H A UMAN NATOMY Seventh Edition Frederic H. Martini, Ph.D. University of Hawaii at Manoa Michael J. Timmons, M.S. Moraine Valley Community College Robert B. Tallitsch, Ph.D. Augustana College with William C. Ober, M.D. Art Coordinator and Illustrator Claire W. Garrison, R.N. Illustrator Kathleen Welch, M.D. Clinical Consultant Ralph T. Hutchings Biomedical Photographer Executive Editor:Leslie Berriman Associate Editor:Katie Seibel Editorial Development Manager:Barbara Yien Editorial Assistant:Nicole McFadden Senior Managing Editor:Deborah Cogan Production Project Manager:Caroline Ayres Director of Media Development:Lauren Fogel Media Producer:Aimee Pavy Production Management and Composition:S4Carlisle Publishing Services, Inc. Copyeditor:Michael Rossa Art Coordinator:Holly Smith Design Manager:Marilyn Perry Interior Designer:Gibson Design Associates Cover Designer:Yvo Riezebos Photo Researcher:Maureen Spuhler Senior Manufacturing Buyer:Stacey Weinberger Marketing Manager:Derek Perrigo Cover Illustration Credit: Bryan Christie Credits and acknowledgments borrowed from other sources and reproduced, with permission, in this textbook appear on the appropriate page within the text or on page845. Copyright © 2012, 2009, 2006 by Frederic H. Martini, Inc., Michael J. Timmons, and Robert B. Tallitsch. Published by Pearson Education, Inc., publishing as Pearson Benjamin Cummings. All rights reserved. Manufactured in the United States of America. This publication is protected by Copyright and permission should be obtained from the publisher prior to any prohibited reproduction, storage in a retrieval system, or transmission in any form or by any means, electronic, mechanical, photocopying, recording, or likewise. To obtain permission(s) to use material from this work, please submit a written request to Pearson Education, Inc., Permissions Department, 1900 E. Lake Ave., Glenview, IL 60025. For information regarding permissions, call (847) 486-2635. Many of the designations used by manufacturers and sellers to distinguish their products are claimed as trademarks. Where those designations appear in this book, and the publisher was aware of a trademark claim, the designations have been printed in initial caps or all caps. Mastering A&P™, Practice Anatomy Lab™ (PAL™), and A&P Flix™ are trademarks, in the U.S. and/or other countries, of Pearson Education, Inc. or its afffiliates. Library of Congress Cataloging-in-Publication Data Martini, Frederic. Human anatomy/Frederic H. Martini, Michael J. Timmons, Robert B. Tallitsch; with William C. Ober, art coordinator and illustrator; Claire W. Garrison, illustrator; Kathleen Welch, clinical consultant; Ralph T. Hutchings, biomedical photographer.—7th ed. p. ; cm. Includes bibliographical references and index. ISBN-13: 978-0-321-68815-6 (student ed.) ISBN-10: 0-321-68815-5 (student ed.) ISBN-13: 978-0-321-73064-0 (exam copy) ISBN-10: 0-321-73064-X (exam copy) 1. Human anatomy. 2. Human anatomy—Atlases. I. Timmons, Michael J. II. Tallitsch, Robert B. III. Title. [DNLM: 1. Anatomy—Atlases. QS 17 M386h 2012] QM23.2.M356 2012 612—dc22 2010022870 ISBN 10: 0-321-68815-5 (Student edition) ISBN 13: 978-0-321-68815-6 (Student edition) ISBN 10: 0-321-76626-1 (Exam copy) ISBN 13: 978-0-321-76626-7 (Exam copy) 1 2 3 4 5 6 7 8 9 10—DOW—14 13 12 11 10 Text and Illustration Team Frederic (Ric) Martini Michael J. Timmons Robert B. Tallitsch Author Author Author Dr. Martini received his Ph.D. from Cornell Michael J. Timmons received his degrees from Dr. Tallitsch received his Ph.D. in physiology University in comparative and functional Loyola University, Chicago. For more than with an anatomy minor from the University anatomy for work on the pathophysiology of three decades he has taught anatomy to nurs- of Wisconsin-Madison. Dr. Tallitsch has stress. In addition to professional publica- ing, EMT, and pre-professional students at been on the biology faculty at Augustana tions that include journal articles and con- Moraine Valley Community College. He was College (Illinois) since 1975. His teaching re- tributed chapters, technical reports, and honored with the Professor of the Year Award sponsibilities include Human Anatomy, Neu- magazine articles, he is the lead author of by MVCC and the Excellence Award from the roanatomy, Histology, and Kinesiology. He is nine undergraduate texts on anatomy or National Institute for Staff and Organizational also a member of the Asian Studies faculty at anatomy and physiology. Dr. Martini is cur- Development for his outstanding contribu- Augustana College, teaching a course in Tra- rently affiliated with the University of Hawaii tions to teaching, leadership, and student ditional Chinese Medicine. In ten out of the at Manoa and has a long-standing bond with learning. He is