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(BQ) Part 1 book The developing human clinically oriented embryology presents the following contents: Introduction to human development, first week of human development, first week of human development, third week of human development, fourth to eighth weeks of human development,...
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Access to the eBook is limited to the first individual who redeems the PIN, located on the inside cover of this book, at studentconsult.inkling.com and may not be transferred to another party by resale, lending or other means THE DEVELOPING HUMAN CLINICALLY ORIENTED EMBRYOLOGY KEITH L MOORE Recipient of the inaugural Henry Gray/Elsevier Distinguished Educator Award in 2007—the American Association of Anatomists’ highest award for excellence in human anatomy education at the medical/dental, graduate, and undergraduate levels of teaching; the Honored Member Award of the American Association of Clinical Anatomists (1994) for significant contributions to the field of clinically relevant anatomy; and the J.C.B Grant Award of the Canadian Association of Anatomists (1984) “in recognition of meritorious service and outstanding scholarly accomplishments in the field of anatomical sciences.” In 2008 Professor Moore was inducted as a Fellow of the American Association of Anatomists The rank of Fellow honors distinguished AAA members who have demonstrated excellence in science and in their overall contributions to the medical sciences In 2012 Dr Moore received an Honorary Doctor of Science degree from The Ohio State University; The Queen Elizabeth II Diamond Jubilee Medal honoring significant contributions and achievements by Canadians; and the Benton Adkins Jr Distinguished Service Award for an outstanding record of service to the American Association of Clinical Anatomists T.V.N (VID) PERSAUD Recipient of the Henry Gray/Elsevier Distinguished Educator Award in 2010—the American Association of Anatomists’ highest award for excellence in human anatomy education at the medical/dental, graduate, and undergraduate levels of teaching; the Honored Member Award of the American Association of Clinical Anatomists (2008) for significant contributions to the field of clinically relevant anatomy; and the J.C.B Grant Award of the Canadian Association of Anatomists (1991) “in recognition of meritorious service and outstanding scholarly accomplishments in the field of anatomical sciences.” In 2010 Professor Persaud was inducted as a Fellow of the American Association of Anatomists The rank of Fellow honors distinguished AAA members who have demonstrated excellence in science and in their overall contributions to the medical sciences In 2003 Dr Persaud was a recipient of the Queen Elizabeth II Golden Jubilee Medal, presented by the Government of Canada for “significant contribution to the nation, the community, and fellow Canadians.” MARK G TORCHIA Recipient of the Norman and Marion Bright Memorial Medal and Award and the Silver Medal of the Chemical Institute of Canada in 1990 for outstanding contributions In 1993 he was awarded the TIMEC Medical Device Champion Award In 2008 and in 2014 Dr Torchia was a nominee for the Manning Innovation Awards, for innovation talent Dr Torchia’s most cherished award has been the Award for Teaching Excellence in 2011 from the Faculty of Medicine, University of Manitoba, and being asked to address the graduating class of 2014 THE DEVELOPING HUMAN CLINICALLY ORIENTED EMBRYOLOGY 10th Edition Keith L Moore, BA, MSc, PhD, DSc, FIAC, FRSM, FAAA Professor Emeritus, Division of Anatomy, Department of Surgery Former Professor and Chair, Department of Anatomy and Associate Dean for Basic Medical Sciences Faculty of Medicine, University of Toronto, Toronto, Ontario, Canada Former Professor and Head of Anatomy, Faculty of Medicine, University of Manitoba, Winnipeg, Manitoba, Canada T.V.N (Vid) Persaud, MD, PhD, DSc, FRCPath (Lond.), FAAA Professor Emeritus and Former Head, Department of Human Anatomy and Cell Science Professor of Pediatrics and Child Health Associate Professor of Obstetrics, Gynecology, and Reproductive Sciences, Faculty of Medicine, University of Manitoba, Winnipeg, Manitoba, Canada Professor of Anatomy, St George’s University, Grenada, West Indies Mark G Torchia, MSc, PhD Associate Professor and Director of Development, Department of Surgery Associate Professor, Department