1 / 2671 Gray's Anatomy, 40th Edition By Susan Standring, PhD, DSc, FKC A universal landmark in medicine ever since Drs. Henry Gray and H.V. Carter published the first edition in 1858, Gray's Anatomy now celebrates its 150th anniversary! Book features: Completely re-imaged, full-colored art program Meticulously enhanced and updated artwork throughout, with 1,800 images now in full color More than 120 completely new radiological, CT, MT and other imaging studies Getting started Additional Features Bibliography: An extended bibliography, To start browsing, use the table of contents on the left. Click to expand the contents compiled over several editions of this of a section or chapter. Clicking the chapter or section title itself will take you to that text, furnishes an exhaustive library of section. Alternatively, search the book using the search function above, or look up a key articles. term in the complete index. Eponyms: Hundreds of eponyms and their definitions are provided. For further information on Expert Consult, view a demo of the site. 2 / 2671 Gray's Anatomy The Anatomical Basis of Clinical Practice Fortieth Edition EDITOR-IN-CHIEF: Susan Standring, PhD, DSc, FKC Emeritus Professor of Anatomy, King's College London, London, UK SECTION EDITORS: Neil R Borley, FRCS, FRCS(Ed), MS Consultant Colorectal Surgeon, Department of Gastrointestinal Surgery, Cheltenham General Hospital, Gloucestershire Hospitals NHS Trust, Cheltenham, UK Patricia Collins, PhD Associate Professor of Anatomy, Anglo-European College of Chiropractic, Bournemouth, UK Alan R Crossman, PhD, DSc Professor of Anatomy, Faculty of Life Sciences, The University of Manchester, Manchester, UK Michael A Gatzoulis, MD, PhD, FESC, FACC Professor of Cardiology, Congenital Heart Disease Consultant Cardiologist, Adult Congenital Heart Centre and Centre for Pulmonary Hypertension, Royal Brompton Hospital, and the National Heart and Lung Institute, Imperial College, London, UK Jeremiah C Healy, MA, MB BChir, MRCP, FRCR Consultant Radiologist, Chelsea and Westminster Hospital; Honorary Senior Lecturer Imperial College, London, UK David Johnson, MA, BM BCh, DM, FRCS(Eng) Consultant in Plastic, Reconstructive and Craniofacial Surgery, Department of Plastic and Reconstructive Surgery, Radcliffe Infirmary, Oxford, UK Vishy Mahadevan, PhD, FRCS(Ed), FRCS(Eng) Professor of Surgical Anatomy and Barbers' Company Reader in Anatomy, Raven Department of Education, The Royal College of Surgeons of England, London, UK Richard LM Newell, BSc, MB BS, FRCS Honorary Consultant Orthopaedic Surgeon, Royal Devon and Exeter Healthcare NHS Trust, Exeter, UK Caroline B Wigley, BSc, PhD Senior Honorary, University Teaching Fellow, College of Medicine and Dentistry Peninsula, Exeter, UK CHURCHILL LIVINGSTONE ELSEVIER is an imprint of Elsevier. © 2008, Elsevier Limited. All rights reserved. Fortieth edition published 2008 First edition JW Parker & Son 1858 The right of Susan Standring, Neil R Borley, Patricia Collins, Alan R Crossman, Michael A Gatzoulis, Jeremiah C Healy, David Johnson, Vishy Mahadevan, Richard LM Newell and Caroline Wigley to be identified as authors of this work has been asserted by them in accordance with the Copyright, Designs and Patents Act 1988. 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. Permissions may be sought directly from Elsevier's Rights Department: phone: (+1) 215 239 3804 (US) or (+44) 1865 843830 (UK); fax: (+44) 1865 853333; e-mail: [email protected] . You may also complete your request on-line via the Elsevier website at http://www.elsevier.com/permissions . Expert Consult ISBN: 978-0-443-06684-9 International Edition ISBN: 978-0-8089-2371-8 British Library Cataloguing in Publication Data A catalogue record for this book is available from the British Library Library of Congress Cataloging in Publication