the recipient of the Excellence in last twelve years the graduating seniors at Au- the Shoals Marine Laboratory, a joint venture Teaching Award by the Illinois Community gustana have designated Dr. Tallitsch as one between Cornell University and the Univer- College Board of Trustees. Professor Timmons, of the “unofficial teachers of the year.” Dr. sity of New Hampshire. Dr. Martini is a Pres- a member of the American Association of Tallitsch is a member of the American Physi- ident Emeritus of the Human Anatomy and Anatomists, has authored several anatomy and ological Society, American Association of Physiology Society, and he is a member of the physiology lab manuals and dissection guides. Anatomists, American Association of Clini- American Association of Anatomists, the His areas of interest include biomedical pho- cal Anatomists, AsiaNetwork, and the Hu- American Physiological Society, the Society tography, crafting illustration programs, and man Anatomy and Physiology Society. In for Integrative and Comparative Biology, and developing instructional technology learning addition to his teaching responsibilities at the International Society of Vertebrate Mor- systems. He chaired the Midwest Regional Hu- Augustana College, Dr. Tallitsch has served phologists. man Anatomy and Physiology Conference and as a visiting faculty member at the Beijing is also a national and regional presenter at the University of Chinese Medicine and Pharma- League for Innovation Conferences on Infor- cology (Beijing, PRC), the Foreign Languages mation Technology for Colleges and Universi- Faculty at Central China Normal University ties and at Human Anatomy and Physiology (Wuhan, PRC), and in the Biology Depart- Society meetings. ment at Central China Normal University (Wuhan, PRC). iii iv Text and Illustration Team Kathleen Welch Ralph T. Hutchings Clinical Consultant Biomedical Photographer William C. Ober Dr. Welch received her M.D. from the Univer- Mr. Hutchings was associated with The Royal Art Coordinator and Illustrator sity of Washington in Seattle and did her resi- College of Surgeons of England for 20 years. dency at the University of North Carolina in An engineer by training, he has focused for Dr. William C. Ober received his undergrad- Chapel Hill. For two years she served as Direc- years on photographing the structure of the uate degree from Washington and Lee Uni- tor of Maternal and Child Health at the LBJ human body. The result has been a series of versity and his M.D. from the University of Tropical Medical Center in American Samoa color atlases, including the Color Atlas of Hu- Virginia. While in medical school, he also and subsequently was a member of the De- man Anatomy, the Color Atlas of Surface studied in the Department of Art as Applied partment of Family Practice at the Kaiser Per- Anatomy, and The Human Skeleton (all pub- to Medicine at Johns Hopkins University. Af- manente Clinic in Lahaina, Hawaii. She has lished by Mosby-Yearbook Publishing). For ter graduation, Dr. Ober completed a resi- been in private practice since 1987. Dr. Welch his anatomical portrayal of the human body, dency in Family Practice and later was on the is a Fellow of the American Academy of Fam- the International Photographers Association faculty at the University of Virginia in the ily Practice and a member of the Hawaii Med- has chosen Mr. Hutchings as the best photog- Department of Family Medicine. He is cur- ical Association and the Human Anatomy and rapher of humans in the twentieth century. He rently a Visiting Professor of Biology at Physiology Society. lives in North London, where he tries to bal- Washington and Lee University and is part of ance the demands of his photographic assign- the Core Faculty at Shoals Marine Labora- ments with his hobbies of early motor cars tory, where he teaches Biological Illustration and airplanes. every summer. The textbooks illustrated by Medical & Scientific Illustration have won numerous design and illustration awards. Claire W. Garrison Illustrator Claire W. Garrison, R.N., B.A., practiced pe- diatric and obstetric nursing before turning to medical illustration as a full-time career. She returned to school at Mary Baldwin Col- lege where she received her degree with dis- tinction in studio art. Following a five-year apprenticeship, she has worked as Dr. Ober’s partner in Medical & Scientific Illustration since 1986. She is on the Core Faculty at Shoals Marine Laboratory and co-teaches the Biological Illustration course. Preface Human Anatomy! Welcome to the Seventh Edition of THROUGH SEVEN EDITIONS,the authors and illustrators have continued to build on this text’s hallmark qualities: its distinctive atlas-style format and its unsurpassed visual presentation of anatomy and anatomical concepts. Our