S e v e n t h E d iti on A T A e x t a n d t l a s with Correlated Cell and Molecular Biology I . A 1 * ' t « S s \ « s 1 , v : V - / 1 , i f ' % • . •• • • • , • % fc# . . • I ■ hW ■ v ? I V '1 .»«• ■&' . 1* -' • :* • - f e t J • • > * /M M m tb S P w . . *i3> V v ■ v T S . l l . ' N \ • v V v W « * * www.medilibros.com W o ja e ch P a w lin a •fihWolters Kluwer H is t o l o g y A T A ext and tla s With Correlated Cell and Molecular Biology Seventh Edition W o jc ie c h P a w lin a Discussing histology education in his eosin-colored tie H i s t o l o g y A T A ext and tla s With Correlated Cell and Molecular Biology Michael H. Ross, PhD (deceased) Professor and Chairman Emeritus Department of Anatomy and Cell Biology University of Florida College of Medicine Gainesville, Florida Wojciech Pawlina, MD, FAAA Professor of Anatomy and Medical Education Fellow of the American Association of Anatomists Chair, Department of Anatomy Department of Obstetrics and Gynecology Director of Procedural Skills Laboratory Mayo Clinic College of Medicine Rochester, Minnesota Seventh Edition iters Kluwer Philadelphia • Baltimore • New York • London Buenos Aires • Hong Kong • Sydney • Tokyo Acquisitions Editor: Crystal Taylor Product Development Editor: Greg Nicholl Editorial Assistant: Joshua Haffher Production Project Manager: David Orzechowski Design Coordinator: Joan Wendt Illustration Coordinator: Jennifer Clements Marketing Manager: Joy Fisher Williams Prepress Vendor: Absolute Service, Inc. 7th edition Copyright © 2016 Wolters Kluwer Health Copyright © 2011, 2006, 2003 Lippincott Williams & Wilkins. Copyright © 1995, 1989 Williams & Wilkins. Copyright © 1985 Harper & Row, Publisher, J. B. Lippincott Company. All rights reserved. This book is protected by copyright. No part of this book may be reproduced or transmitted in any form or by any means, including as photocopies or scanned-in or other electronic copies, or utilized by any information storage and retrieval system without written permission from the copyright owner, except for brief quotations embodied in critical articles and reviews. Materials appearing in this book prepared by individuals as part of their official duties as U.S. government employees are not covered by the above-mentioned copyright. To request permission, please contact Wolters Kluwer Health at Two Commerce Square, 2001 Market Street, Philadelphia, PA 19103, via email at [email protected], or via our website at lww.com (products and services). 9 8 7 6 5 4 3 2 1 Printed in China Library of Congress Cataloging-in-Publication Data Ross, Michael H., author. Histology : a text and atlas : with correlated cell and molecular biology / Michael H. Ross, Wojciech Pawlina.—Seventh edition, p. ; cm. Includes index. ISBN 978-1-4511-8742-7 I. Pawlina, Wojciech, author. II. Title. [DNLM: 1. Histology—Atlases. QS 517] QM551 611’.018—dc23 2014032437 This work is provided “as is,” and the publisher disclaims any and all warranties, expressed or implied, including any warranties as to accuracy, comprehensiveness, or currency of the content of this work. This work is no substitute for individual patient assessment based on healthcare professionals’ examination of each patient and consideration of, among other things, age, weight, gender, current or prior medical conditions, medication history, laboratory data, and other factors unique to the patient. The publisher does not provide medical advice or guidance, and this work is merely a reference tool. Healthcare professionals, and not the publisher, are solely responsible for the use of this work including all medical judgments and for any resulting diagnosis and treatments. Given continuous, rapid advances in medical science and health information, independent professional verification of medi­ cal diagnoses, indications, appropriate pharmaceutical selections and dosages, and treatment options should be made and healthcare professionals should consult a variety of sources. When prescribing medication, healthcare professionals are advised to consult the product information sheet (the manufacturer’s package insert) accompanying each drug to verify, among other things, conditions of use, warnings, and side effects and identify any changes in dosage schedule or contradictions, particularly if the medication to be administered is new, infrequently used, or has a narrow therapeutic range. To the maximum extent per­ mitted under applicable law, no responsibility is assumed by the publisher for any injury and/or damage to persons or property, as a matter of products liability, negligence law or otherwise, or from any reference to or use by any person of this work. LWW.com This edition is dedicated to Teresa Pawlina, my wife, colleague, and best friend whose love, patience, and endurance created a safe haven for working on this textbook and to my children Conrad Pawlina and Stephanie Pawlina Fixell and her husband Ryan Fixell whose stimulation and excitement are always contagious. Preface This seventh edition of Histology: A Text and Atlas with Reader-friendly innovations have been implemented. Correlated Cell and Molecular Biology continues its tradition Similar to the previous edition of this book, the aim is to of introducing