Data A catalog record for this book is available from the Library of Congress 3 / 2671 Notice No responsibility is assumed by the Publishers, Editors or Contributors for any loss or injury and/or damage to persons or property arising out of or related to any use of the material contained in this book. It is the responsibility of the treating practitioner, relying upon independent expertise and knowledge of the patient, to determine the best treatment and method of application for the patient. Every effort has been made to contact holders of copyright to obtain permission to reproduce copyright material. However, if any have been inadvertently overlooked, the Publisher will be pleased to make the necessary arrangements at the first opportunity. The Publisher Printed in Spain Last digit is the print number: 9 8 7 6 5 4 3 2 1 Commissioning Editors: Madelene Hyde and Inta Ozols Development Editors: Martin Mellor, Gavin Smith and Alison Whitehouse Head of Development (Global Medicine): Louise Cook Editorial Assistants: Julie Goolsby and Liam Jackson Project Managers: Glenys Norquay and Bryan Potter Illustration Managers: Merlyn Harvey, Bruce Hogarth and Gillian Richards Illustrators (39th and 40th editions): Antbits (Paul Banville, Michael Courtney, Jonathan Dimes, Paul Richardson, Richard Tibbitts), Robert Britton, Joe Chovan, Peter Cox, Ethan Danielson, Brian Evans, Sandie Hill, Bruce Hogarth, Gillian Lee, Debbie Maizels, Annabel Milne, Philip Wilson, Martin Woodward Page Design: Jayne Jones Cover Design: Stewart Larking Copyeditors: Carolyn Holleyman, with Alyson Colley Proofreader: Ian Ross Indexer: Susan Boobis Marketing Managers: Jeremy Bowes (UK) and Lisa Damico (US) 4 / 2671 COVER IMAGE ‘Tractography reconstruction of the perisylvian language pathways' from Catani M, Jones DK & ffytche DH, Ann Neurol 2005; 57(1):8–16 with permission of John Wiley & Sons. After almost 120 years this tractography reconstruction of the language pathways has updated the anatomy of the language networks. Image courtesy of Marco Catani, Natbrainlab, Centre for Neuroimaging Sciences and Section of Brain Maturation, Department of Psychological Medicine, Institute of Psychiatry, King's College London. 5 / 2671 PREFACE The 40th edition of Gray's Anatomy celebrates 150 years of continuous publication of an extraordinary book. Although this edition looks very different from the first edition (see Ruth Richardson's historical introduction on page xvii), the essential aim of Henry Gray and the illustrator Henry Vandyke Carter, to describe the clinically relevant anatomy of the human body, particularly (but not exclusively) for the practicing surgeon, has not changed over the years. Anatomy may be struggling to retain its foothold in many undergraduate medical curricula, but it is enjoying a renaissance at the postgraduate level. A detailed knowledge of clinical anatomy which would have been undreamed of in the first edition, is now essential for clinicians working in technologically driven fields such as imaging, endoscopic and robotic surgery. Nine Section Editors (Neil Borley, Pat Collins, Alan Crossman, Michael Gatzoulis, Jeremiah Healy, David Johnson, Vishy Mahadevan, Richard Newell and Caroline Wigley) have worked with me in preparing the 40th edition. They have brought their extensive experience as anatomists, cell biologists and clinicians to the task, and I thank them for their dedication and enthusiastic support. Pat Collins, Jeremiah Healy and Caroline Wigley also worked closely with all members of the editorial team in updating the text and artworks for embryology, imaging and microstructure respectively throughout the book. Harold Ellis has meticulously edited the section on eponyms which is available on the website, and has commented on many of the chapters. Each Section Editor was assisted