approach for this text has been to provide a seamless learning system with closely integrated art and text. The illustrations do more than provide occasional support for the narrative; they are partners with the text in conveying information and helping students understand structures and relationships in a way that distinguishes this human anatomy textbook from all others. New to the Seventh Edition In approaching this Seventh Edition, we paid particular attention to the most difficult topics in human anatomy and to areas identified by students and reviewers. Our primary goal was to build upon the strengths of the previous edition while addressing the changing needs of today’s students. The changes described below are intended to enhance student learning and increase student engagement. • A more visual and dynamic presentation of clinical information.Select Clinical Notes covering key clinical topics now feature new, dramatic layouts that integrate illustrations, photos, and text in a way that makes reading easy and science relevant (see pp.108–109, 127, 132–133). Clinical Cases, which appear at the end of each body system section, now include patient photos and diagnostic images (see pp.110–111, 501–502, 602–604). Every Clinical Case begins with a photo of the patient and his/her background information, making the case personal and real to the students. Diagnostic images (photos, x-rays, and MRI scans) also appear within the narrative. • Over 65 new and visually stunning histology photomicrographs.These photomicrographs appear in chapters 3, 4, 5, 13, 19–21, and 23–27. The slides prepared for these photos match the types v vi Preface of slides that beginning students will encounter in the anatomy • New “Hot Topics: What’s New in Anatomy” highlight cur- lab. rent research.These brief boxes introduce students to new • New spiral scans.Using the most up-to-date imaging technique peer-reviewed anatomical research findings that have been available, these spiral scans (see Figures8.16and 22.16) provide published within the past two years. This feature appears in students with unparallelled views of anatomical structures and chapters 2–5, 10, 13, 19, 21, and 23–28. introduce them to a new imaging technique that is increasingly • Increased focus on learning methodology.Each chapter now used in clinical settings. These spiral scan images have been pro- opens with concrete Student Learning Outcomesinstead of vided by Fovia, Inc., and by TeraRecon, Inc. learning objectives. • Improved presentation of figures.Figure legends now appear In addition, approximately 85 percent of the figures in this consistently above figures, and the detailed figure captions that edition are either new or have been revised. Some figures were describe parts within figures now appear within the figures. updated for increased visual appeal to students (see Figures 1.1, This new figure presentation style guides students through 4.1, and 4.12). In many figures, areas of detail have been revised multi-part figures and compels them to read the part captions to improve clarity. All bone photos in chapters 6 and 7 received a as they view each part of a figure. The result is easier reading new silhouette treatment that results in a cleaner, more and improved understanding of figures. contemporary look and makes bone markings easier to see. The • A reorganized and streamlined presentation of the nervous presentation of boxes and banners has been improved to better system chapters (Chapters 13–18).These chapters have been organize many figures (see Figures 9.11, 26.6, and 23.7). The reorganized to take a “bottom up” rather than a “top down” ap- overlay of illustrations on surface anatomy photos has been proach to make the nervous system easier for instructors to continued in this edition to provide students with a better present and students to understand. Specifically, the discussion understanding of where structures are located within the human of the spinal cord started in Chapter 14 (The Nervous System: body. The information derived from superficial and deep The Spinal Cord and Spinal Nerves) now continues in Chapter dissections is more easily understood as a result of a new heading 15 (The Nervous System: Sensory and Motor Tracts of the style that has been continued in many of the figures (see Figure Spinal Cord) so that sensory and motor tracts of the spinal cord 23.14b). are covered beforethe brain and cranial nerves in Chapter 16 The following section provides a detailed description of this (The Nervous System: The Brain and Cranial Nerves). Addi- edition’s chapter-by-chapter revisions. tionally, Chapter 16 also presents the brain and cranial nerve in- formation in a “bottom up” sequence, starting with the brain stem and ending with the cerebrum. Preface vii Chapter-by-Chapter Revisions Specific chapter-by-chapter revisions, with select examples, include: • New material was added, and existing material has been clarified, in the dis- cussions of the clavicle, scapula, humerus, pelvic girdle, patella, tibia, and the 1 Foundations: An Introduction to Anatomy arches of the foot. • Twelve illustrations are either new or have been significantly revised. 