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Gray's Anatomy: The Anatomical Basis of Clinical Practice, 41eSusan Standring PhD DSc
In 1858, Drs. Henry Gray and Henry Vandyke Carter created a book for their surgical colleagues that established an enduring standard among anatomical texts. After more than 150 years of continuous publication, Gray's Anatomy remains the definitive, comprehensive reference on the subject, offering ready access to the information you need to ensure safe, effective practice.
This 41st edition has been meticulously revised and updated throughout, reflecting the very latest understanding of clinical anatomy from field leaders around the world. The book's traditional lavish art programme and clear text have been further honed and enhanced, while major advances in imaging techniques and the new insights they bring are fully captured in new state-of-the-art X-ray, CT, MR, and ultrasonic images.
The completely reconfigured accompanying eBook version is richly enhanced with additional content and media, covering all the body regions, cell biology and embryogenesis. This unlocks a whole new level of related information, interactivity and understanding, in keeping with the spirit of innovation that has characterized Gray's Anatomy since its inception.
- Presents the most detailed and dependable coverage of anatomy available anywhere.
- Regional organization collects all relevant material on each body area together in one place, making access to core information easier for clinical readers.
- Anatomical information is matched with key clinical information where relevant.
- Numerous clinical discussions emphasize considerations that may affect medical care.
- Each chapter has been edited by experts in their field, ensuring access to the very latest evidence-based information on that topic.
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- The downloadable Expert Consult eBook version included with your purchase allows you to search all of the text, figures, references and videos from the book on a variety of devices.
- Carefully selected electronic enhancements include additional text, tables, illustrations, labelled imaging and videos - as well as 24 specially invited 'Commentaries' on new and emerging topics related to anatomy.
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Grays Anatomy-The Anatomical Basis of Clinical Practice -41 E (2015) [PDF] [UnitedVRG].pdf GRAY'S Anatomy Commissioning Editors: Jeremy Bowes and Madelene Hyde Development Editors: Humayra Rahman Khan and Poppy Garraway Content Development Manager: Louise Cook Project Managers: Julie Taylor and Joanna Souch Production Manager: Deena Burgess Illustrators (41st edition): Antbits (Paul Richardson, Richard Tibbitts), Robert Britton, Joe Chovan, Peter Cox, Ethan Danielson, Brian Evans, Sandie Hill, Bruce Hogarth, Gillian Lee, Debbie Maizels, Annabel Milne, Amanda Williams, Philip Wilson, Martin Woodward Page and Cover Design: Christian Bilbow Copyeditor: Wendy Lee Proofreaders: Susan Stuart and Glenys Norquay Indexer: Jan Ross, Merrall-Ross International Ltd Marketing Manager: Melissa Darling Multimedia Producer: Megan Graieg Inkling Development Specialist: Kim Benson EDITOR-IN-CHIEF Susan Standring MBE, PhD, DSc, FKC, Hon FAS, Hon FRCS Emeritus Professor of Anatomy King's College London London, UK SECTION EDITORS Anatomy G RA Y' S The Anatomical Basis of Clinical Practice Neel Anand MD Clinical Professor of Surgery Director, Spine Trauma, Minimally Invasive Spine Surgery Spine Center Cedars Sinai Medical Center Los Angeles, CA, USA Rolfe Birch MChir, FRCPS(Glasg), FRCS(Ed), FRCS(Eng) Retired Consultant in Charge War Nerve Injury Clinic, Defence Medical Rehabilitation Centre, Surrey; Retired Head, Peripheral Nerve Injury Unit, Royal National Orthopaedic Hospital; Professor in Neurological Orthopaedic Surgery, University College of London London, UK Patricia Collins BSc, PhD, FHEA Professor of Anatomy Anglo-European College of Chiropractic Bournemouth, UK Editor for Embryology and Development Alan R Crossman BSc, PhD, DSc Professor Emeritus University of Manchester Manchester, UK Michael Gleeson MD, FRCS, FRACS, FDS Professor of Skull Base Surgery University College London The National Hospital for Neurology and Neurosurgery London, UK Girish Jawaheer MD, FRCS(Eng), FRCS(Paed) Consultant Paediatric Surgeon Great North Children's Hospital, Royal Victoria Infirmary Newcastle upon Tyne NHS Foundation Trust Newcastle upon Tyne, UK; Formerly Specialty Tutor for Paediatric Surgery Royal College of Surgeons of England London, UK Editor for Paediatric Anatomy Ariana L Smith MD Associate Professor of Urology Director of Pelvic Medicine and Reconstructive Surgery Penn Medicine, Perelman School of Medicine University of Pennsylvania Health System Philadelphia, PA, USA Jonathan D Spratt MA(Cantab), FRCS(Eng), FRCR Clinical Director of Diagnostic Radiology City Hospitals Sunderland NHS Foundation Trust Sunderland, UK; Former Anatomy Examiner for the Royal College of Surgeons of England and Royal College of Radiologists Editor for Imaging Anatomy Mark D Stringer BSc, MS, FRCP, FRCS, FRCS(Ed), FRACS Professor of Paediatric Surgery Christchurch Hospital; Honorary Professor of Anatomy University of Otago Dunedin, New Zealand R Shane Tubbs MS, PA-C, PhD Chief Scientific Officer Seattle Science Foundation, Seattle, WA, USA; Professor of Human Gross and Developmental Anatomy Department of Anatomical Sciences St. George's University, Grenada, West Indies; Professor Centre of Anatomy and Human Identification University of Dundee, Dundee, UK Richard Tunstall BMedSci, PhD, PGCLTHE FHEA Head of Clinical Anatomy and Imaging Warwick Medical School University of Warwick, UK; University Hospitals Coventry and Warwickshire NHS Trust Coventry, UK; Visiting Professor of Anatomy St George's University, Grenada, West Indies; Editor for Surface Anatomy Alan J Wein MD, PhD(Hon), FACS Founders Professor and Chief of Urology Director, Urology Residency Program Penn Medicine, Perelman School of Medicine University of Pennsylvania Health System Philadelphia, PA, USA Caroline B Wigley BSc, PhD University of Exeter Medical School Exeter, UK Editor for Cell and Tissue Microstructure FORTY-FIRST EDITION http://M.Chir © 2016, Elsevier Limited. All rights reserved. Forty-first edition published 2016 Fortieth edition published 2008 First edition JW Parker & Son 1858 The right of Susan Standring, Neel Anand, Rolfe Birch, Patricia Collins, Alan R Crossman, Michael Gleeson, Girish Jawaheer, Ariana Smith, Jonathan D Spratt, Mark D Stringer, R Shane Tubbs, Richard Tunstall, Alan J Wein and Caroline B 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 photocopy- ing, 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@example.com. You may also complete your request online via the Elsevier website at http://www.elsevier.com/permissions. ISBN: 978-0-7020-5230-9 978-0-7020-6306-0 978-0-7020-6851-5 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 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 copy- right material. However, if any have been inadvertently overlooked, the Publisher will be pleased to make the necessary arrangements at the first opportunity. The Publisher COVER IMAGE Reconstruction of the short and long connections of the hand region in the living human brain using advanced diffusion tractography developed by the NatBrainLab (www.natbrainlab.com). The connections of the hand region resemble a 'poppy flower' with a green stem representing the long projection fibres and four red 'petals' connecting the precentral cortex to postcentral and premotor cortices. This network is important for motor learning and execution of reaching and grasping hand movements (see chapter 25). (Image courtesy of Dr Marco Catani, originally published in Catani M, Dell'Acqua F, Vergani F, et al; Short frontal lobe connections of the human brain. 2012 Cortex 48:273–91.) The publisher's policy is to use paper manufactured from sustainable forests Get the most out of your Gray's Anatomy Included in your purchase is a rich variety of BONUS electronic content, to supplement and enhance the printed book. Look out for this icon , indicating where there is related further electronic text, images, tables, labelled imaging or video material. In addition, you can access 24 'Gray's Commentaries', especially written (electronic only) articles on new and emerging topics related to anatomy. This icon highlights where one relates directly to a topic in the printed book. Don't miss out on any of this additional electronic content – see the inside front cover for your access instructions. Forty-First Edition mailto:firstname.lastname@example.com http://www.elsevier.com/permissions http://www.natbrainlab.com v CONTENTS Video 1.5.1 – Diagnostic histopathology by electron microscopy Video 1.5.2 – Serial block face scanning electron microscopy (SBFSEM) Roland A Fleck SECTION 2: EMBRYOGENESIS Section Editor: Patricia Collins 8 Preimplantation development 163 Alison Campbell, Patricia Collins 9 Implantation and placentation 171 Eric Jauniaux, Graham J Burton 10 Cell populations at gastrulation 181 Patricia Collins 11 Embryonic induction and cell division 189 Patricia Collins 12 Cell populations at the start of organogenesis 193 Patricia Collins 13 Early embryonic circulation 200 Patricia Collins 14 Pre- and postnatal development 205 Patricia Collins, Girish Jawaheer 15 Development of the limbs 218 Cheryll Tickle Commentaries 2.1 Human anatomy informatics Jonathan BL Bard, Paul N Schofield 2.2 An evolutionary consideration of pharyngeal development Anthony Graham, Victoria L Shone Videos Video 8.1 – Human in vitro fertilization and early development Alison Campbell Video 9.1 – Ultrasound features of the maternal placental blood flow Eric Jauniaux Video 14.1 – Ultrasound features of the fetus at 26 weeks Jonathan D Spratt, Patricia Collins SECTION 3: NEUROANATOMY Section Editor: Alan R Crossman 16 Overview of the nervous system 227 Alan R Crossman, Richard Tunstall 17 Development of the nervous system 238 Zoltán Molnár 18 Ventricular system and subarachnoid space 271 Jacob Bertram Springborg, Marianne Juhler Preface ix Preface Commentary The continuing relevance of anatomy in current surgical practice and research R Shane Tubbs Acknowledgements x Contributors xi Historical introduction A brief history of Gray's Anatomy Ruth Richardson Anatomical nomenclature xvi Bibliography of selected titles xviii SECTION 1: CELLS, TISSUES AND SYSTEMS Section Editor: Caroline B Wigley 1 Basic structure and function of cells 4 Abraham L Kierszenbaum 2 Integrating cells into tissues 28 Caroline