by a group of Contributors – experienced anatomists and clinicians (sometimes both) who contributed text and/or artworks and original micrographs. The page proofs were scrutinized by a core team of Reviewers and finally a panel of International Reviewers: their comments have been incorporated into the text and I acknowledge their critical input. Working at this level of detail I have often been impressed by how much anatomy remains controversial or is simply unknown: surgeons, radiologists and embryologists often disagree passionately about structural relationships or developmental processes. Gray's continues to flag up these uncertainties and to present different perspectives. As a general rule, the orientation of diagrams and photographs throughout the book has been standardized to show the left side of the body, irrespective of whether a lateral or medial view is presented, and transverse sections are viewed from below to facilitate comparison with clinical images. Clinicopathological examples have been selected where the pathology is either a direct result of, or a consequence of, the anatomy, or where the anatomical features are instrumental in the diagnosis/treatment/management of the condition. All of the pre 39th edition artworks, and the great majority of the images and micrographs of histological and embryological specimens have been replaced: wherever possible, the photomicrographs illustrate human histology and embryology, and non human sources have been acknowledged in the captions. New artworks have either been generated de novo or have been taken from other texts (principally Sobotta's Atlas of Anatomy, Books 1 and 2 or Gray's Atlas of Anatomy), making this the first full-colour edition of Gray's Anatomy. I am often asked why Gray's Anatomy does not contain even more detailed surgical and radiological anatomy, with further examples of anatomical variants and of laparoscopic and endoscopic anatomy, and why we exclude all but a very few references. The answer is that we would love to do this, and to include some systematic anatomy, but we have reached a point where there is no room for more material in a single volume. Short reference lists are provided at the ends of each chapter to guide further reading, and a list of general texts and references covering material presented in more than one chapter, e.g. the distribution of angiosomes, appears on page xxiv. The Bibliography, which collated all of the references cited in a number of earlier editions 35 –38, is retained on the website. I offer my sincere thanks to the editorial team at Elsevier, initially under the leadership of Inta Ozols and latterly of Madelene Hyde, for their guidance, professionalism, good humour and unfailing support. In particular, I thank Alison Whitehouse, Gavin Smith, Martin Mellor and Louise Cook, for being at the end of a phone or e-mail whenever I needed advice. I am especially grateful to my dear husband, Guy Standring, for his tolerance while he has shared his life with Gray's Anatomy, and I dedicate my work in this book to him. Susan Standring May 2008 6 / 2671 ACKNOWLEDGEMENTS Within individual figure captions, we have acknowledged all figures kindly loaned from other sources. However, we would particularly like to thank the following authors who have generously loaned so many figures from other books published by Elsevier: Drake R, Vogl W, Mitchell AWM 2004 Gray's Anatomy for Students. Edinburgh: Churchill Livingstone. Drake R, Vogl W, Mitchell AWM, Tibbitts R, Richardson P 2008 Gray's Atlas of Anatomy. Edinburgh: Churchill Livingstone. Putz R, Pabst R 2006 Sobotta: Atlas der Anatomie des Menschen. Band 1. Kopf, Hals, obere Extremitat. 