of Human Anatomy and Cell Sciences Director, Centre for the Advancement of Teaching and Learning, University of Manitoba, Winnipeg, Manitoba, Canada 1600 John F Kennedy Blvd Ste 1800 Philadelphia, PA 19103-2899 THE DEVELOPING HUMAN, TENTH EDITION INTERNATIONAL EDITION Copyright © 2016 by Elsevier, Inc All rights reserved ISBN: 978-0-323-31338-4 ISBN: 978-0-323-31347-6 No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, recording, or any information storage and retrieval system, without permission in writing from the publisher Details on how to seek permission, further information about the Publisher’s permissions policies and our arrangements with organizations such as the Copyright Clearance Center and the Copyright Licensing Agency, can be found at our website: www.elsevier.com/permissions This book and the individual contributions contained in it are protected under copyright by the Publisher (other than as may be noted herein) Notices Knowledge and best practice in this field are constantly changing As new research and experience broaden our understanding, changes in research methods, professional practices, or medical treatment may become necessary Practitioners and researchers must always rely on their own experience and knowledge in evaluating and using any information, methods, compounds, or experiments described herein In using such information or methods they should be mindful of their own safety and the safety of others, including parties for whom they have a professional responsibility With respect to any drug or pharmaceutical products identified, readers are advised to check the most current information provided (i) on procedures featured or (ii) by the manufacturer of each product to be administered, to verify the recommended dose or formula, the method and duration of administration, and contraindications It is the responsibility of practitioners, relying on their own experience and knowledge of their patients, to make diagnoses, to determine dosages and the best treatment for each individual patient, and to take all appropriate safety precautions To the fullest extent of the law, neither the Publisher nor the authors, contributors, or editors, assume any liability for any injury and/or damage to persons or property as a matter of products liability, negligence or otherwise, or from any use or operation of any methods, products, instructions, or ideas contained in the material herein Previous editions copyrighted 2013, 2008, 2003, 1998, 1993, 1988, 1982, 1977, and 1973 Library of Congress Cataloging-in-Publication Data Moore, Keith L., author The developing human : clinically oriented embryology / Keith L Moore, T.V.N (Vid) Persaud, Mark G Torchia.—10th edition p ; cm Includes bibliographical references and index ISBN 978-0-323-31338-4 (pbk : alk paper)—ISBN 978-0-323-31347-6 (international edition : alk paper) I Persaud, T V N., author. II Torchia, Mark G., author. III Title [DNLM: Embryology QS 604] QM601 612.6′4018—dc23 2015001490 Content Strategist: Meghan Ziegler Senior Content Development Specialist: Jennifer Ehlers Publishing Services Manager: Patricia Tannian Senior Project Manager: Kristine Feeherty Design Direction: Margaret Reid The cover images show a magnetic resonance image of a 27-week-old fetus in the uterus (Courtesy Dr Deborah Levine, Beth Israel Deaconess Medical Center, Boston, Massachusetts) The photograph of the baby (Kennedy Jackson) was taken days after her birthday She is wrapped in a knitted cocoon that symbolizes the uterus Printed in the United States of America Last digit is the print number: 9 8 7 6 5 4 3 2 In Loving Memory of Marion My best friend, wife, colleague, mother of our five children and grandmother of our nine grandchildren, for her love, unconditional support, and understanding Wonderful memories keep you ever near our hearts —KLM and family For Pam and Ron I should like to thank my eldest daughter, Pam, who assumed the office duties previously carried out by her mother, Marion She has also been helpful in so many other ways (e.g., reviewing the text) I am also grateful to my son-in-law, Ron Crowe, whose technical skills have helped me utilize the new technology when I was improving this book —KLM For Gisela My lovely wife and best friend, for her endless support and patience; our three children—Indrani, Sunita, and Rainer (Ren)—and grandchildren (Brian, Amy, and Lucas) —TVNP For Barbara, Muriel, and