B Wigley 3 Nervous system 42 Helmut Kettenmann 4 Blood, lymphoid tissues and haemopoiesis 68 Andrew JT George 5 Functional anatomy of the musculoskeletal system 81 Michael A Adams 6 Smooth muscle and the cardiovascular and lymphatic systems 123 Jeremy PT Ward 7 Skin and its appendages 141 John A McGrath, Joey E Lai-Cheong Commentaries 1.1 Fluorescence microscopy in cell biology today Dylan M Owen 1.2 Stem cells in regenerative medicine Jonathan M Fishman, Paolo De Coppi, Martin A Birchall 1.3 Merkel cells Ellen A Lumpkin 1.4 Metaplasia Jonathan MW Slack, Leonard P Griffiths, David Tosh 1.5 Electron microscopy in the twenty-first century Roland A Fleck 1.6 The reaction of peripheral nerves to injury Rolfe Birch Videos Video 1.1 – Mitosis in a cell with fluorescently-labelled chromosomes and microtubules Jonathon Pines, Daisuke Izawa CONTENTS vi 19 Vascular supply and drainage of the brain 280 Paul D Griffiths 20 Spinal cord: internal organization 291 Monty Silverdale 21 Brainstem 309 Duane E Haines 22 Cerebellum 331 Jan Voogd 23 Diencephalon 350 Ido Strauss, Nir Lipsman, Andres M Lozano 24 Basal ganglia 364 Tipu Aziz, Erlick AC Pereira 25 Cerebral hemispheres 373 Guilherme C Ribas Commentary 3.1 The resting human brain and the predictive potential of the default mode network Stefano Sandrone Videos Video 18.1 – Interactive 3D rotation of the subarachnoid space Video 18.2 – Interactive 3D rotation of the ventricles and cisterns Jose C Rios Video 19.1 – Rotational angiography of an intracranial aneurysm Paul D Griffiths SECTION 4: HEAD AND NECK Section Editor: Michael Gleeson 26 Head and neck: overview and surface anatomy 404 Michael Gleeson, Richard Tunstall Head and Neck 27 External skull 416 Sue Black 28 Intracranial region 429 Juan C Fernandez-Miranda 29 Neck 442 John C Watkinson, Michael Gleeson 30 Face and scalp 475 Simon Holmes Upper Aerodigestive Tract 31 Oral cavity 507 Barry KB Berkovitz 32 Infratemporal and pterygopalatine fossae and temporomandibular joint 534 Barrie T Evans 33 Nose, nasal cavity and paranasal sinuses 556 Claire Hopkins 34 Pharynx 571 Stephen McHanwell 35 Larynx 586 Stephen McHanwell 36 Development of the head and neck 605 Gillian M Morriss-Kay Special Senses 37 External and middle ear 624 Michael Gleeson 38 Inner ear 641 David N Furness 39 Development of the ear 658 Susan Standring 40 Development of the eye 661 Jane C Sowden 41 Orbit and accessory visual apparatus 666 John G Lawrenson, Ronald H Douglas 42 Eye 686 Ronald H Douglas, John G Lawrenson Commentaries 4.1 Surgery of the skull base Juan C Fernandez-Miranda 4.2 The role of three-dimensional imaging in facial anatomical assessment Vikram Sharma, Bruce Richard 4.3 Anatomy of facial ageing Bryan C Mendelson, Chin-Ho Wong Videos Video 28.1 – 3D surface rotation of the sella turcica in the horizontal plane Video 28.2 – 3D surface rotation of the sella turcica in the multiaxial plane Video 28.3 – 3D surface rotation of the sella turcica in the vertical plane Michael D Luttrell Video 30.1 – Pan-facial fractures Video 30.2 – Postoperative cranio-orbital imaging Video 30.3 – A comminuted zygomatic fracture (plus Le Fort I) pattern Video 30.4 – A comminuted zygomatic fracture pattern – post reduction Simon Holmes Video 32.1 – Temporomandibular joint arthroscopy demonstrating intracapsular anatomy of the joint Gary Warburton Video 32.2 – Endoscopic anatomy of the infratemporal and pterygopalatine fossae Carl H Snyderman, Juan C Fernandez-Miranda Video 4.2.1 – 3D anatomical imaging of the face Vikram Sharma SECTION 5: BACK Section Editor: Neel Anand 43 Back 710 Eli M Baron, Richard Tunstall 44 Development of the back 751 Bodo EA Christ, Martin Scaal 45 Spinal cord and spinal nerves: gross anatomy 762 Eli M Baron Commentary 5.1 Minimally invasive surgical corridors to the lumbar spine Y Raja Rampersaud CONTENTS vii SECTION 6: PECTORAL GIRDLE AND UPPER LIMB Section Editor: Rolfe Birch 46 Pectoral girdle and upper limb: overview and surface anatomy 776 Rolfe Birch, Richard Tunstall 47 Development of the pectoral girdle and upper limb 794 Cheryll Tickle 48 Shoulder girdle and arm 797 Simon M Lambert 49 Elbow and forearm 837 Leela C Biant 50 Wrist and hand 862 Alistair C Ross Commentaries 6.1 Injuries of the supraclavicular brachial plexus Rolfe Birch 6.2 Nerves at risk from musculoskeletal injury Rolfe Birch 6.3 Thoracic outlet syndromes Rolfe Birch Videos Video 46.1 – Upper limb surface anatomy Rolfe Birch Video 50.1 – Movements of the hand Rolfe Birch Video 50.2 – Wrist block: surface anatomy Dominic Harmon SECTION 7: THORAX Section Editor: Jonathan D Spratt 51 Thorax: overview and surface anatomy 898 Jonathan D Spratt, Richard Tunstall 52 Development of the thorax 905 Andrew Bush (lungs), Patricia Collins (thoracic walls), Antoon FM Moorman (heart) 53 Chest wall and breast 931 Thomas Collin, Julie Cox Lungs and Diaphragm 54 Pleura, lungs, trachea and bronchi 953 Horia Muresian 55 Diaphragm and phrenic nerves 970 Marios Loukas Heart and Mediastinum 56 Mediastinum 976 Horia Muresian 57 Heart 994 Marios Loukas 58 Great vessels 1024 Marios Loukas Commentaries 7.1 Technical aspects and applications of diagnostic radiology Jonathan D Spratt 7.2 Endobronchial ultrasound Natalie M Cummings Videos Video 52.1 – Animation of the pattern of contraction of the early heart tube Antoon FM Moorman SECTION 8: ABDOMEN AND PELVIS Section Editor (Abdomen): Mark D Stringer Section Editors (Pelvis): Ariana L Smith and Alan J Wein 59 Abdomen and pelvis: overview and surface anatomy 1033 Mark D Stringer, Ariana L Smith, Alan J Wein, Richard Tunstall 60 Development of the peritoneal cavity, gastrointestinal tract and its adnexae 1048 Patricia Collins 61 Anterior abdominal wall 1069 Michael J Rosen, Clayton C Petro, Mark D Stringer 62 Posterior abdominal wall and retroperitoneum 1083 Alexander G Pitman, Donald Moss, Mark D Stringer 63 Peritoneum and peritoneal cavity 1098 Paul H Sugarbaker Gastrointestinal Tract 64 Abdominal oesophagus and stomach 1111 Hugh Barr, L Max Almond 65 Small intestine 1124 Simon M Gabe 66 Large intestine 1136 Peter J Lunniss Abdominal Viscera 67 Liver 1160 J Peter A Lodge 68 Gallbladder and biliary tree 1173 Mark D Stringer 69 Pancreas 1179 Mohamed Rela, Mettu Srinivas Reddy 70 Spleen 1188 Andy Petroianu 71 Suprarenal (adrenal) gland 1194 Nancy Dugal Perrier Urogenital System 72 Development of the urogenital system 1199 Patricia Collins, Girish Jawaheer, Richard M Sharpe 73 True pelvis, pelvic floor and perineum 1221 John OL Delancey 74 Kidney and ureter 1237 Thomas J Guzzo, Drew A Torigian CONTENTS viii 75 Bladder, prostate and urethra 1255 Serge Ginzburg, Anthony T Corcoran, Alexander Kutikov 76 Male reproductive system 1272 Marc Goldstein, Akanksha Mehta 77 Female reproductive system 1288 Lily A Arya, Nadav Schwartz Commentaries 8.1 The neurovascular bundles of the prostate Robert P Myers 8.2 Real-time microscopy of the upper and lower gastrointestinal tract and the hepatobiliary–pancreatic system during endoscopy Martin Götz Videos Video 63.1 – Surgical exploration of the peritoneal cavity Paul H Sugarbaker Video 75.1 – Laparoscopic view of bladder filling and emptying in relation to the rectovesical pouch Video 75.2 – Laparoscopic view of anterior abdominal wall and ligaments Serge Ginzberg, Anthony T Corcoran, Alexander Kutikov SECTION 9: PELVIC GIRDLE AND LOWER LIMB Section Editor: R Shane Tubbs 78 Pelvic girdle and lower limb: overview and surface anatomy 1316 Nihal Apaydin, Richard Tunstall 79 Development of the pelvic girdle and lower limb 1334 Cheryll Tickle 80 Pelvic girdle, gluteal region and thigh 1337 Mohammadali M Shoja 81 Hip 1376 Donald A Neumann 82 Knee 1383 Brion Benninger 83 Leg 1400 Robert J Spinner, Benjamin M Howe 84 Ankle and foot 1418 Anthony V D'Antoni Commentaries 9.1 Nerve biomechanics Kimberly S Topp 9.2 Functional anatomy and biomechanics of the pelvis Andry Vleeming, Frank H Willard 9.3 Articularis genus Stephanie J Woodley Videos Video 78.1 – Lower limb surface anatomy Rolfe Birch Video 84.1 – Ankle block: surface anatomy Dominic Harmon Index 1453 Bonus imaging collection Section 2 2.1 Human oocyte undergoing fertilization, cell division, blastocyst development and hatching in vitro Section 3 3.1 MRI head: axial T2-weighted 3.2 MRI head: coronal T2-weighted 3.3 MRI head: sagittal T2-weighted Section 4 4.1 CT neck: axial post-IV contrast 4.2 CT neck: coronal post-IV contrast Section 7 7.1 CT chest, abdomen and pelvis: axial post-IV contrast 7.2 CT chest, abdomen and pelvis: coronal post-IV contrast 7.3 CT chest, abdomen and pelvis: sagittal post-IV contrast Section 8 8.1 MRI male pelvis: axial T1-weighted Section 9 9.1 MRI male pelvis: coronal T1-weighted Eponyms Historical bibliography References cited in earlier editions, up to and including the thirty-eighth edition ix PREFACE 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, or a consequence, of the anatomy, or where the anatomical features are instrumental in the diagnosis/ treatment/management of the condition. Wherever possible, the photo micrographs illustrate human histology and embryology; nonhuman sources are acknowledged in the captions. In an ideal world, anatomical terminology would satisfy both anat omists and clinicians. For the avoidance of doubt, the same word should be agreed and used for each structure that is described, whether in the anatomy laboratory or the clinic. In the real world, this goal is achieved with varying degrees of success; alternative terms (co)exist and may (and frequently do) confuse or frustrate. Currently, Terminologia Anatomica (TA)1 is the reference source for the terminology for macro scopic anatomy; the text of the fortyfirst edition of Gray's Anatomy is almost entirely TAcompliant. However, where terminology is at vari ance with, or, more likely, is not included in, the TA, the alternative term that is chosen either is cited in the relevant consensus document or position paper – e.g. 