health science students to histology correlated provide more ready access to important concepts and essen­ with cell and molecular biology. As in previous editions, this tial information. Changes introduced in the sixth edition, book is a combination “text-atlas” in that the standard text­ such as bolded key terms, clinical information in blue text, book descriptions of histologic principles are supplemented and a fresh design for clinical correlation folders, were all by an array of schematics, tissue and cell images, and clini­ enthusiastically approved by the new generation of textbook cal photographs. In addition, the separate atlas sections now users and have been maintained in this edition. Important conclude each chapter to provide large-format, labeled atlas concepts have been revised and are listed as sentence head­ plates accompanied by legends that highlight and summarize ings. Dominant features of cells, tissues, and organs have the elements of microscopic anatomy. Histology: A Text and been summarized into short phrases and formatted into Atlas is, therefore, “two books in one.” bulleted lists clearly identifiable in the body of the text by The following significant modifications have been made oversized, colored bullets. Essential terms within each spe­ to this edition: cific section are introduced within the text in eye-catching, oversized, bold, red font. Text containing clinical informa­ “Histology 101 ” sections have been added at the end of tion and the latest research findings is presented in blue, with each chapter. These new sections contain essential informa­ terminology pertaining to diseases, conditions, symptoms, or tion for a quick review of the material listed in a bullet-point causative mechanisms in oversized bolded blue. Each clinical format and are perfect for students who find themselves on the folder contains updated clinical text with more illustrations eve of quizzes or examinations. These reader-friendly sections and drawings easily found within each chapter and visually are designed for fast information retrieval with concepts and appealing to keep readers turning page after page. facts listed in separate boxes. More features have been added. In understanding that All figures in this book have been carefully revised and students are pressed for time and require stimulation when updated. Many schematics and flowcharts have additionally reading several hundred pages of text, we continue to enhance been redrawn. More than one-third of all figures have been this textbook with pedagogic features, including: replaced by new drawings designed to show the latest inter­ pretation of molecular, cellular, and tissue concepts based on • “Histology 101” sections at the end of each chapter recent discoveries in molecular research. All drawings main­ • Summary tables including a review table on the character­ tain a uniform style throughout the chapters with a palette of istics of lymphatic organs eye-pleasing colors. Several conceptual drawings have been • More Clinical Correlation and Functional Considerations aligned side by side with photomicrographs, a feature car­ Folders, which contain clinical information related to the ried over from the sixth edition that was widely agreeable to symptoms, photomicrographs of diseased tissues or or­ reviewers, students, and faculty members. gans, short histopathologic descriptions, and treatment of Cellular and molecular biology content has been specific diseases updated. Text material introduced in the sixth edition has • Updated and relabeled atlas plates been updated to include the latest advancements in cellu­ • New figures, illustrations, and high-resolution digital pho­ lar and molecular biology, stem cell biology, cellular mark­ tomicrographs, more than one-third of which have been ers, and cell signaling. The seventh edition focuses on target redrawn for greater clarity and conceptual focus concepts to help students with overall comprehension of the • A bright, energetic new text design that sets off the new subject matter. To accommodate reviewers’ suggestions, the illustrations and photos and makes navigation of the text seventh edition integrates new information in cell biology even easier than before with clinical correlates, which readers will see as new clinical information items in blue text and clinical folders. For ex­ As in the last six editions, all changes have been made with ample, within the adipose tissue discussion, the reader might students in mind. We strive for clarity and concision to aid stu­ also discover a cell biology topic regarding white-to-brown dent comprehension of the subject matter, familiarity with the fat transdifferentiation. Also added is a basic discussion on latest information, and application of newfound knowledge. virtual microscopy, a new approach used in the majority of U.S. histology courses. Wojciech Pawlina Acknowledgments First and foremost, I wish to thank the creator of this book, Dr. Michael H. Ross, my mentor, colleague, and dear friend for his confidence in my ability to carry on with this project, so the future generations of students studying histology would benefit from his visionary idea of integrating text and atlas into a single