Erik Nothing could ever mean more to me than each of you Thank you for your support and your love —MGT For Our Students and Their Teachers To our students: We hope you will enjoy reading this book, increase your understanding of human embryology, pass all of your exams, and be excited and well prepared for your careers in patient care, research, and teaching You will remember some of what you hear, much of what you read, more of what you see, and almost all of what you experience To their teachers: May this book be a helpful resource to you and your students We appreciate the numerous constructive comments we have received over the years from both students and teachers Your remarks have been invaluable to us in improving this book Contributors CONTRIBUTORS FIGURES AND IMAGES (SOURCES) David D Eisenstat, MD, MA, FRCPC Professor, Departments of Pediatrics, Medical Genetics and Oncology, Faculty of Medicine and Dentistry, University of Alberta; Director, Division of Pediatric Immunology, Hematology, Oncology, Palliative Care, and Environmental Health, Department of Pediatrics, Stollery Children’s Hospital and the University of Alberta; Inaugural Chair, Muriel and Ada Hole Kids with Cancer Society Chair in Pediatric Oncology, Edmonton, Alberta, Canada We are grateful to the following colleagues for the clinical images they have given us for this book and also for granting us permission to use figures from their published works: Jeffrey T Wigle, PhD Principal Investigator, Institute of Cardiovascular Sciences, St Boniface Hospital Research Centre; Associate Professor, Department of Biochemistry and Medical Genetics, University of Manitoba, Winnipeg, Manitoba, Canada CLINICAL REVIEWERS Albert E Chudley, MD, FRCPC, FCCMG Professor, Department of Pediatrics and Child Health; Professor, Department of Biochemistry and Medical Genetics, University of Manitoba, Winnipeg, Manitoba, Canada Steve Ahing, DDS Faculty of Dentistry, University of Manitoba, Winnipeg, Manitoba, Canada Figure 19-20F Franco Antoniazzi, MD Department of Pediatrics, University of Verona, Verona, Italy Figure 20-4 Dean Barringer and Marnie Danzinger Figure 6-7 †Volker Becker, MD Pathologisches Institut der Universität, Erlangen, Germany Figures 7-18 and 7-21 J.V Been, MD Department of Pediatrics, Maastricht University Medical Centre, Maastricht, The Netherlands Figure 10-7C Michael Narvey, MD, FRCPC, FAAP Section Head, Neonatal Medicine, Health Sciences Centre and St Boniface Hospital; Associate Professor of Pediatrics and Child Health, University of Manitoba, Winnipeg, Manitoba, Canada Beryl Benacerraf, MD Diagnostic Ultrasound Associates, P.C., Boston, Massachusetts, USA Figures 13-29A, 13-35A, and 13-37A Kunwar Bhatnagar, MD Department of Anatomical Sciences and Neurobiology, School of Medicine University of Louisville, Louisville, Kentucky, USA Figures 9-33, 9-34, and 19-10 †Deceased vii viii CONTRIBUT O R S David Bolender, MD Department of Cell Biology, Neurobiology, and Anatomy, Medical College of Wisconsin, Milwaukee, Wisconsin, USA João Carlos Fernandes Rodrigues, MD Servico de Dermatologia, Hospital de Desterro, Lisbon, Portugal Figure 19-5B Figure 14-14BC Dr Mario João Branco Ferreira Servico de Dermatologia, Hospital de Desterro, Lisbon, Portugal Frank Gaillard, MB, BS, MMed Department of Radiology, Royal Melbourne Hospital, Australia Figures 4-15 and 9-19B Figure 19-5A Albert E Chudley, MD, FRCPC, FCCMG Department of Pediatrics and Child Health, Section of Genetics and Metabolism, Children’s Hospital, University of Manitoba, Winnipeg, Manitoba, Canada Figures 4-6, 9-38, 11-19AB, 11-28A, 12-24, 12-42, 12-43, 14-11, 15-6, 16-13DE, 16-14, 16-15, 17-14, 17-33, 17-36, 18-20, 18-21, 18-23, 19-9, 20-3, 20-5, 20-6CD, 20-7, 20-8, 20-13, 20-14, 20-17, and 20-19A Blaine M Cleghorn, DMD, MSc Faculty of Dentistry, Dalhousie University, Halifax, Nova Scotia, Canada Figures 19-19 and 19-20A-E Dr M.N Golarz De Bourne St George’s University Medical School, True Blue, Grenada Figure 11-21 Heather Dean, MD, FRCPC Department of Pediatrics and Child Health, University of Manitoba, Winnipeg, Manitoba, Canada Figures 12-28 and 20-18 Gary Geddes, MD Lake Oswego, Oregon, USA Figure 14-14A Barry H Grayson, MD, and Bruno L Vendittelli, MD New York University Medical Center, Institute of Reconstructive Plastic Surgery, New York, New York, USA Figure 9-40 Christopher R Harman, MD, FRCSC, FACOG Department of