'European Position Paper on the Anatomical Terminology of the Internal Nose and Paranasal Sinuses'2 and the Inter national Interdisciplinary Consensus Statement on the 'Nomenclature of the Veins of the Lower Limbs'3 – or enjoys widespread clinical usage: for example, the use of attitudinally appropriate terms in cardiology (see Chapter 57). The continued use of eponyms is contentious.4 Pro ponents of their retention argue that some eponyms are entrenched in medical language and are (therefore) indispensable, that they facilitate communication because their use is so pervasive and that they serve to remind us of the humanism of medicine. Detractors argue that eponyms are inherently inaccurate, nonscientific and often undeserved. In this edition of Gray's Anatomy, synonyms and eponyms are given in paren theses on first usage of a preferred term and not shown thereafter in the text; an updated list of eponyms remains available in the ebook for reference purposes. I offer my sincere thanks to the editorial team at Elsevier, initially under the leadership of Madelene Hyde and latterly of Jeremy Bowes, for their guidance, professionalism, good humour and unfailing support. In particular, I thank Poppy Garraway, Humayra Rahman Khan, Wendy Lee, Joanna Souch, Julie Taylor, Jan Ross and Louise Cook, for being at the end of a phone or available by email whenever I needed advice or support. I dedicate my work on the fortyfirst edition of Gray's Anatomy to the memory of my late husband, Guy Standring. Susan Standring January 2015 'Anatomy is the basis of medical discourse.' (Hippocrates, De locis in homine 2) Looking through an almost complete set of the previous editions of Gray's Anatomy, I am struck by the marked difference in size between the first and fortieth editions. That progressive increase in girth has occurred pari passu with groundbreaking advances in basic science and clinical medicine over the past 155 years. Anatomy has become a far wider discipline than Henry Gray, Henry van Dyke Carter or any of their students could have envisaged. Fields such as cell biology, molecular genetics, neuroanatomy, embryology and bioinformatics either had not emerged or were in their infancy in 1858. Techniques that today inform our view of the internal landscape of the body – such as specialized types of light and electron microscopy; imaging modalities, including Xrays, magnetic resonance imaging, computed tomography and ultra sonography; the use of 'soft' perfusion techniques and frozenthawed, unembalmed cadavers for dissectionbased studies; and the advances in information technology that enable endoscopic and robotic surgery and facilitate minimally invasive access to structures previously consid ered inaccessible – were all unknown. As each development entered mainstream scientific or clinical use, the new perspectives on the body it afforded, whether at submicroscopic or macroscopic level, filtered into the pages of Gray's Anatomy: for example, the introduction of Xray plates (twentyseventh edition, 1938) and electron micrographs (thirty second edition, 1958). In the Preface to the first edition, Henry Gray wrote that 'This Work is intended to furnish the Student and Practitioner with an accurate view of the Anatomy of the Human Body, and more especially the application of this science to Practical Surgery.' We remain true to his intention. An appropri ate knowledge of clinically relevant, evidencebased anatomy is an essential element in the armamentarium of a practising clinician; indeed, 'If anything, the relevance of anatomy in surgery is more impor tant now than at any other time in the past' (Tubbs, in Preface Com mentary, which accompanies this volume). In my Preface to the fortieth edition, I intimated that the book was quite literally in danger of breaking its binding if any more pages were added. In order to avoid this unfortunate occurrence, the fortyfirst edition contains a significant amount of material that is exclusively electronic, in the form of 77,000 words of additional text, 300 artworks and tables, 28 videos and 24 specially invited commentaries on topics as diverse as electron microscopy and fluorescence microscopy; the neurovascular bundles of the prostate; stem cells in regenerative medi cine; the anatomy of facial ageing; and technical aspects and applica tions of diagnostic radiology. In keeping with the expectation that anatomy should be evidencebased, the fortyfirst edition contains many more references in the ebook than could be included in the thirtyninth and fortieth printed editions. Neel Anand, Rolfe Birch, Pat Collins, Alan Crossman, Michael Gleeson, Ariana Smith, Jonathan Spratt, Mark Stringer, Shane Tubbs, Alan Wein and Caroline Wigley brought a wealth of scholarship and experience as anatomists, cell biologists and clinicians to their roles as Section Editors. I thank them for their dedication and enthusiastic support, in selecting and interacting with the authors in their Sections and for meeting deadlines, despite the everincreasing demands on their time from university and/or hospital managers. Pat Collins, Girish Jawaheer, Richard Tunstall and Caroline Wigley worked closely with many authors to update the text and artworks for organogenesis, paediatric anatomy, evidencebased surface anatomy and microstruc ture, respectively, across Sections 3 to 9. Jonathan Spratt acted as both a Section Editor (thorax) and an indefatigable 'go to' for sourcing images throughout the book; in the latter capacity, he has produced a superb collection of additional labelled images, available in the ebook (see Bonus imaging collection). Over a hundred highly experi enced anatomists and clinicians contributed text, often extensively revised from the previous edition, and/or artworks, original micro graphs or other images to individual chapters. 1Terminologia Anatomica (1998) is the joint creation of the Federative Committee on Anatomical Terminology (FCAT) and the Member Associations of the Interna tional Federation of Associations of Anatomists (IFAA). 2Lund VJ, Stammberger H, Fokkens WJ et al 2014 European position paper on the anatomical terminology of the internal nose and paranasal sinuses. Rhinol Suppl 24:1–34. 3Caggiati A, Bergan JJ, Gloviczki P et al; International Interdisciplinary Consensus Committee on Venous Anatomical Terminology 2005 Nomenclature of the veins of the lower limb: extensions, refinements, and clinical application. J Vasc Surg 41:719–24. 4Amarnani A, Brodell RT, Mostow EN 2013 Finding the evidence with eponyms. JAMA Dermatol 149:664–5; Fargen KM, Hoh BL 2014 The debate over eponyms. Clin Anat 27:1137–40; Lo WB, Ellis H 2010 The circle before Willis: a historical account of the intracranial anastomosis. Neurosurgery 66:7–18; Ma L, Chung KC 2012 In defense of eponyms. Plast Reconstr Surg 129:896e–8e. e1 The continuing relevance of anatomy in current surgical practice and research PREFACE COMMENTARY When our anatomy forebears embarked on the uncharted study of the human body, they did so without reference. Their focus was to chart and map the body simply to learn and describe intricacies never chroni- cled before. The anatomical 'map' we use today came about thanks to figures such as da Vinci, Vesalius, Cheselden and, more recently, Henry Gray. On the shoulders of these giants, we see farther than our predeces- sors. In The Metalogicon, published in 1159, John Salisbury recognized the profound observation of French philosopher Bernard of Chartres, who declared that '...we are like dwarfs on the shoulders of giants, so that we can see more than they, and things at a greater distance, not by virtue of any sharpness of sight on our part, or any physical distinction, but because we are carried high and raised up by their giant size'. So, with the gross anatomy of man presumed, by many scholars, to have been described and understood long ago, how does the modern anatomist bring rel- evance to the continued study of morphology? Is there any uncharted territory for the modern anatomist to plot in order to sustain our field of study and for it to continue to be perceived as relevant to an educa- tional world, and to medical and dental curricula in which the time allotted to anatomical study has significantly waned? Simply put, yes. Henry Gray, based on the title of his original text, Anatomy, Descriptive and Surgical, knew very well that there was a need to refocus the lenses of teaching and research in the anatomical sciences, and to expand and explore their surgical relevance. Our gross anatomical map of the human body must continue to be updated and legends must continue to be placed on that map to incorporate modern advances in technol- ogy. New methods of surgery, such as laparoscopy and endoscopy, as well as the use of the surgical microscope, offer the opportunity to view the human form in a different light and in greater surgical detail than ever before. If anything, the relevance of anatomy in surgery is more important now than at any other time in the past. The modern surgeon must take what is learned macroscopically, in the dissection room, and apply this knowledge to structures seen under magnification and through instruments that provide a surgical field that is, at times, just millimetres in diameter. Therefore, attention to anatomical detail is of vital importance as references and anatomical landmarks are mini- mized in the surgical theatre of the new millennium. As mentioned before, early anatomists dissected with curiosity about the unknown and gained knowledge that would become a prerequisite for proper surgical manœuvres. Today, as anatomists, our anatomical knowledge should create in us a curiosity about what we can do with the knowledge that we have gained. The ability to apply that knowledge offers an opportunity to be an integral part of the ever-progressing field of surgery. For example, today, surgical problems are often the impetus for dissection studies, which can influence the way in which surgery is performed and, moreover, can sway the way in which anatomy is taught (e.g. redefining a focus in condensed curricula and with decreased work hours for house officers). Surgically, dissection studies have allowed us to manipulate known human anatomy and to solve, for example, complex neurological problems. As an illustration of the surgical rele- vance of modern-day anatomical studies for neurological pathologies, we have conducted, in my laboratory, cadaveric feasibility studies that suggested that the phrenic nerve could be reinnervated in high quadri- plegic patients who are ventilator-dependent (a morbid condition with an associated high mortality rate) by using the intact, adjacent accessory nerve (i.e. neurotization) (Tubbs et al 2008a) (Fig. 1.6.1). The theory behind this investigation was that the functioning accessory nerve would be used to form a new circuit between it and the dysfunctional phrenic nerve, and that this would allow recovery of diaphragm func- tion. For this technique, a longitudinal incision was made along the lower half of the posterior border of sternocleidomastoid. Dissection was then performed in order to identify both the accessory nerve at this level, at its entrance into trapezius, and the phrenic nerve crossing anterior to scalenus anterior. The medial half of the accessory nerve was then split away from its lateral half and