8 The Skeletal System: Articulations • Changes were made in terminology according to the Terminologia Anatomica (TA). • Seven illustrations are either new or have been significantly revised. • New material was added and existing material clarified for better student 2 Foundations: The Cell comprehension. • Fifteen illustrations are either new or have been significantly revised. 9 The Muscular System: Skeletal Muscle Tissue and • Changes were made in terminology according to the TA and Terminologia Muscle Organization Histologica (TH). • The presentation order of some material was rearranged in order to facilitate • Eight illustrations are either new or have been significantly revised. student learning. • Considerable material within the chapter was revised to better facilitate stu- dent comprehension and learning. 3 Foundations: Tissues and Early Embryology 10 The Muscular System: Axial Musculature • Nineteen illustrations are either new or have been significantly revised. • Seventeen new photomicrographs were added. • Five illustrations are either new or have been significantly revised. • Changes were made in terminology according to the TAand TH. • Two new photomicrographs were added. • The presentation order of some material was rearranged in order to facilitate • The sections entitled “Muscles of the Vertebral Column” and “Muscles of the student learning. Perineum and the Pelvic Diaphragm” have been updated and clarified. • New material was added to update the chapter according to current histolog- ical research. 11 The Muscular System: Appendicular Musculature 4 The Integumentary System • Nine illustrations are either new or have been significantly revised. • A new section entitled “Factors Affecting Appendicular Muscle Function” • Fourteen illustrations are either new or have been significantly revised. was added to this chapter in the Sixth Edition and has been revised for this • Four new photomicrographs were added. Seventh Edition. This section helps students work through the process of • Changes were made in terminology according to the TAand TH. understandingthe actions of skeletal muscles at a joint. This section also ex- • New material was added to the discussion of the epidermis, and the existing plains the concept of the action line of a muscle, and how students, once they material was revised for easier comprehension. have determined the action line, may apply three simple rules in order to de- termine the action of a muscle at that joint. 5 The Skeletal System: Osseous Tissue and Skeletal Structure 12 Surface Anatomy and Cross-Sectional Anatomy • Eleven illustrations are either new or have been significantly revised. • Nine illustrations are either new or have been significantly revised. • Two new photomicrographs were added. • New material was added to the discussion of bone remodeling and repair, and 13 The Nervous System: Neural Tissue the existing material was revised for easier reading and comprehension. • New material was added to the discussion of the cells of bone to match cur- • Five illustrations are either new or have been significantly revised. rent histological terminology and research. • Two new photomicrographs were added. • The sections entitled “Neuroglia of the CNS” and “Synaptic Communication” 6 The Skeletal System: Axial Division were updated in order to match current research findings in the field. • Twenty-three illustrations are either new or have been significantly revised. 14 The Nervous System: The Spinal Cord and Spinal Nerves • New material was added to the discussion of the bones of the cranium to match current anatomical terminology and research. • Seven illustrations are either new or have been significantly revised. • New material was added, and existing material has been clarified, in the dis- • The discussion of the meninges of the spinal cord was expanded. cussions of the vertebral regions. • The discussion of the sectional anatomy of the spinal cord was expanded, with particular emphasis on the revision of the section on “Organization of 7 The Skeletal System: Appendicular Division the Gray Matter.” • The section on “Spinal Nerves” has been rewritten in order to facilitate stu- • Twenty-one illustrations are either new or have been significantly dent learning and comprehension. revised. viii Preface • The sections on “The Brachial Plexus” and “The Lumbar and Sacral Plexuses” • All sections of this chapter were revised, either partially or totally, to make were rewritten to make them easier to understand. them easier to understand. 