Obstetrics, Gynecology, and Reproductive Sciences, Women’s Hospital and University of Maryland, Baltimore, Maryland, USA Figures 7-17 and 12-23 †Jean Hay, MSc Department of Anatomy, University of Manitoba, Winnipeg, Manitoba, Canada Figure 17-25 Blair Henderson, MD Department of Radiology, Health Sciences Centre, University of Manitoba, Winnipeg, Manitoba, Canada Figure 13-6 Marc Del Bigio, MD, PhD, FRCPC Department of Pathology (Neuropathology), University of Manitoba, Winnipeg, Manitoba, Canada Figures 17-13, 17-29 (inset), 17-30BC, 17-32B, 17-37B, 17-38, 17-40, and 17-42A David D Eisenstat, MD, MA, FRCPC Manitoba Institute of Cell Biology, Department of Human Anatomy and Cell Science, University of Manitoba, Winnipeg, Manitoba, Canada Figure 17-2 Lyndon M Hill, MD Magee-Women’s Hospital, Pittsburgh, Pennsylvania, USA Figures 11-7 and 12-14 †Klaus V Hinrichsen, MD Medizinische Fakultät, Institut für Anatomie, Ruhr-Universität Bochum, Bochum, Germany Figures 5-12A, 9-2, and 9-26 Dr Jon and Mrs Margaret Jackson Figure 6-9B Vassilios Fanos, MD Department of Pediatrics, University of Verona, Verona, Italy Figure 20-4 †Deceased C O N T R I B U TORS Evelyn Jain, MD, FCFP Breastfeeding Clinic, Calgary, Alberta, Canada Figure 9-24 John A Jane, Sr., MD David D Weaver Professor of Neurosurgery, Department of Neurological Surgery, University of Virginia Health System, Charlottesville, Virginia, USA Figure 14-12 Margaret Morris, MD, FRCSC, MEd Professor of Obstetrics, Gynaecology, and Reproductive Sciences, Women’s Hospital and University of Manitoba, Winnipeg, Manitoba, Canada Figure 12-46 Stuart C Morrison, MD Section of Pediatric Radiology, The Children’s Hospital, Cleveland Clinic, Cleveland, Ohio, USA Figures 7-13, 11-20, 17-29E, and 17-41 Robert Jordan, MD St George’s University Medical School, True Blue, Grenada Figures 6-6B and 7-25 John B Mulliken, MD Children’s Hospital Boston, Harvard Medical School, Boston, Massachusetts, USA Figure 9-42 Linda J Juretschke, MD Ronald McDonald Children’s Hospital, Loyola University Medical Center, Maywood, Illinois, USA Figure 7-31 W Jerry Oakes, MD Children’s Hospital Birmingham, Birmingham, Alabama, USA Figure 17-42B Dagmar K Kalousek, MD Department of Pathology, University of British Columbia, Children’s Hospital, Vancouver, British Columbia, Canada Figures 8-11AB, 11-14A, 12-12C, 12-16, and 20-6AB †Dwight Parkinson, MD Departments of Surgery and Human Anatomy & Cell Science, University of Manitoba, Winnipeg, Manitoba, Canada Figure 17-14 E.C Klatt, MD Department of Biomedical Sciences, Mercer University School of Medicine, Savannah, Georgia, USA Figure 7-16 Wesley Lee, MD Division of Fetal Imaging, William Beaumont Hospital, Royal Oak, Michigan, USA Figures 13-20 and 13-30A Deborah Levine, MD, FACR Departments of Radiology and Obstetric & Gynecologic Ultrasound, Beth Israel Deaconess Medical Center, Boston, Massachusetts, USA Figures 6-8, 6-15, 8-10, 9-43CD, 17-35B, and cover image (magnetic resonance image of 27-week fetus) Maulik S Patel, MD Consultant Pathologist, Surat, India Figure 4-15 Dr Susan Phillips Department of Pathology, Health Sciences Centre, Winnipeg, Manitoba, Canada Figure 18-6 Srinivasa Ramachandra, MD Figure 9-13A †Dr M Ray Department of Human Genetics, University of Manitoba, Winnipeg, Manitoba, Canada Figure 20-12B E.A (Ted) Lyons, OC, MD, FRCPC, FACR Departments of Radiology, Obstetrics & Gynecology, and Human Anatomy & Cell Science, Division of Ultrasound, Health Sciences Centre, University of Manitoba, Winnipeg, Manitoba, Canada ix Martin H Reed, MD, FRCPC Department of Radiology, University of Manitoba and Children’s Hospital, Winnipeg, Manitoba, Canada Figure 11-27 Figures 3-7, 3-9, 4-1, 4-13, 5-19, 6-1, 6-10, 6-12, 7-23, 7-26, 7-29, 11-19CD, 12-45, and 13-3 †Deceased ... 9-38, 11 -19 AB, 11 -28A, 12 -24, 12 -42, 12 -43, 14 -11 , 15 -6, 16 -13 DE, 16 -14 , 16 -15 , 17 -14 , 17 -33, 17 -36, 18 -20, 18 - 21, 18 -23, 19 -9, 20-3, 20-5, 20-6CD, 20-7, 20-8, 20 -13 , 20 -14 , 20 -17 , and 20 -19 A Blaine... Maastricht, The Netherlands Figure 10 -7C Prem S Sahni, MD Formerly of the Department of Radiology, Children’s Hospital, Winnipeg, Manitoba, Canada Figures 8 -11 C, 10 -7B, 10 -13 , 11 -4C, 11 -28B, 12 -16 , 12 -17 ,... Membranes 10 7 Placenta 10 7 Decidua 10 9 Development of Placenta 10 9 Placental Circulation 11 1 Placental Membrane 11 3 Functions of Placenta 11 4 Placental Endocrine Synthesis and Secretion 11 7 The