transected at its entrance into muscle. This distally disconnected medial half of the nerve was then swung medially to the phrenic nerve, which had been transected proxi- mally. The two nerves were then sutured together without tension. This 'rearranging' of human anatomy has now been employed clinically with success. Yang et al (2011) used our study results to treat a 44-year-old man with complete spinal cord injury at the C2 level. Clinically, left diaphragm activity was decreased and the right diaphragm was com- pletely paralysed. Four weeks after surgery, training of the synchronous activities of trapezius and inspiration was conducted. Six months after surgery, motion was observed in the previously paralysed right dia- phragm. Evaluation of lung function indicated improvements in vital capacity and tidal volume. The patient was able to sit in a wheelchair and conduct activities without assisted ventilation 12 months after surgery. For the surgeon, such manipulation of anatomy requires a comprehensive understanding not only of normal anatomy but also of what might occur functionally by rewiring such nerves. For example, patients undergoing this surgery will initially need to think of moving their trapezius to activate their diaphragm. With time, this will not be the case. Similar illustrations of the plasticity of the brain have been seen in patients undergoing hypoglossal to facial nerve neurotization procedures; these patients at first need to think of moving their tongue in order for their facial muscles to contract. Rewiring of nerves has been addressed in other studies. Thus, we have shown, first in a cadaveric study (Hansasuta et al 2001) and then clinically (Wellons et al 2009), that the medial pectoral nerve can be sectioned near its entrance into the deep surface of pectoralis major and swung round and sewn into the musculocutaneous nerve (Fig. 1.6.2). If this procedure is successful, axonal regrowth from the medial pectoral nerve into the musculocutaneous nerve (about 1 mm/day) will re-establish function in the anterior arm muscles; the loss of clinically significant function of the dually innervated pectoralis major is minimal and the functional gain of having the anterior arm muscles work is significant (Wellons et al 2009). Being able to bring the hand to the mouth and feed oneself is a task that most take for granted. In children with birth-related injuries to the upper brachial plexus (i.e. Erb's palsy), this movement is often the difference between waiting to be fed or feeding oneself. This method has been used at our institution for over 15 years with an 80% success rate, where success is measured as the patient regaining function of arm flexion. Another example of what we have termed 'reverse translational research in anatomy' (i.e. from the bed to the bench and back) is the location of new anatomical diversionary sites (in this case, the medul- lary cavity of the ilium) that could be used in patients with cerebrospi- nal fluid absorption problems (i.e. hydrocephalus) and in whom the traditionally used receptacles for absorbing this diverted cerebrospinal fluid (e.g. peritoneal and pleural cavities, heart) are not options, as a consequence of e.g. malabsorption or local infection (Tubbs et al 2015) (Fig. 1.6.3). This alternative site has, for the first time, just been used and with success (unpublished data). Although not proven clinically, an earlier study in primates showed that the manubrium of the sternum could also be used as a distal receptacle for cerebrospinal fluid collec- tion (Tubbs et al 2011). After tubing was tunnelled from the cannulated ventricle, the distal tubing was inserted subcutaneously into the supe- rior aspect of the midline manubrium, where a small hole had been drilled. Up to 50 ml of saline per hour could be infused into the primate sternum without vital sign changes. This study, and the study using the ilium as a depository, both demonstrate the anatomical continuity between the bony medullary cavities and the vascular system. Such positive effects on patient outcomes not only make the study of human anatomy from a slanted perspective extremely gratifying, but are also practical since the results have direct application in the surgical theatre. In addition to surgical anatomy playing a role in new uses of the normal anatomy, this field can also explore and direct new surgical approaches where the goals are to make surgery more effective and R Shane Tubbs The conTinuing relevance of anaTomy in currenT surgical pracTice and research e2 treatment, resulted in a more limited laminectomy and myelotomy, and, in one case, assisted in identifying a residual spinal cord tumour. It was also useful in the fenestration of a multilevel spinal arachnoid cyst and in confirming communication of fluid spaces in the setting of a complex holocord syrinx. Endoscopy aided the visualization of the spinal cord to ensure the absence of tethering in the case of split spinal cord malformation. These endoscopic approaches were only possible by knowing the normal anatomy and how it appears in a confined field of view, as first seen in the anatomy laboratory. Lastly, the anatomist can add to the relevance of anatomy for the surgeon with studies that have an impact on the identification or avoid- ance of important structures during operative manœuvres (i.e. anatomi- cal landmark studies). My group has defined surgical landmarks for anatomical structures such as the superior and inferior gluteal nerves (Apaydin et al 2013, Apaydin et al 2009); vein of Labbé (Tubbs et al 2012); sigmoid sinus (Tubbs et al 2009a); amygdala (Tubbs et al minimally invasive, and involve fewer complications. For example, we have performed feasibility studies looking at a wide range of novel approaches that might be used by the surgeon. These include a dorsal approach to the carpal tunnel for an entrapped median nerve (Tubbs et al 2005a); an anterior approach to the sciatic nerve potentially com- pressed by piriformis via the obturator foramen (Tubbs, unpublished data); an anterior approach to the upper thoracic vertebrae for spine fusion procedures (Tubbs et al 2010a); an intra-abdominal laparoscopic approach to decompress the pudendal nerve (Loukas et al 2008); and midline endoscopic approaches to the fourth ventricle with application to decompressing a 'trapped' fourth ventricle, as is seen in some cases of hydrocephalus (Tubbs et al 2004). We have also explored the feasibil- ity in cadavers of using endoscopy for exploration of pathologies of the thecal sac (Chern et al 2011). In a series of children with intraspinal pathology (arachnoid cyst, spinal cord tumour, holocord syrinx and split cord malformation), intradural spinal endoscopy was a useful Fig. 1.6.1 A schematic representation of the anatomically defined technique of using the accessory nerve for neurotization of the phrenic nerve with application to patients with high cervical quadriplegia who are ventilator- dependent. With nerve regrowth, axons from the intact and functioning accessory nerve travel into the phrenic nerve to reinnervate this nerve and restore diaphragmatic function. In this example, only one-half of the accessory nerve is used in order to maintain some function of trapezius. (Drawn by Mr David Fisher.) Fig. 1.6.2 The neurotization of the musculocutaneous nerve with the medial pectoral nerve (inset). Similar to the example illustrated in Figure 1.6.1, such a method of nerve repair is employed in the hope that a patient with an upper brachial plexus injury and anterior arm muscles that are dysfunctional can regain function by regrowth of axons from the intact medial pectoral nerve into and along the musculocutaneous nerve. (Drawn by Mr David Fisher.) The continuing relevance of anatomy in current surgical practice and research e3 (Loukas et al 2006); long thoracic nerve (Tubbs et al 2006b); anterior interosseous nerve (Tubbs et al 2006c); accessory nerve (Tubbs et al 2005b); lumbar plexus and its branches (Tubbs et al 2005c); trochlear nerve (Tubbs and Oakes 1998); and frontal sinus (Tubbs et al 2002). Such studies might assist in decreasing the morbidity and increasing the efficiency of surgical approaches and certainly illustrate the surgical relevance of anatomy. Moreover, this list exemplifies the multitude of anatomical structures that may be given greater surgical relevance by addressing how they may be more accurately located in the operating theatre. In this day and age, if anatomists are not to lose their footing and simply be considered teachers of an old and outdated discipline, the onus is on us to renew interest in our field with timely and salient studies that gird the loins of a profession that is in danger of becoming extinct. It is my opinion, and that of others, that one effective way to achieve this is to remind the world by demonstrations such as those listed here that the study of anatomy is as clinically relevant today as it was at its humble beginnings. Considering the adage that anatomy is the oldest child of Mother Medicine, the fact that surgical problems and anatomical studies go hand in hand is obvious – anatomical research is not a 'dead' science! The modern relevance of anatomy to surgical practice and research must not be underestimated. Fig. 1.6.3 The technique used in a patient with hydrocephalus to divert cerebrospinal fluid from the cerebral ventricles to the ilium. The enlarged ventricles are cannulated with a catheter connected to a subcutaneous valve that drains into tubing tunnelled under the skin and then implanted into the medullary cavity of the ilium; here, the cerebrospinal fluid is absorbed into the vascular system. The techniques described in Figures 1.6.2 and 1.6.3, based on surgical problems and manipulation of known anatomy for surgical benefit, were evaluated and studied in the anatomy laboratory, and have now been used clinically. (Drawn by Mr David Fisher.) Fig. 1.6.4 A superior view of the cranium, with the underlying superior sagittal sinus, cortical veins and lateral lacunae illustrated. This study explored the relationship between the underlying lateral lacunae and the overlying coronal and sagittal sutures, and made measurements between these structures. Neurosurgically, the initial placement of burr-holes avoids the midline in order to prevent damage to the superior sagittal sinus. However, the intracranial entrance of the drill often injures more laterally placed lacunae. Using surface anatomy based on anatomical landmarks, a neurosurgeon can be more aware of the locations of these underlying structures while performing craniotomies. Such landmarks have now been used by neurosurgeons at our institution. (Drawn by Mr David Fisher.) 