanocytes, 1091 52 Acquired and Congenital Dermal Hypermelanosis, 1003 57 Common Benign Neoplasms of Melanocytes, 1093 Sacral Spot of Infancy, 1003 Pigmented Spindle Cell Nevi, 1093 Sang Ju Lee, Seung Kyung-Hann, & Sungbin Im Julie V. Schaffer & Jean L. Bolognia Nevus of Ota, 1006 Speckled Lentiginous Nevus (Nevus Spilus), 1098 Sang Ju Lee, Seung Kyung-Hann, & Sungbin Im Julie V. Schaffer & Jean L. Bolognia Nevus of Ito, 1012 Melanocytic (Nevocellular) Nevi and Their Biology, Sang Ju Lee, Seung Kyung-Hann, & Sungbin Im 1112 Phakomatosis Pigmentovascularis, 1013 Julie V. Schaffer & Jean L. Bolognia Sang Ju Lee, Seung Kyung-Hann, & Sungbin Im 58 Rare Benign Neoplasms of Melanocytes, 1148 Other Congenital Dermal Melanocytosis, 1015 Nevus Aversion Phenomenon, 1148 Sang Ju Lee, Seung Kyung-Hann, & Sungbin Im James J. Nordlund Acquired Dermal Melanocytosis, 1016 Melanotic Neuroectodermal Tumor of Infancy, 1148 Sang Ju Lee, Seung Kyung-Hann, & Sungbin Im Julie V. Schaffer & Jean L. Bolognia Carleton–Biggs Syndrome, 1017 Pilar Neurocristic Hamartoma, 1157 Sang Ju Lee, Seung Kyung-Hann, & Sungbin Im Julie V. Schaffer & Jean L. Bolognia Acquired Bilateral Nevus of Ota-like Macules (ABNOM), 1017 Section 8: Treatment of Pigmentary Disorders, 1163 Sang Ju Lee, Seung Kyung-Hann, & Sungbin Im 59 Topical Treatment of Pigmentary Disorders, 1165 Blue Macules Associated with Progressive Systemic Rebat M. Halder & James J. Nordlund Sclerosis, 1018 60 Chemophototherapy of Pigmentary Disorders, 1175 Sang Ju Lee, Seung Kyung-Hann, & Sungbin Im Rebat M. Halder & James J. Nordlund 53 Mixed Epidermal and Dermal Hypermelanoses and 61 UVB Therapy for Pigmentary Disorders, 1183 Hyperchromias, 1020 Thierry Passeron & Jean-Paul Ortonne , 1020 62 Sunscreens and Cosmetics, 1188 Sang Ju Lee, Seung Kyung-Hann, & Sungbin Im James J. Nordlund & Rebat M. Halder Melanosis from Melanoma, 1023 63 Surgical Treatments of Pigmentary Disorders, 1191 Sang Ju Lee, Seung Kyung-Hann, & Sungbin Im Rebat M. Halder & James J. Nordlund 54 Drug-induced or -related Pigmentation, 1026 64 Laser Treatment of Pigmentary Disorders, 1198 Peter A. Lio & Arthur J. Sober Rebat M. Halder, Lori M. Hobbs, & James J. Nordlund Section 6: Disorders of Pigmentation of the Nails and Mucous Membranes, 1055 Index, 1205 55 The Melanocyte System of the Nail and its Disorders, Plate section can be found between pages 494 and 495 1057 Robert Baran, Christophe Perrin, Luc Thomas, & Ralph Braun

ix Contributors

Zalfa Abdel-Malek PhD Raymond E. Boissy PhD Research Professor of Dermatology, Department of Dermatology, Professor of Dermatology and Cell Biology, Department of University of Cincinnati College of Medicine, Cincinnati, OH, USA Dermatology, University of Cincinnati College of Medicine, Cincinnati, OH, USA Joerg Albrecht MD Department of Dermatology, University of Pennsylvania, Jean L. Bolognia MD Philadelphia, PA, USA Professor of Dermatology, Department of Dermatology, Yale University School of Medicine, New Haven, CT, USA Mayra Alvarez-Franco (deceased) Formerly of the Departments of Dermatology and Pathology, Yale Barbara Boone MD University School of Medicine, New Haven, CT, USA Resident in Dermatology, Universitair Ziekenhuis Ghent, Ghent, Belgium Joseph T. Bagnara PhD Professor Emeritus, Department Cell Biology and Anatomy, Ralph P. Braun MD University of Arizona College of Medicine, Tucson, AZ, USA Assistant Professor, Pigmented Skin Lesion Unit, Department of Dermatology, University Hospital, Geneva, Switzerland Philippe Bahadoran MD PhD Assistant Professor of Medicine, Service de Dermatologie, Debra L. Breneman MD Hôpital l’Archet, Nice, France Associate Professor of Dermatology, University of Cincinnati, Cincinnati, OH, USA Debdutta Bandyopadhyay PhD Instructor of Dermatology, Huffington Center on Aging, Baylor Stefania Briganti PhD College of Medicine, Houston, TX, USA San Gallicano Dermatological Institute, Rome, Italy