22 Auflage. Munchen: Elsevier. Much of the illustration in Gray's Anatomy has as its basis the work of illustrators and photographers who contributed towards earlier editions, their figures sometimes being retained almost unchanged though more often being used as the foundation for figures that are new to this edition. The Publishers would like to thank them. For illustration reused from the 39th edition, we particularly thank its photographer Sarah-Jane Smith and the artists listed on page iv. The Publishers would also like to thank: Anne-Marie Childs BSc, MB ChB, MRCP, FRCPCH, Consultant Paediatric Neurologist, The General infirmary at Leeds; James M McCall BSc, MBBS, MRCP, FRCR, Consultant Radiologist, Chelsea and Westminster Hospital, London, and Romney JE Pope MA, MRCP, FRCR, Consultant Radiologist, Royal Marsden and Chelsea and Westminster Hospitals, London, for their help in preparing material for the new edition. 7 / 2671 CONTRIBUTORS Michael A Adams, BSc, PhD Reader in Spine Biomechanics, Department of Anatomy, University of Bristol, Bristol, UK Andrew Amis, PhD, DSc(Eng), FIMechE Professor of Orthopaedic Biomechanics, Imperial College London, London, UK Robert H Anderson, BSc, MD, FRCPath Formerly Professor of Paediatric Cardiac Morphology, Institute of Child Health, University College, London, UK Professor of Paediatrics, Medical University of South Carolina, Charleston, South Carolina, USA Tipu Aziz, BSc, MBBS, MD, FRCS, FRCS(SN), DMedSci Professor of Neurosurgery, University of Oxford, Oxford, UK Timothy J Beale, MBBS FRCS FRCR Consultant Radiologist, Royal National Throat Nose and Ear Hospital, London, UK Sue M Black, OBE, BSc, PhD, DSc, FRSE, Hon FRCPS (Glasg) Professor, Head of Anatomy and Human Identification, College of Life Sciences, University of Dundee, Dundee, UK Nikolai Bogduk, BSc (Med), MB BS, PhD, MD, DSc, DipAnat, Dip Pain Med, FAFRM, FAFMM, FFPM(ANZCA) Professor of Pain Medicine, University of Newcastle, Newcastle, New South Wales, Australia Peter R Braude, PhD, FRCOG, FMedSci Professor and Head of Department of Women's Health, King's College London Honorary Consultant in Reproductive Medicine, Guy's and St Thomas' Foundation Trust, London, UK Jonathan L Brown, MA, MSc, MD, FRCP, DTMH Professor and Consultant Physician and Gastroenterologist, Gloucestershire Hospitals NHS Trust, Gloucester, UK Anthony Bull, BEng, DIC CEng, PhD, FIMechE Reader in Musculoskeletal Mechanics, Department of Bioengineering, Imperial College London, London, UK Andrew Bush, MB BS (Hons), MA MD FRCP, FRCPCH Professor of Paediatric Respirology, Department of Paediatric Respiratory Medicine, Imperial College and Royal Brompton Hospital, London, UK Declan JP Cahill, BSc, MSc, FRCS (Urol) Consultant Urologist, Department of Urology, Guy's and St Thomas' NHS Trust, London, UK Cécile Chalouni, PhD Technical Director and Associate Research Scientist in Cell Biology, Department of Cell Biology, Ludwig Institute for Cancer Research, Yale University School of Medicine, New Haven, CT, USA Ashish Chandra, FRCPath, DipRCPath (Cyto) Consultant Histopathologist, Guy's and St. Thomas' Hospital NHS Foundation Trust, London, UK Roger Chinn, MB BS, MRCP, FRCR Consultant Radiologist, Department of Radiology, Chelsea and Westminster NHS Foundation Trust, London, UK Bodo EA Christ, MD, PhD, h.c. Professor, Director of the Institute of Anatomy and Cell Biology, University of Freiburg, Freiburg, Germany Alfred Cutner, MD, FRCOG Consultant Gynaecologist, Department of Obstetrics and Gynaecology, University College London Hospitals, London, UK Catriona L Davies, MB BS, MRCP, FRCR Consultant Radiologist, Chelsea and Westminster Hospital, London, UK Patricia Dolan, BSc, PhD Reader in Biomechanics, Department of Anatomy, University of Bristol, Bristol, UK Ronald H Douglas, BSc, PhD Professor of Visual Science, Department of Optometry and Visual Science, City University, London, UK Justin A Durham, BDS, MFDS, RCS (Ed), FHEA Clinical Fellow and Honorary