15 The Nervous System: Sensory and Motor Tracts of the 23 The Lymphoid System Spinal Cord • Eight illustrations are either new or have been significantly revised. • Two new illustrations have been included and eight others have been signifi- • Four new photomicrographs were added. cantly revised. • All sections of this chapter were updated in order to match current research • All sections of this chapter were revised, either partially or totally, to make findings in the field. them easier to understand. • All sections of this chapter were revised, either partially or totally, to make • At the request of reviewers and instructors, the section dealing with Higher- them easier to understand. Order Functions has been deleted. 24 The Respiratory System 16 The Nervous System: The Brain and Cranial Nerves • Seven illustrations are either new or have been significantly revised. • Ten illustrations have been significantly revised. • Two new photomicrographs were added. • Revisions were made to reflect the current histological information on the respiratory system. 17 The Nervous System: Autonomic Division • All sections of this chapter were revised, either partially or totally, to make • Seven illustrations are either new or have been significantly revised. them easier to understand. • All sections of this chapter were revised, either partially or totally, to make them easier to understand. 25 The Digestive System • Thirteen illustrations are either new or have been significantly revised. 18 The Nervous System: General and Special Senses • Thirteen new photomicrographs were added. • Seven illustrations are either new or have been significantly revised. • Revisions were made to reflect the current histological information on the • All sections of this chapter were revised, either partially or totally, to make various organs of the digestive system. them easier to understand. • All sections of this chapter were revised, either partially or totally, to make them easier to understand. 19 The Endocrine System 26 The Urinary System • Five illustrations are either new or have been significantly revised. • Five new photomicrographs were added. • Seven illustrations are either new or have been significantly revised. • All sections of this chapter were revised, either partially or totally, to make • Six new photomicrographs were added. them easier to understand. • Revisions were made to reflect the current histological information on the various organs of the urinary system. • All sections of this chapter were revised, either partially or totally, to make 20 The Cardiovascular System: Blood them easier to understand. • Six illustrations are either new or have been significantly revised. • Five new photomicrographs were added. 27 The Reproductive System • All sections of this chapter were updated in order to match current research findings in the field. • Seven illustrations are either new or have been significantly revised. • Six new photomicrographs were added. • Revisions were made to reflect the current histological information on the 21 The Cardiovascular System: The Heart various organs of the male and female reproductive systems. • Eight illustrations are either new or have been significantly revised. • All sections of this chapter were revised, either partially or totally, to make • One new photomicrograph was added. them easier to understand. • The sections on “The Intercalated Discs” and “Coronary Blood Vessels” were rewritten in order to reflect new research findings in the field and to make 28 The Reproductive System: Embryology and Human them easier to understand. Development 22 The Cardiovascular System: Vessels and Circulation • All of the Embryology Summaries have been revised. • Eleven illustrations are either new or have been significantly revised. • All sections of this chapter were updated in order to match current research findings in the field. Acknowledgments The creative talents brought to this project by our artist team, William Ober, We are also indebted to the Pearson Benjamin Cummings staff, whose efforts M.D., Claire Garrison, R.N., and Anita Impagliazzo, M.F.A., are inspiring and were vital to the creation of this edition. A special note of thanks and apprecia- valuable beyond expression. Bill, Claire, and Anita worked intimately and tire- tion goes to the editorial staff at Benjamin Cummings, especially Leslie Berri- lessly with us, imparting a unity of vision to the book while making each illus- man, Executive Editor, for her dedication to the success of this project, and tration clear and beautiful. Their superb art program is greatly