2010b); buccal branch of the trigeminal nerve (Tubbs et al 2010c); radial nerve and posterior interosseous branch (Cox et al 2010, Tubbs et al 2006a); perineal branch of the posterior femoral cutaneous nerve (Tubbs et al 2009b); lateral lacunae (Tubbs et al 2008b) (Fig. 1.6.4); basal vein of Rosenthal (Tubbs et al 2007); greater occipital nerve REFERENCES Apaydin N, Bozkurt M, Loukas M et al 2009 The course of the inferior gluteal nerve and surgical landmarks for its localization during posterior approaches to hip. Surg Radiol Anat 31:415-18. Apaydin N, Kendir S, Loukas M et al 2013 Surgical anatomy of the superior gluteal nerve and landmarks for its localization during minimally inva- sive approaches to the hip. Clin Anat 26:614–20. The conTinuing relevance of anaTomy in currenT surgical pracTice and research e4 Tubbs RS, Miller JH, Cohen-Gadol AA et al 2010b Intraoperative anatomic landmarks for resection of the amygdala during medial temporal lobe surgery. Neurosurgery 66:974–7. Tubbs RS, Miller J, Loukas M et al 2009b Surgical and anatomical landmarks for the perineal branch of the posterior femoral cutaneous nerve: impli- cations in perineal pain syndromes. Laboratory investigation. J Neuro- surg 111:332–5. Tubbs RS, Oakes WJ 1998 Relationships of the cisternal segment of the trochlear nerve. J Neurosurg 89:1015–19. Tubbs RS, Pearson B, Loukas M 2008a Phrenic nerve neurotization utilizing the spinal accessory nerve: technical note with potential application in patients with high cervical quadriplegia. Childs Nerv Syst 24:1341–4. Tubbs RS, Salter EG, Custis JW et al 2006b Surgical anatomy of the cervical and infraclavicular parts of the long thoracic nerve. J Neurosurg 104: 792–5. Tubbs RS, Salter EG, Sheetz J et al 2005a Novel surgical approach to the carpal tunnel: cadaveric feasibility study. Clin Anat 18:350–6. Tubbs RS, Salter EG, Wellons JC 3rd et al 2005b Superficial landmarks for the spinal accessory nerve within the posterior cervical triangle. J Neu- rosurg Spine 3:375–8. Tubbs RS, Salter EG, Wellons JC 3rd et al 2005c Anatomical landmarks for the lumbar plexus on the posterior abdominal wall. J Neurosurg Spine 2:335–8. Tubbs RS, Salter EG, Wellons JC 3rd et al 2006a Superficial surgical land- marks for identifying the posterior interosseous nerve. J Neurosurg 104:796–9. Tubbs RS, Tubbs I, Loukas M et al 2015 Ventriculoiliac shunt: a cadaveric feasibility study. J Neurosurg Pediatr 15:310–12. Tubbs RS, Wellons JC 3rd, Salter G et al 2004 Fenestration of the superior medullary velum as treatment for a trapped fourth ventricle: a feasibility study. Clin Anat 17:82–7. Wellons JC, Tubbs RS, Pugh JA et al 2009 Medial pectoral nerve to muscu- locutaneous nerve neurotization for the treatment of persistent birth- related brachial plexus palsy: an 11-year institutional experience. J Neurosurg Pediatr 3:348–53. Yang ML, Li JJ, Zhang SC 2011 Functional restoration of the paralyzed dia- phragm in high cervical quadriplegia via phrenic nerve neurotization utilizing the functional spinal accessory nerve. J Neurosurg Spine 15: 190–4. Chern JJ, Gordon AS, Naftel RP et al 2011 Intradural spinal endoscopy in children. J Neurosurg Pediatr 8:107–11. Cox CL, Riherd D, Tubbs RS et al 2010 Predicting radial nerve location using palpable landmarks. Clin Anat 23:420–6. Hansasuta A, Tubbs RS, Grabb PA 2001 Surgical relationship of the medial pectoral nerve to the musculocutaneous nerve: a cadaveric study. Neuro- surgery 48:203–6. Loukas M, El-Sedfy A, Tubbs RS et al 2006 Identification of greater occipital nerve landmarks for the treatment of occipital neuralgia. Folia Morphol (Warsz) 65:337–42. Loukas M, Louis RG Jr, Tubbs RS et al 2008 Intra-abdominal laparoscopic pudendal canal decompression – a feasibility study. Surg Endosc 22: 1525–32. Tubbs RS, Bauer D, Chambers MR 2011 A novel method for cerebrospinal fluid diversion: a cadaveric and animal study. Neurosurgery 68:491–4. Tubbs RS, Custis JW, Salter EG et al 2006c Quantitation of and superficial surgical landmarks for the anterior interosseous nerve. J Neurosurg 104:787–91. Tubbs RS, Elton S, Salter G et al 2002 Superficial surgical landmarks for the frontal sinus. J Neurosurg 96:320–2. Tubbs RS, Johnson PC, Loukas M et al 2010c Anatomical landmarks for localizing the buccal branch of the trigeminal nerve on the face. Surg Radiol Anat 3:933–5. Tubbs RS, Louis RG Jr, Song YB et al 2012 External landmarks for identifying the drainage site of the vein of Labbé: application to neurosurgical procedures. Br J Neurosurg 26:383–5. Tubbs RS, Loukas M, Callahan JD et al 2010a A novel approach to the upper anterior thoracic spine: a cadaveric feasibility study. J Neurosurg Spine 13:346–50. Tubbs RS, Loukas M, Louis RG Jr et al 2007 Surgical anatomy and landmarks for the basal vein of Rosenthal. J Neurosurg 106:900–2. Tubbs RS, Loukas M, Shoja MM et al 2008b Lateral lakes of Trolard: anatomy, quantitation, and surgical landmarks. Laboratory investigation. J Neu- rosurg 108:1005–9. Tubbs RS, Loukas M, Shoja MM et al 2009a Surface landmarks for the junc- tion between the transverse and sigmoid sinuses: application of the 'strategic' burr hole for suboccipital craniotomy. Neurosurgery 65: 37–41. x 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 RL, Vogl AW, Mitchell A (eds), Gray's Anatomy for Students, 2nd ed. Elsevier, Churchill Livingstone. Copyright 2010. Drake RL, Vogl AW, Mitchell A, Tibbitts R, Richardson P (eds), Gray's Atlas of Anatomy. Elsevier, Churchill Livingstone. Copyright 2008. Waschke J, Paulsen F (eds), Sobotta Atlas of Human Anatomy, 15th ed. Elsevier, Urban & Fischer. Copyright 2013. Acknowledgements for paediatric anatomy content in chapter 45 to Ritchie Marcus, MD and Guirish A. Solanki, MD, Birmingham Children's Hospital, UK, and for chapter 81 to Christopher Edward Bache, MBChB, FRCS (Tr & Orth), Birmingham, UK. The editors would like to thank all contributors and illustrators to the previous editions of Gray's Anatomy, including the fortieth and thirty-ninth editions. 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, and sometimes being used as the foundation for figures that are new to this edition. xi CONTRIBUTORS TO THE FORTY-FIRST EDITION Graham J Burton MD, DSc, FMedSci Mary Marshall and Arthur Walton Professor of the Physiology of Reproduction Centre for Trophoblast Research University of Cambridge Cambridge, UK Andrew Bush MD, FRCP, FRCPCH, FERS Professor of Paediatrics and Head of Section (Paediatrics) Imperial College; Professor of Paediatric Respirology National Heart and Lung Institute; Consultant Paediatric Chest Physician Royal Brompton and Harefield NHS Foundation Trust Paediatric Respiratory Medicine London, UK Alison Campbell BSc(Hons), MMedSci, DipRCPath Group Director of Embryology CARE Fertility Nottingham, UK Bodo EA Christ MD Professor and Former Chairman Department of Molecular Embryology University of Freiburg Freiburg, Germany Thomas Collin MBBS, FRCS(Plast) Consultant Plastic and Reconstructive Surgeon University Hospital of North Durham Department of Plastic Surgery Durham, UK Patricia Collins BSc, PhD, FHEA Professor of Anatomy Anglo-European College of Chiropractic Bournemouth, UK; Editor for Embryology and Development Anthony T Corcoran MD Assistant Professor of Urologic Oncology and Minimally Invasive Surgery Department of Urology SUNY Stony Brook School of Medicine Stony Brook, NY, USA Julie Cox FRCS(Eng), FRCR Consultant Radiologist City Hospitals Sunderland NHS Foundation Trust Sunderland, UK Alan R Crossman BSc, PhD, DSc Professor Emeritus University of Manchester Manchester, UK Michael A Adams BSc, PhD Professor of Biomechanics Centre for Comparative and Clinical Anatomy University of Bristol, UK L Max Almond MB, ChB, MRCS, MD Senior Registrar in Gastrointestinal Surgery West Midlands Deanery Birmingham, UK Neel Anand MD Clinical Professor of Surgery Director, Spine Trauma, Minimally Invasive Spine Surgery Spine Center Cedars Sinai Medical Center Los Angeles, CA, USA Nihal Apaydin MD, PhD Associate Professor of Anatomy Department of Anatomy and Brain Research Center Ankara University Faculty of Medicine Ankara, Turkey Lily A Arya MD, MS Associate Professor of Obstetrics and Gynecology Perelman School of Medicine University of Pennsylvania Department of Obstetrics and Gynecology Philadelphia, PA, USA Tipu Aziz FMedSci Professor of Neurosurgery John Radcliffe Hospital University of Oxford Oxford, UK Jonathan BL Bard MA, PhD Emeritus Professor of Development and Bioinformatics School of Biomedical Sciences University of Edinburgh Edinburgh, UK Eli M Baron MD Clinical Associate Professor of Neurosurgery Spine Surgeon, Cedars Sinai Department of Neurosurgery Cedars Sinai Spine Center, Cedars Sinai Medical Center Los Angeles, CA, USA Hugh Barr MD(Dist), ChM, FRCS(Eng), FRCS(Ed), FHEA, FODI Consultant General and Gastrointestinal Surgeon Oesophagogastric Resection Unit Gloucestershire Royal Hospital Gloucester, UK Brion Benninger MD, MSc Professor, Executive Director Medical Anatomy Center – Innovation and Technology Research McDaniel Surgical, Radiological & Education Research Lab Departments of Medical Anatomical Sciences & Neuromuscular Medicine Western University of Health Sciences, Lebanon, Oregon Faculty Orthopaedics & Surgical Residency Training Faculty Sports Medicine Fellowship Training Samaritan Health Services, Corvallis, Oregon USA Barry KB Berkovitz BDS, MSc, PhD, FDS, LDSRCS(Eng) Emeritus Reader in Dental Anatomy Anatomy Department King's College London London, UK; Visiting Professor Oman Dental College Oman Leela C Biant BSc(Hons), MBBS, AFRCSEd, FRCSEd(Tr & Orth), MSres(Lond), MFSTEd Consultant Trauma and Orthopaedic Surgeon Royal Infirmary of Edinburgh; Honorary Senior Lecturer University of Edinburgh NRS Career Clinician Scientist Fellow Edinburgh, UK Rolfe Birch MChir, FRCPS(Glasg), FRCS(Ed), FRCS(Eng) Retired Consultant in Charge War Nerve Injury Clinic, Defence Medical Rehabilitation Centre, Surrey; Retired Head, Peripheral Nerve Injury Unit, Royal National Orthopaedic Hospital; Professor in Neurological Orthopaedic Surgery, University College of London London, UK Martin A Birchall MD, FRCS, FMedSci Professor of Laryngology Consultant Otolaryngologist, Ear Institute University College London and Royal National Throat Nose and Ear Hospital University College Hospitals NHS Foundation Trust London, UK Sue Black OBE, BSc, PhD, DSc, FRSE, FRAI, FRCP, FSB Professor of Anatomy and Forensic Anthropology Centre for Anatomy and Human Identification University of Dundee Scotland, UK The editors would like to acknowledge and offer