Specialist Registrar in Oral Surgery, School of Dental Sciences, Newcastle University, Newcastle, UK William C Earnshaw, PhD, FRSE Professor and Wellcome Trust Principal Research Fellow, Wellcome Trust Centre for Cell Biology, University of Edinburgh, Edinburgh, UK David M Evans, FRCS Consultant Hand Surgeon, Clinical Director, The Hand Clinic, Windsor, UK Paul A Felts, PhD Senior Lecturer, Centre for Anatomy and Human Identification, University of Dundee, Dundee, UK 8 / 2671 Alex Freeman, MSc, MBBS, MD, MRCPath Consultant Urological Pathologist, University College London Hospitals NHS Foundation Trust, London, UK David Furness, PhD Reader in Auditory Neuroscience, MacKay Institute of Communication and Neuroscience, The School of Life Sciences Keele University, Keele, UK Andrew JT George, MA, PhD, FRCPath, FRSA Professor of Molecular Immunology, Department of Immunology, Division of Medicine, Faculty of Medicine, Imperial College London, London, UK Jonathan M Glass, BSc, MB BS, FRCS (Urol) Urologist, Department of Urology, Guy's and St Thomas' Hospitals, London, UK Michael Gleeson, MD, FRCS Consultant Otolaryngologist and Skull Base Surgeon to Guy's, Kings and St. Thomas' Hospitals, London Professor of Skull Base Surgery, National Hospital for Neurology and Neurosurgery, London Honorary Skull Base Surgeon to the Great Ormond Street Hospital for Sick Children, London, UK David JA Goldsmith, MA, FRCP Consultant Nephrologist, Kidney Unit, Guy's Hospital, London, UK Anthony Graham, BSc (Hons), PhD Professor of Developmental Biology, MRC Centre for Developmental Neurobiology and Division of Anatomy and Human Sciences, School of Biomedical and Health Sciences, Guy's Campus, King's College London, London, UK Paul D Griffiths, MB ChB, PhD, FRCR Professor of Radiology, Academic Unit of Radiology, Division of Clinical Science, University of Sheffield, Sheffield, UK Gerald PH Gui, MS, FRCS, FRCS(Ed) Consultant Surgeon, Academic Surgery (Breast Unit), Royal Marsden NHS Foundation Trust, London, UK Chinmay M Gupte, FRCS(Tr & Orth), PhD, MA (Oxon), MRCS Specialist Registrar, St Mary's Hospital and North West Thames, London, UK Carole M Hackney, BSc (Hons), PhD Professor of Auditory Neuroscience, MacKay Institute of Communication and Neuroscience, School of Life Sciences, Keele University, Keele, UK Peter A Helliwell, FIBMS, CSci, CBA, Cert Ed Chief Scientist and Manager, Molecular and Cell Biology Unit, Department of Cellular Pathology, Royal Cornwall Hospital, Truro, UK John M Hutson, AO Professor of Paediatric Surgery, University of Melbourne, Department of Urology, Royal Children's Hospital, Parkville, Australia Alan Jackson, BSc (Hons), PhD, MRCP, FRCR, FRCP Professor of Radiology Imaging, Science and Biomedical Engineering, Wolfson Molecular Imaging Centre, The University of Manchester, Manchester, UK Bruce W Jafek, MD, FACS, FRSM Professor and Former Chairman (1976–1998), Department of Otolaryngology, Head and Neck Surgery, University of Colorado School of Medicine, Denver, Colorado, USA Eric R Jauniaux, MD, PhD, MRCOG Professor and Consultant in Materno-Fetal Medicine, Department of Obstetrics and Gynaecology, University College London Medical School, London, UK Jonathan C Kentish, MA, PhD Professor of Cellular Cardiology, Cardiovascular Division, King's College London, London, UK Aadil A Khan, BM BCh, MPH, MRCS(Eng) Speciality Trainee in Plastic Surgery, Oxford Region, UK Khan Nasir, MBBS, MRCP, FRCR Consultant Radiologist, Chelsea and Westminster Hospital Royal Marsden Hospital, London, UK W Niall A Kirkpatrick, BDS, MB BS, MD, FRCS, FRCS(Plast) Consultant Craniofacial Plastic Surgeon, Craniofacial Unit, Chelsea and Westminster Hospital, London, UK John G Lawrenson, BSc, PhD, MCOptom Professor of Clinical Visual Science, Department of Optometry and Visual Science, City University, London, UK