book. While preparing this seventh edition, I have very much missed him, frequently recalling our meetings and discussions. He will forever be present in my heart and thoughts. Changes to the seventh edition arise largely from comments and suggestions by students who have taken the time and effort to send me e-mails of what they like about the book and, more importantly, how the book might be improved to help them better learn histology. I have also received thoughtful comments from my first-year histology students who always have an eye for improvement. I am grateful to them for the keen sense by which they sharpen this work. Many of my colleagues who teach histology and cell biology courses all over the world have, likewise, been helpful in creating this new edition. Many have suggested a stronger emphasis on clinical relevance, which I strive to continu­ ally engage as new research makes itself known. Others have provided new photomicrographs, access to their virtual slide collections or new tables, or have pointed out where existing diagrams and figures need to be redrawn. Specifically, I owe my thanks to the following reviewers, who have spent time to provide me with constructive feedback in planning this seventh edition. Baris Baykal, MD Rita Colella, PhD Giilhane Military Medical Academy University of Louisville School of Medicine Ankara, Turkey Louisville, Kentucky Irwin Beitch, PhD Iris M. Cook, PhD Quinnipiac University State University of New York Westchester Community College Hamden, Connecticut Valhalla, New York Paul B. Bell, Jr., PhD Andrea Deyrup, MD, PhD University of Oklahoma University of South Carolina School of Medicine Norman, Oklahoma Greenville, South Carolina Jalaluddin Bin Mohamed, MBBS, PhD Tamira Elul, PhD Touro University College of Osteopathic Medicine National Defence University of Malaysia Vallejo, California Kuala Lumpur, Malaysia Bruce E. Felgenhauer, PhD David E. Birk, PhD University of Louisiana at Lafayette University of South Florida, College of Medicine Lafayette, Louisiana Tampa, Florida G. Ian Gallicano, PhD Christy Bridges, PhD Georgetown University School of Medicine Mercer University School of Medicine Washington, DC Macon, Georgia Joaquin J. Garcia, MD Craig A. Canby, PhD Mayo Clinic College of Medicine Des Moines University Rochester, Minnesota Des Moines, Iowa Ferdinand Gomez, MS Stephen W. Carmichael, PhD Florida International University, Herbert Wertheim College Mayo Clinic College of Medicine of Medicine Rochester, Minnesota Miami, Florida Pike See Cheah, PhD Amos Gona, PhD Universiti Putra Malaysia University of Medicine & Dentistry of New Jersey Serdang, Selangor, Malaysia Newark, New Jersey John Clancy, Jr., PhD Ervin M. Gore, PhD Loyola University Medical Center Middle Tennessee State University Maywood, Illinois Murfreesboro, Tennessee vii Joseph P. Grande, MD, PhD Beverley Kramer, PhD Mayo Clinic College of Medicine University of the Witwatersrand Rochester, Minnesota Johannesburg, South Africa Joseph A. Grasso, PhD Craig Kuehn, PhD University of Connecticut Health Center Western University of Health Sciences Farmington, Connecticut Pomona, California Brian H. Hallas, PhD Nirusha Lachman, PhD New York Institute of Technology Mayo Clinic College of Medicine Old Westbury, New York Rochester, Minnesota Arthur R. Hand, DDS Priti S. Lacy, PhD University of Connecticut School of Dental Medicine Des Moines University, College of Osteopathic Medicine Farmington, Connecticut Des Moines, Iowa Charlene Hoegler, PhD H. Wayne Lambert, PhD Pace University West Virginia University Pleasantville, New York Morgantown, West Virginia Michael N. Horst, PhD Gavin R. Lawson, PhD Mercer University School of Medicine Western University of Health Sciences Macon, Georgia Bridgewater, Virginia Christopher Horst Lillig, PhD Susan LeDoux, PhD Ernst-Moritz Arndt University of Greifswald University of South Alabama Greifswald, Germany Mobile, Alabama Jim Hutson, PhD Karen Leong, MD Texas Tech University Drexel University College of Medicine Lubbock, Texas Philadelphia, Pennsylvania John-Olov Jansson, MD, PhD Kenneth M. Lerea, PhD University of Gothenburg New York Medical College Gothenburg, Sweden Valhalla, New York Cynthia J. M. Kane, PhD A. Malia Lewis, PhD University of Arkansas for Medical Sciences Loma Linda University Little Rock, Arkansas Loma Linda, California G. M. Kibria, MD Frank Liuzzi, PhD National Defence University of Malaysia Lake Erie College of Osteopathic Medicine Kuala Lumpur, Malaysia Bradenton, Florida Thomas S. King, PhD Donald J. Lowrie, Jr., PhD University of Texas Health Science Center at University of Cincinnati College of Medicine San Antonio Cincinnati, Ohio San Antonio, Texas Andrew T. Mariassy, PhD Penprapa S. Klinkhachorn, PhD Nova Southeastern University College of West Virginia University Medical Sciences Morgantown, West Virginia Fort Lauderdale, Florida Bruce M. Koeppen, MD, PhD Rajar am-Gilkes Mathangi, MBBS, MSc University of Connecticut Health Center St. George s University School of Medicine Farmington, Connecticut True Blue, Grenada, West Indies Andrew Koob, PhD Geoffrey W. McAuliffe, PhD University of Wisconsin River Falls Robert Wood Johnson Medical School River Falls, Wisconsin Piscataway, New Jersey