grateful thanks for the input of all previous editions' contributors, without whom this new edition would not have been possible. Contributors to the forty-first edition xii Natalie M Cummings BSc(Med Sci), MB ChB, MPhil, MD, MRCP(Ed) Consultant Respiratory Physician University Hospital of North Durham Durham, UK Anthony V D'Antoni MS, DC, PhD Clinical Professor and Director of Anatomy Department of Pathobiology Sophie Davis School of Biomedical Education City University of New York; Adjunct Associate Professor Division of Pre-Clinical Sciences and Department of Surgery New York College of Podiatric Medicine New York, NY, USA Paolo De Coppi MD, PhD Professor of Paediatric Surgery; Head of Stem Cells and Regenerative Medicine; Consultant Paediatric Surgeon Great Ormond Street Hospital UCL Institute of Child Health London, UK John OL DeLancey MD Norman F Miller Professor of Gynecology Department of Obstetrics and Gynecology Professor, Department of Urology University of Michigan Medical School Ann Arbor, MI, USA Ronald H Douglas BSc, PhD Professor of Visual Science Division of Optometry and Visual Science School of Health Sciences City University London London, UK Barrie T Evans BDS(Hons), MB BCh, FRCS(Eng), FRCS(Ed), FDSRCS(Eng), FFDRCS(Ire) Consultant Oral and Maxillofacial Surgeon Southampton University Hospitals; Honorary Senior Lecturer in Surgery to Southampton University Medical School; Civilian Consultant Advisor in Oral and Maxillofacial Surgery to the Royal Navy; Past President, British Association of Oral and Maxillofacial Surgeons Southampton, UK Juan C Fernandez-Miranda MD Associate Professor of Neurological Surgery; Associate Director, Center for Cranial Base Surgery; Director, Surgical Neuroanatomy Laboratory University of Pittsburgh Medical Center Pittsburgh, PA, USA Jonathan M Fishman BM BCh(Oxon), MA(Cantab), MRCS(Eng), DOHNS, PhD Clinical Lecturer University College London London, UK Roland A Fleck PhD, FRCPath, FRMS Reader and Director, Centre for Ultrastructural Imaging King's College London London, UK David N Furness BSc, PhD Professor of Cellular Neuroscience School of Life Sciences Keele University Newcastle-under-Lyme, UK Simon M Gabe MD, MSc, BSc(Hons), MBBS, FRCP Consultant Gastroenterologist and Honorary Senior Lecturer; Co-Chair of the Lennard-Jones Intestinal Failure Unit, St Mark's Hospital Middlesex, UK Andrew JT George MA, PhD, DSc, FRCPath, FSB Deputy Vice Chancellor (Education and International) Professor of Immunology Brunel University London, UK Serge Ginzburg MD Assistant Professor of Urologic Oncology Division of Urology Fox Chase Cancer Center; Department of Urology Albert Einstein Medical Center Philadelphia, PA, USA Michael Gleeson MD, FRCS, FRACS Hons, FDS Hons Professor of Skull Base Surgery University College London The National Hospital for Neurology and Neurosurgery London, UK Marc Goldstein MD, DSc(Hon), FACS Matthew P Hardy Distinguished Professor of Reproductive Medicine and Urology; Surgeon-in-Chief, Male Reproductive Medicine and Surgery Cornell Institute for Reproductive Medicine and Department of Urology Weill Cornell Medical Center; Adjunct Senior Scientist, Population Council, Center for Biomedical Research New York, NY, USA Martin Götz MD, PhD Professor, Interdisciplinary Endoscopy Universitätsklinikum Tübingen Tübingen, Germany Anthony Graham BSc, PhD Professor of Developmental Biology MRC Centre for Developmental Neurobiology King's College London London, UK Leonard P Griffiths MB ChB, MRCP(UK) Registrar in Gastroenterology and General Internal Medicine Royal United Hospital Bath; Clinical Research Fellow University of Bath Bath, UK Paul D Griffiths PhD, FRCR, FMedSci Professor of Radiology, Academic Unit of Radiology University of Sheffield Sheffield, UK Thomas J Guzzo MD, MPH Vice-Chief of Urology Assistant Professor of Urology Perelman School of Medicine University of Pennsylvania Philadelphia, PA, USA Duane E Haines PhD, FAAAS, FAAA Professor, Department of Neurobiology and Anatomy; Professor, Department of Neurology Wake Forest School of Medicine Winston-Salem, NC; Professor Emeritus, University of Mississippi Medical Center Jackson, MS, USA Peter A Helliwell FIBMS, Cert BA, Cert Ed Head Biomedical Scientist Department of Cellular Pathology Royal Cornwall Hospitals Trust Truro, UK Simon Holmes BDS, MBBS, FDS, RCS, FRCS Professor of Craniofacial Traumatology Department of Oral and Maxillofacial Surgery Royal London Hospital, Queen Mary University of London London, UK Claire Hopkins MA (Oxon), FRCS (ORLHNS), DM Consultant Ear, Nose and Throat Surgeon Guy's and St Thomas' Hospitals; Reader in ENT King's College London London, UK Benjamin M Howe MD Assistant Professor of Radiology Mayo Clinic Rochester, MN, USA Daisuke Izawa PhD Assistant Professor, Laboratory of Chromosome Dynamics Institute of Molecular and Cellular Biosciences University of Tokyo Tokyo, Japan Eric Jauniaux MD, PhD, FRCOG Professor in Obstetrics and Fetal Medicine Academic Department of Obstetrics and Gynaecology UCL EGA Institute for Women's Health University College London London, UK Girish Jawaheer MD, FRCS(Eng), FRCS(Paed) Consultant Paediatric Surgeon Great North Children's Hospital, Royal Victoria Infirmary Newcastle upon Tyne NHS Foundation Trust Newcastle upon Tyne, UK; Formerly Specialty Tutor for Paediatric Surgery Royal College of Surgeons of England London, UK; Editor for Paediatric Anatomy Marianne Juhler MD, DMSc Consultant Neurosurgeon Copenhagen University Hospital; Professor of Neurosurgery University Clinic of Neurosurgery Copenhagen, Denmark Helmut Kettenmann PhD Professor, Charité Universitätsmedizin Berlin Max Delbrück Center for Molecular Medicine in the Helmholtz Society Berlin, Germany Contributors to the forty-first edition xiii Abraham L Kierszenbaum MD, PhD Medical (Clinical) Professor Emeritus The Sophie Davis School of Biomedical Education The City University of New York New York, NY, USA Alexander Kutikov MD, FACS Associate Professor of Urologic Oncology Department of Surgical Oncology Fox Chase Cancer Center, Temple University Health System Philadelphia, PA, USA Joey E Lai-Cheong BMedSci(Hons), MBBS, PhD, MRCP(UK) Consultant Dermatologist King Edward VII Hospital (Frimley Health NHS Foundation Trust) Windsor, UK Simon M Lambert BSc, MBBS, FRCS, FRCS(Ed) (Orth) Consultant Orthopaedic Surgeon Shoulder and Elbow Service Royal National Orthopaedic Hospital Trust Stanmore, Middlesex; Honorary Senior Lecturer Institute of Orthopaedics and Musculoskeletal Science University College London London, UK John G Lawrenson MSc(Oxon), PhD, FCOptom Professor of Clinical Visual Science Division of Optometry and Visual Science City University London London, UK Nir Lipsman MD, PhD Neurosurgery Resident University of Toronto Toronto, ON, Canada J Peter A Lodge MD, FRCS Professor of Surgery Hepatobiliary and Transplant Unit St James's University Hospital Leeds, UK Marios Loukas MD, PhD Professor, Department of Anatomical Sciences Dean of Basic Sciences St George's University Grenada, West Indies Andres M Lozano MD, PhD, FRCSC, FRSC, FCAHS Professor and Chairman, Dan Family Chair in Neurosurgery University of Toronto Department of Neurosurgery Toronto Western Hospital Toronto, ON, Canada Ellen A Lumpkin PhD Associate Professor of Somatosensory Biology Columbia University College of Physicians and Surgeons Departments of Dermatology and of Physiology and Cellular Biophysics New York, NY, USA Peter J Lunniss BSc, MS, FRCS Retired Senior Lecturer Academic Surgical Unit, St Bartholomew's and The London Medical College, Queen Mary University London; Retired Honorary Consultant Colorectal Surgeon Royal London and Homerton Hospitals London, UK the late Joseph Mathew MBBS, FMCPath, FRCPath, CertTLHE, PGCE, CertBusStud, FHEA Consultant in Histopathology Department of Histopathology Royal Cornwall Hospitals Trust Truro, UK John A McGrath MD, FRCP, FMedSci Professor of Molecular Dermatology St John's Institute of Dermatology King's College London London, UK Stephen McHanwell BSc, PhD, FHEA, FLS, CBiol FSB, NTF Professor of Anatomical Sciences School of Medical Education and School of Dental Sciences Faculty of Medical Sciences Newcastle University Newcastle upon Tyne, UK Akanksha Mehta MD Assistant Professor of Urology, Emory University School of Medicine Atlanta, GA, USA Bryan C Mendelson FRCS(Ed), FRACS, FACS Head of Faculty Melbourne Advanced Facial Anatomy Course; Private Practitioner, Centre for Facial Plastic Surgery Melbourne, VIC, Australia Zoltán Molnár MD, DPhil Professor of Developmental Neuroscience Department of Physiology, Anatomy and Genetics University of Oxford Oxford, UK Antoon FM Moorman MD, PhD Professor of Embryology and Molecular Biology of Cardiovascular Diseases Department of Anatomy, Embryology and Physiology University of Amsterdam, Academic Medical Center Amsterdam, The Netherlands Gillian M Morriss-Kay DSc Emeritus Professor of Developmental Anatomy Department of Physiology, Anatomy and Genetics University of Oxford Oxford, UK Donald Moss MB, BS, FRACS, FACS Consultant Urologist Ballarat, VIC, Australia Horia Muresian MD, PhD Head of Cardiovascular Surgery Department University Hospital of Bucharest Bucharest, Romania; Visiting Professor, St George's University School of Medicine Grenada, West Indies Robert P Myers MD, MS, FACS Professor Emeritus Department of Urology Mayo Clinic Rochester, MN, USA Donald A Neumann PT, PhD, FAPTA Professor of Physical Therapy Marquette University Milwaukee, WI, USA Dylan Myers Owen PhD Lecturer in Experimental Biophysics Department of Physics and Randall Division of Cell and Molecular Biophysics King's College London London, UK Erlick AC Pereira MA(Camb), DM(Oxf), FRCS(Eng), FRCS(NeuroSurg), MBPsS, SFHEA Senior Clinical Fellow in Complex Spinal Surgery Guy's and St Thomas' Hospitals National Hospital of Neurology and Neurosurgery London, UK Nancy Dugal Perrier MD, FACS Professor, Anderson Cancer Center Department of Surgical Oncology Houston, TX, USA Clayton C Petro MD General Surgery Resident; Allen Research Scholar Department of General Surgery University Hospitals Case Medical Center Cleveland, OH, USA Andy Petroianu MD, PhD Professor of Surgery Department of Surgery School of Medicine of the Federal University of Minas Gerais Belo Horizonte, Minas Gerais, Brazil Jonathon Pines PhD, FMedSci Director of Research in Cell Division University of Cambridge Cambridge, UK Alexander G Pitman BMedSci, MBBS, MMed(Rad), FRANZCR, FAANMS Professorial Fellow Department of Anatomy and Neuroscience University of Melbourne Parkville, VIC, Australia Y Raja Rampersaud MD, FRCSC Associate Professor, Division of Orthopaedic Surgery and