Justin Lee, BSc, MBBS, MRCS(Eng), FRCR Consultant Radiologist, Department of Radiology, Chelsea and Westminster Hospital, London, UK David Lowe, MD, FRCS, FRCPath, FIBiol 9 / 2671 Consultant Histopathologist, The London Clinic, London, UK Andres Lozano, MD, PhD, FRCSC Professor, Division of Neurosurgery, University of Toronto, Toronto, Ontario, Canada Joseph Mathew, MBBS, FMCPath, FRCPath, CertTLHE, PGCE, CertBusStud, FHEA Consultant in Histopathology, Department of Histopathology, Royal Cornwall Hospitals Trust, Truro, UK Stephen McHanwell, BSc, PhD Professor of Anatomical Sciences, School of Dental Sciences, The Dental School, Framlington Place, Newcastle-Upon-Tyne, UK Mary-Clare C Miller, MA (Cantab), MB BS, MRCS (Eng) Specialty Registrar in Plastic Surgery, London Deanery, London, UK Sukhbinder Minhas, MD, FRCS (Urol) Consultant Uro-andrologist and Senior Lecturer, The Institute of Urology, University College Hospital, London, UK Zoltán Molnár, MD, DPhil Professor of Developmental Neuroscience, Department of Physiology, Anatomy and Genetics, University of Oxford, Oxford, UK Michael Monteiro, BDS, FDSRCS, MB BCH, MRCS Specialist Registrar, Department of Oral and Maxillofacial Surgery, Brighton and Sussex University Hospitals NHS Trusts, Brighton, UK Louise A Moore, MB ChB, Dip Paeds, FRANZCR Consultant Radiologist, Department of Radiology, Tauranga Hospital, Tauranga, New Zealand Antoon FM Moorman, PhD Professor of Embryology and Molecular, Biology of Cardiovascular Diseases, Head, Department of Anatomy and Embryology, Academic Medical Center, Amsterdam, The Netherlands Gillian M Morriss-Kay, BSc (Hons), PhD, DSc Professor Emeritus, Department of Physiology, Anatomy and Genetics, University of Oxford, Oxford, UK Timothy AJ Mould, MA (Oxon), DM (Oxon), FRCOG Consultant Gynaecological Oncologist, University College London Hospitals Gynaecological Cancer Centre, University College London Hospitals, London, UK Jagdeep Nanchahal, BSc, PhD, MBBS, FRCS(plast), FRACS Professor of Hand, Plastic and Reconstructive Surgery, Kennedy Institute of Rheumatology and Charing Cross Hospital, Imperial College, London, UK David Neary, MD, FRCP Professor of Neurology, The Greater Manchester Neuroscience Centre, Salford Royal Hospital, University of Manchester, Manchester, UK Niri S Niranjan, MBBS, LRCP, MS, FRCS, FRCS(Plast), FRCS Ed, FRCS Eng Consultant Plastic, Reconstructive and Cosmetic Surgeon, St. Andrew's Centre for Burns and Plastic Surgery, Broomfield Hospital, Chelmsford, UK Simon Padley, BSc, MBBS, FRCP, FRCR Consultant Radiologist, Royal Brompton and Harefield NHS Trust, Honorary Senior Lecturer, Imperial College Medical School, London, UK Pranav P Pandya, BSc, MRCOG, MD Consultant and Honorary Senior Lecturer in Fetal Medicine and Obstetrics, Institute for Women's Health, Department of Obstetrics and Gynaecology, University College London Hospitals, London, UK Terence A Partridge, BSc, PhD, FMed. Sci Research Faculty, Research Center for Genetic Medicine, Children's National Medical Center, Washington, DC, USA Hannes Petersen, MD, PhD Head of Department and Associate Professor, Department of Anatomy and Otorhinolaryngology, Head and Neck Surgery, University of Iceland and Landspitali University Hospital, Reykjavik, Iceland Raj Prasad, MS, MCh, FRCS Consultant Surgeon, Hepatobiliary and Transplant Unit, St James's University Hospital, Leeds, UK Donald J Reid, PhD Lecturer in Oral Anatomy, The Dental School, School of Dental Sciences, Newcastle-Upon-Tyne, UK Patricia A Reynolds, BDS, MBBS, MAODE(Open), PhD, FDSRCS(Eng)(Edin) Professor of Dental Education, King's College London Dental Institute, London, UK Paul D Robinson, PhD, MBBS, BDS, FDSRCS Oral and Maxillofacial Surgeon (Retired from), Department of Maxillofacial Surgery, King's College London Dental Institute, London, UK 10 / 2671