enhanced by the Katie Seibel, Associate Editor, for her management of the text and its supple- incomparable bone and cadaver photographs of Ralph T. Hutchings, formerly ments. Thanks also to Barbara Yien, Editorial Development Manager, and of The Royal College of Surgeons of England. In addition, Dr. Pietro Motta, Pro- Nicole McFadden, Editorial Assistant. We express thanks to Aimee Pavy, Media fessor of Anatomy, University of Roma, La Sapienza, provided several superb Producer, and Sarah Young-Dualan, Senior Media Producer, for their work on SEM images for use in the text. We also gratefully acknowledge Shay Kilby, Ken the media programs that support Human Anatomy,especially Mastering A & P™ Fineman, and Steve Sandy of Fovia, Inc., and Donna Wefers and Cormac Dono- and Practice Anatomy Lab™ (PAL™). Thanks also to Caroline Ayres, Production van of TeraRecon, Inc., for creating and providing the 3-D spiral scans that ap- Supervisor, for her steady hand managing this complex text; and Debbie Cogan, pear in this edition. Norine Strang, Holly Smith, Maureen Spuhler, and Donna Kalal for their roles We are deeply indebted to Jim Gibson of Graphic Design Associates for his in the production of the text. wonderful work and suggestions in the design aspect of the Seventh Edition of We are very grateful to Paul Corey, President, and Frank Ruggirello, Edito- Human Anatomy. Jim provided new insight into the design concept, and most rial Director, for their continued enthusiasm and support of this project. We ap- of the design changes and innovations in this edition of Human Anatomyreflect preciate the contributions of Derek Perrigo, Marketing Manager, who keeps his Jim’s expertise. finger on the pulse of the market and helps us meet the needs of our customers, We would like to acknowledge the many users and reviewers whose advice, and the remarkable and tireless Pearson Science sales reps. comments, and collective wisdom helped shape this text into its final form. We are also grateful that the contributions of all of the aforementioned peo- Their passion for the subject, their concern for accuracy and method of presen- ple have led to this text receiving the following awards: The Association of Med- tation, and their experience with students of widely varying abilities and back- ical Illustrators Award, The Text and Academic Authors Award, the New York grounds have made the revision process interesting and educating. International Book Fair Award, the 35th Annual Bookbuilders West Award, and the 2010 Text and Academic Authors Association “Texty” Textbook Excellence Reviewers Award. We would also like to thank Steven Bassett of Southeast Community Col- Lori Anderson,Ridgewater College lege; Kelly Johnson of University of Kansas; Jason LaPres of North Harris Col- Tamatha R. Barbeau,Francis Marion University lege; Agnes Yard of University of Indianopolis; and Michael Yard of Indiana Steven Bassett,Southeast Community College University-Purdue University at Indianapolis for their work on the media and Martha L. Dixon,Diablo Valley College print supplements for this edition. Cynthia A. Herbrandson,Kellogg Community College Finally, we would like to thank our families for their love and support dur- Judy Jiang,Triton College ing the revision process. We could not have accomplished this without the help Kelly Johnson,University of Kansas of our wives—Kitty, Judy, and Mary—and the patience of our children—P.K., Michael G. Koot,Michigan State University Molly, Kelly, Patrick, Katie, Ryan, Molly, and Steven. George H. Lauster,Pulaski Technical College No three people could expect to produce a flawless textbook of this scope Robert G. MacBride,Delaware State University and complexity. Any errors or oversights are strictly our own rather than those Les MacKenzie,Queen’s University of the reviewers, artists, or editors. In an effort to improve future editions, we Christopher McNair,Hardin-Simmons University ask that readers with pertinent information, suggestions, or comments con- Qian F. Moss,Des Moines Area Community College cerning the organization or content of this textbook send their remarks to Tim R. Mullican,Dakota Wesleyan University Robert Tallitsch directly, by the e-mail address below, or care of Publisher, Ap- John Steiner,College of Alameda plied Sciences, Pearson Benjamin Cummings, 1301 Sansome Street, San Fran- Lucia J. Tranel,Saint Louis College of Pharmacy cisco, CA 94111. Maureen Tubbiola,Saint Cloud State University Jacqueline Van Hoomissen,University of Portland Frederic H. Martini,Haiku, HI Michael Yard,Indiana University-Purdue University at Indianapolis Michael J. Timmons,Orland Park, IL Scott Zimmerman,Missouri State University Robert B. Tallitsch,Rock Island, IL John M. Zook,Ohio University ([email protected]) ix