Neurosurgery Department of Surgery University of Toronto Toronto, ON, Canada Contributors to the forty-first edition xiv Mettu Srinivas Reddy MS, FRCS, PhD Consultant Surgeon Institute of Liver Disease and Transplantation Global Health City Chennai, India Mohamed Rela MS, FRCS, DSc Director, Institute of Liver Disease and Transplantation Global Health City, Chennai, India; Professor of Liver Surgery Institute of Liver Studies, King's College Hospital London, UK Guilherme C Ribas MD Professor of Surgery University of São Paulo Medical School; Neurosurgeon, Hospital Israelita Albert Einstein São Paulo, Brazil; Visiting Professor of Neurosurgery University of Virginia Charlottesville, VA, USA Bruce Richard MBBS, MS, FRCS(Plast) Consultant Plastic Surgeon Birmingham Children's Hospital Birmingham, UK Michael J Rosen MD Professor of Surgery; Chief, Division of Gastrointestinal and General Surgery Case Medical Center Case Western Reserve University University Hospitals of Cleveland Cleveland, OH, USA Alistair C Ross MB, FRCS Consultant Orthopaedic Surgeon The Bath Clinic Bath, UK Stefano Sandrone PhD student Neuroscientist, NatBrainLab Sackler Institute of Translational Neurodevelopment Department of Forensic and Neurodevelopmental Sciences Institute of Psychiatry, Psychology and Neuroscience King's College London London, UK Martin Scaal PhD Professor of Anatomy and Developmental Biology Institute of Anatomy II University of Cologne Cologne, Germany Paul N Schofield MA, DPhil University Reader in Biomedical Informatics Department of Physiology, Development and Neuroscience University of Cambridge Cambridge, UK Nadav Schwartz MD Assistant Professor, Maternal Fetal Medicine Department of Obstetrics and Gynecology, Perelman School of Medicine University of Pennsylvania Philadelphia, PA, USA Vikram Sharma BSc(Hons), MBBS(Lon), MRCS(Eng), PG(Cert) Clinical Research Fellow Nuffield Department of Surgical Sciences University of Oxford Oxford, UK Richard M Sharpe BSc, Msc, PhD, FRSE Professor and Group Leader MRC Centre for Reproductive Health The Queen's Medical Research Institute University of Edinburgh Edinburgh, UK Mohammadali M Shoja MD Research Fellow Department of Neurosurgery University of Alabama at Birmingham Birmingham, AL, USA Victoria L Shone PhD, MSc, BSc Research Associate in Developmental Biology King's College London London, UK Monty Silverdale MD, PhD, FRCP Consultant Neurologist Salford Royal NHS Foundation Trust; Honorary Senior Lecturer in Neuroscience University of Manchester Manchester, UK Jonathan MW Slack MA, PhD, FMedSci Emeritus Professor, University of Bath Bath, UK; Emeritus Professor, University of Minnesota, Minneapolis, MN, USA Ariana L Smith MD Associate Professor of Urology Director of Pelvic Medicine and Reconstructive Surgery Penn Medicine, Perelman School of Medicine University of Pennsylvania Health System Philadelphia, PA, USA Carl H Snyderman MD, MBA Professor of Otolaryngology and Neurological Surgery Co-Director, UPMC Center for Cranial Base Surgery University of Pittsburgh Medical Center Pittsburgh, PA, USA Jane C Sowden PhD Professor of Developmental Biology and Genetics UCL Institute of Child Health University College London London, UK Robert J Spinner MD Chair, Department of Neurologic Surgery Burton M Onofrio, MD Professor of Neurosurgery; Professor of Orthopedics and Anatomy Mayo Clinic Rochester, MN, USA Jonathan D Spratt MA(Cantab), FRCS(Eng), FRCR Clinical Director of Diagnostic Radiology City Hospitals Sunderland NHS Foundation Trust Sunderland, UK; Visiting Professor of Anatomy Former anatomy examiner for the Royal College of Surgeons of England and Royal College of Radiologists Editor for Imaging Anatomy Jacob Bertram Springborg MD, PhD Consultant Neurosurgeon; Associate Professor of Neurosurgery University Clinic of Neurosurgery Copenhagen University Hospital Copenhagen, Denmark Susan Standring MBE, DSc, FKC, Hon FAS, Hon FRCS Emeritus Professor of Anatomy King's College London London, UK Ido Strauss MD, PhD Department of Neurosurgery Toronto Western Hospital Toronto, ON, Canada Mark D Stringer BSc, MS, FRCP, FRCS, FRCS(Ed), FRACS Professor of Paediatric Surgery Christchurch Hospital; Honorary Professor of Anatomy University of Otago Dunedin, New Zealand Paul H Sugarbaker MD, FACS, FRCS Medical Director, Center for Gastrointestinal Malignancies; Chief, Program in Peritoneal Surface Oncology MedStar Washington Hospital Center Washington, DC, USA Cheryll Tickle MA, PhD Emeritus Professor Department of Biology and Biochemistry University of Bath Bath, UK Kimberly S Topp PT, PhD, FAAA Professor and Chair, Department of Physical Therapy and Rehabilitation Science Professor, Department of Anatomy University of California, San Francisco San Francisco, CA, USA Drew A Torigian MD, MA, FSAR Associate Professor of Radiology; Clinical Director, Medical Image Processing Group Department of Radiology Hospital of the University of Pennsylvania Philadelphia, PA, USA David Tosh BSc, PhD Professor of Stem Cell and Regenerative Biology Centre for Regenerative Medicine University of Bath Bath, UK R Shane Tubbs MS, PA-C, PhD Chief Scientific Officer Seattle Science Foundation, Seattle, WA, USA; Professor of Human Gross and Developmental Anatomy Department of Anatomical Sciences St. George's University, Grenada, West Indies; Professor Centre of Anatomy and Human Identification University of Dundee, Dundee, UK Richard Tunstall BMedSci, PhD, PGCLTHE FHEA Head of Clinical Anatomy and Imaging Warwick Medical School University of Warwick, UK; University Hospitals Coventry and Warwickshire NHS Trust Coventry, UK; Visiting Professor of Anatomy St George's University, Grenada, West Indies Editor for Surface Anatomy Contributors to the forty-first edition xv Andry Vleeming PhD Professor of Clinical Anatomy University of New England College of Osteopathic Medicine Biddeford, ME, USA; Department of Rehabilitation Sciences and Physiotherapy Faculty of Medicine and Health Sciences Ghent University Ghent, Belgium Jan Voogd MD Emeritus Professor of Anatomy Department of Neuroscience Erasmus Medical Center Rotterdam, The Netherlands Bart Wagner BSc, CSci, FIBMS, Dip Ult Path. Chief Biomedical Scientist Electron Microscopy Unit Histopathology Department Royal Hallamshire Hospital (Sheffield Teaching Hospitals) Sheffield, UK Gary Warburton DDS, MD, FDSRCS, FACS Associate Professor; Program Director and Division Chief Oral and Maxillofacial Surgery University of Maryland Dental School Baltimore, MD, USA Jeremy PT Ward BSc, PhD Head of Department of Physiology; Professor of Respiratory Cell Physiology Department of Physiology King's College London London, UK John C Watkinson MSc, MS, FRCS, DLO Consultant ENT, Head and Neck and Thyroid Surgeon Queen Elizabeth Hospital University of Birmingham NHS Trust Birmingham, UK Alan J Wein MD, PhD(Hon), FACS Founders Professor and Chief of Urology Director, Urology Residency Program Penn Medicine, Perelman School of Medicine University of Pennsylvania Health System Philadelphia, PA, USA Caroline B Wigley BSc, PhD University of Exeter Medical School Exeter, UK Editor for Cell and Tissue Microstructure Frank H Willard PhD Professor of Anatomy University of New England College of Osteopathic Medicine Biddeford, Maine, USA Chin-Ho Wong MBBS, MRCS(Ed), MMed(Surg), FAMS(Plast Surg) Plastic Surgeon, Private Practice Singapore Stephanie J Woodley PhD, MSc, BPhty Senior Lecturer Department of Anatomy University of Otago Dunedin, New Zealand e5 HISTORICAL INTRODUCTION required to operate on real patients, or on soldiers injured at Sebastopol or some other battlefield. The book they planned together was a practi- cal one, designed to encourage youngsters to study anatomy, help them pass exams, and assist them as budding surgeons. It was not simply an anatomy textbook, but a guide to dissecting procedure, and to the major operations. Gray and Carter belonged to a generation of anatomists ready to infuse the study of human anatomy with a new, and respectable, scien- tificity. Disreputable aspects of the profession's history, acquired during the days of body-snatching, were assiduously being forgotten. The Anatomy Act of 1832 had legalized the requisition of unclaimed bodies from workhouse and hospital mortuaries, and the study of anatomy (now with its own Inspectorate) was rising in respectability in Britain. The private anatomy schools that had flourished in the Regency period were closing their doors, and the major teaching hospitals were erecting new, purpose-built dissection rooms (Richardson 2000). The best-known student works when Gray and Carter had qualified were probably Erasmus Wilson's Anatomist's Vade Mecum, and Elements of Anatomy by Jones Quain. Both works were small – pocket-sized – but Quain came in two thick volumes. Both Quain's and Wilson's works were good books in their way, but their small pages of dense type, and even smaller illustrations, were somewhat daunting, seeming to demand much nose-to-the-grindstone effort from the reader. The planned new textbook's dimensions and character were serious matters. Pocket manuals were commercially successful because they appealed to students by offering much knowledge in a small compass. But pocket-sized books had button-sized illustrations. Knox's Manual of Human Anatomy, for example, was a good book, but was only 6 inches by 4 (15 × 10 cm) and few of its illustrations occupied more than one- third of a page. Gray and Carter must have discussed this matter between themselves, and with Gray's publisher, JW Parker & Son, before deci- sions were taken about the size and girth of the new book, and espe- cially the size of its illustrations. While Gray and Carter were working on the book, a new edition of Quain's was published; this time it was a 'triple-decker' – in three volumes – of 1740 pages in all. The two men were earnestly engaged for the following 18 months in work for the new book. Gray wrote the text, and Carter created the illustrations; all the dissections were undertaken jointly. Their working days were long – all the hours of daylight, eight or nine hours at a stretch – right through 1856, and well into 1857. We can infer from the warmth of Gray's appreciation of Carter in his published acknowledge- ments that their collaboration was a happy one. The Author gratefully acknowledges the great services he has derived in the execution of this work, from the assistance of his friend, Dr. H. V. Carter, late Demonstrator of Anatomy at St George's Hospital. All the drawings from which the engravings were made, were executed by him. (Gray 1858) With all the dissections done, and Carter's inscribed wood-blocks