Scott D. Bangert MD Murray H. Brilliant PhD Banner Good Samaritan Hospital, Department of Medical Lindholm Professor of Genetics, Department of Pediatrics, Education, Phoenix, AZ, USA University of Arizona College of Medicine, Tucson, AZ, USA

Robert Baran MD Lieve Brochez MD PhD Head of Nail Disease Center, Cannes, France Professor of Dermatology, Department of Dermatology, University Hospital of Ghent, Ghent, Belgium Gregory S. Barsh MD PhD Professor of Genetics and Pediatrics, Beckman Center for Molecular Cynthia J. Burk MD and Genetic Medicine, Stanford University School of Medicine, Section of Dermatology, University of Arizona Health Sciences Stanford, CA, USA Center, Tucson, AZ, USA

Eulalia Baselga MD Stella D. Calobrisi MD Consultant in Pediatric Dermatology, Hospital de la Santa Creu I Medical Director, The Dermatology Clinic, Boca Raton, FL, USA Sant Pau, Barcelona, Spain Tania Cestari MD PhD Dorothy C. Bennett MA PhD Associate Professor of Dermatology, Department of Dermatology, Professor of Cell Biology, St George’s Hospital, University of University of Rio Grande do Sul, Hospital de Clínicas de Porto London, London, UK Alegre, Brazil x CONTRIBUTORS

Henry H. L. Chan MBBS MSc (Clin Derm) MD FRCP José C. García-Borrón PhD FRCP(Ed) FRCP(Glasg) FHKCP FHKAM (Med) Professor of Biochemistry and Molecular Biology, School of Specialist in Dermatology and Honorary Clinical Associate Medicine, University of Murcia, Murcia, Spain Professor, University of Hong Kong and Chinese University of Hong Kong, Hong Kong Rosemary J. Geary MD President, East Valley Dermatology Center, Chandler, AZ, USA Peter Man Hon Chan MD Clinical Research Fellow, University of Minnesota, Eagan, MN, Vanessa C. Gray-Schopfer PhD USA Signal Transduction Team, Cellular and Molecular Biology Section, Institute of Cancer Research, London, UK Peggy L. Chern MD Department of Dermatology, University of Iowa Hospitals and Pearl E. Grimes MD Clinics, Iowa City, IA, USA Vitiligo and Pigmentation Institute of Southern California, David Geffen School of Medicine, University of California at Los Angeles, Mary K. Cullen MD Los Angeles, CA, USA Department of Cell Biology and Physiology, Clinical Instructor, Washington University School of Medicine, St Louis, MO, USA Vlada Groysman MD Section of Dermatology, University of Arizona Health Sciences Beth A. Drolet MD Center, Tucson, AZ, USA Department of Dermatology, The Medical College of Wisconsin, Milwaukee, WI, USA Ruth Halaban PhD Senior Research Scientist, Department of Dermatology, Yale Carol A. Erickson PhD University School of Medicine, New Haven, CT, USA Professor of Molecular Cellular Biology, University of California at Davis, Davis, CA, USA Rebat M. Halder MD Professor and Chairman, Department of Dermatology, Howard Nancy B. Esterly MD University, Washington, DC, USA Professor of Dermatology and Pediatrics, Department of Dermatology, The Medical College of Wisconsin, Milwaukee, MD PhD WI, USA Seung-Kyung Hann Director, Korea Institute of Vitiligo Research, Yongsan-Gu, Seoul, Korea Dong Fang MD PhD Staff Scientist, The Wistar Institute, Philadelphia, PA, USA Rudolf Happle MD Professor of Dermatology, Department of Dermatology, Philipp’s Sharon A. Foley MD University of Marburg, Marburg, Germany Department of Dermatology, Washington University School of Medicine, St Louis, MO, USA Vincent J. Hearing PhD Chief, Pigment Cell Biology Section, Laboratory of Cell Biology, Peter S. Friedmann MD FRCP FMedSci National Institutes of Health, Bethesda, MD, USA Professor of Dermatology and Head of Division, Dermatopharmacology Unit, Southampton General Hospital, Southampton, UK Meenhard Herlyn DVM, DSc Professor and Chair of Program of Molecular and Cellular Oncogenesis, The Wistar Institute, Philadelphia, PA, USA Charles S. Fulk MD FACP Department of Pathology, Vanderbilt University, Nashville, TN, USA Stan P. Hill MD Denver, CO, USA Anne-Sophie J. Gadenne MD Department of Dermatology, University of Cincinnati College of Lori M. Hobbs MD Medicine, Cincinnati, OH, USA Assistant Professor, Department of Dermatology, Howard University, Washington, DC, USA William A. Gahl MD PhD Clinical Director, National Human Genome Research Institute, Thomas J. Holstein PhD National Institutes of Health, Bethesda, MD, USA Professor Emeritus, Roger Williams University, Bristol, RI, USA

Sheila S. Galbraith MD Yoshiaki Hori MD PhD (deceased) Assistant Professor of Dermatology, Dermatology Department, Formerly Honorary Professor, Department of Dermatology, Kyushu Medical College of Wisconsin, Milwaukee, WI, USA University, Fukuoka, Japan

xi