at the engravers, Gray took six months' leave from his teaching at St George's to work as a personal doctor for a wealthy family. It was probably as good a way as any to get a well-earned break from the dissecting room and the dead-house (Nicol 2002). Carter sat the examination for medical officers in the East India Company, and sailed for India in the spring of 1858, when the book was still in its proof stages. Gray had left a trusted colleague, Timothy Holmes, to see it through the press. Holmes's association with the first edition would later prove vital to its survival. Gray looked over the final galley proofs, just before the book finally went to press. THE FIRST EDITION The book Gray and Carter had created together, Anatomy, Descriptive and Surgical, appeared at the very end of August 1858, to immediate Gray's Anatomy is now on its way to being 160 years old. The book is a rarity in textbook publishing in having been in continuous publication on both sides of the Atlantic Ocean, since 1858. One and a half centu- ries is an exceptionally long era for a textbook. Of course, the volume now is very different from the one Mr Henry Gray first created with his colleague Dr Henry Vandyke Carter, in mid-Victorian London. In this introductory essay, I shall explain the long history of Gray's, from those Victorian days right up to today. The shortcomings of existing anatomical textbooks probably impressed themselves on Henry Gray when he was still a student at St George's Hospital Medical School, near London's Hyde Park Corner, in the early 1840s. He began thinking about creating a new anatomy textbook a decade later, while war was being fought in the Crimea. New legislation was being planned that would establish the General Medical Council (1858) to regulate professional education and standards. Gray was twenty-eight years old, and a teacher himself at St George's. He was very able, hard-working and highly ambitious, already a Fellow of the Royal Society, and of the Royal College of Surgeons. Although little is known about his personal life, his was a glittering career so far, achieved while he served and taught on the hospital wards and in the dissecting room (Fig. 1) (Anon 1908). Gray shared the idea for the new book with a talented colleague on the teaching staff at St George's, Henry Vandyke Carter, in November 1855. Carter was from a family of Scarborough artists, and was himself a clever artist and microscopist. He had produced fine illustrations for Gray's scientific publications before, but could see that this idea was a much more complex project. Carter recorded in his diary: Little to record. Gray made proposal to assist by drawings in bringing out a Manual for students: a good idea but did not come to any plan … too exacting, for would not be a simple artist (Carter 1855). Neither of these young men was interested in producing a pretty book, or an expensive one. Their purpose was to supply an affordable, accurate teaching aid for people like their own students, who might soon be Fig. 1 Henry Gray (1827–1861) is shown here in the foreground, seated by the feet of the cadaver. The photograph was taken by a medical student, Joseph Langhorn. The room is the dissecting room of St George's Hospital Medical School in Kinnerton Street, London. Gray is shown surrounded by staff and students. When the photo was taken, on 27 March 1860, Carter had left St George's, to become Professor of Anatomy and Physiology at Grant Medical College, in Bombay (nowadays Mumbai). The second edition of Gray's Anatomy was in its proof stages, to appear in December 1860. Gray died just over a year later, in June 1861, at the height of his powers. Historical introduction e6 Fig. 2 Henry Vandyke Carter (1831–1897). Carter was appointed Honorary Surgeon to Queen Victoria in 1890. acclaim. Reviews in The Lancet and the British Medical Journal were highly complimentary, and students flocked to buy. It is not difficult to understand why it was a runaway success. Gray's Anatomy knocked its competitors into a cocked hat. It was considerably smaller and more slender than the doorstopper with which modern readers are familiar. The book held well in the hand, it felt substantial, and it contained everything required. To contemporaries, it was small enough to be portable, but large enough for decent illustrations: 'royal octavo' – 912 × 6 inches (24 × 15 cm) – about two-thirds of modern A4 size. Its medium-size, single-volume format was far removed from Quain, yet double the size of Knox's Manual. Simply organized and well designed, the book explains itself confi- dently and well; the clarity and authority of the prose are manifest. But what made it unique for its day was the outstanding size and quality of the illustrations. Gray thanked the wood engravers Butterworth and Heath for the 'great care and fidelity' they had displayed in the engrav- ings, but it was really to Carter that the book owed its extraordinary success. The beauty of Carter's illustrations resides in their diagrammatic clarity, quite atypical for their time. The images in contemporary anatomy books were usually 'proxy-labelled': dotted with tiny numbers or letters (often hard to find or read) or bristling with a sheaf of num- bered arrows, referring to a key situated elsewhere, usually in a footnote, which was sometimes so lengthy it wrapped round on to the following page. Proxy labels require the reader's eye to move to and fro: from the structure to the proxy label to the legend and back again. There was plenty of scope for slippage, annoyance and distraction. Carter's illustra- tions, by contrast, unify name and structure, enabling the eye to assimi- late both at a glance. We are so familiar with Carter's images that it is hard to appreciate how incredibly modern they must have seemed in 1858. The volume made human anatomy look new, exciting, accessible and do-able. The first edition was covered in a brown bookbinder's cloth embossed all over in a dotted pattern, and with a double picture-frame border. Its spine was lettered in gold blocking: sense of calamity. The grand old medical man Sir Benjamin Brodie, Sergeant-Surgeon to the Queen, and the great supporter of Gray to whom Anatomy had been dedicated, cried forlornly: 'Who is there to take his place?' (Anon 1908). But old JW Parker ensured the survival of Gray's by inviting Timothy Holmes, the doctor who had helped proof-read the first edition, and who had filled Gray's shoes at the medical school, to serve as Editor for the next edition. Other long-running anatomy works, such as Quain, remained in print in a similar way, co-edited by other hands (Quain 1856). Holmes (1825–1907) was another gifted St George's man, a scholar- ship boy who had won an exhibition to Cambridge, where his brilliance was recognized. Holmes was a Fellow of the Royal College of Surgeons at 28. John Parker junior had commissioned him to edit A System of Surgery (1860–64), an important essay series by distinguished surgeons on subjects of their own choosing. Many of Holmes's authors remain important figures, even today: John Simon, James Paget, Henry Gray, Ernest Hart, Jonathan Hutchinson, Brown-Séquard and Joseph Lister. Holmes had lost an eye in an operative accident, and he had a gruff manner that terrified students, yet he published a lament for young Parker that reveals him capable of deep feeling (Holmes 1860). John Parker senior's heart, however, was no longer in publishing. His son's death had closed down the future for him. The business, with all its stocks and copyrights, was sold to Messrs Longman. Parker retired to the village of Farnham, where he later died. With Holmes as editor, and Longman as publisher, the immediate future of Gray's Anatomy was assured. The third edition appeared in 1864 with relatively few changes, Gray's estate receiving the balance of his royalty after Holmes was paid £100 for his work. THE MISSING OBITUARY Why no obituary appeared for Henry Gray in Gray's Anatomy is curious. Gray had referred to Holmes as his 'friend' in the preface to the first edition, yet it would also be true to say that they were rivals. Both had just applied for a vacant post at St George's, as Assistant Surgeon. Had Gray lived, it is thought that Holmes may not have been appointed, despite his seniority in age (Anon 1908). Later commentators have suggested, as though from inside knowl- edge, that Holmes's 'proof-reading' included improving Gray's writing … with 'DESCRIPTIVE AND SURGICAL' in small capitals underneath. Gray's Anatomy is how it has been referred to ever since. Carter was given credit with Gray on the book's title page for undertaking all the dissec- tions on which the book was based, and sole credit for all the illustra- tions, though his name appeared in a significantly smaller type, and he was described as the 'Late Demonstrator in Anatomy at St George's Hospital' rather than being given his full current title, which was Profes- sor of Anatomy and Physiology at Grant Medical College, Bombay. Gray was still only a Lecturer at St George's and he may have been aware that his words had been upstaged by the quality of Carter's anatomical images. He need not have worried: Gray is the famous name on the spine of the book. Gray was paid £150 for every thousand copies sold. Carter never received a royalty payment, just a one-off fee at publication, which may have allowed him to purchase the long-wished-for microscope he took with him to India (Fig. 2). The first edition print-run of 2000 copies sold out swiftly. A parallel edition was published in the United States in 1859, and Gray must have been deeply gratified to have to revise an enlarged new English edition in 1859–60, though he was surely saddened and worried by the death of his publisher, John Parker junior, at the young age of 40, while the book was going through the press. The second edition came out in the December of 1860 and it too sold like hot cakes, as indeed has every subsequent edition. The following summer, in June 1861, at the height of his powers and full of promise, Henry Gray died unexpectedly at the age of only 34. Gray had contracted smallpox while nursing his nephew. A new strain of the disease was more virulent than the one with which Gray had been vaccinated as a child; the disease became confluent, and Gray died in a matter of days. Within months, the whole country would be pitched into mourning for the death of Prince Albert. The creative era over which he had pre- sided – especially the decade that had flowered since the Great Exhibi- tion of 1851 – would be history. THE BOOK SURVIVES Anatomy Descriptive and Surgical could have died too. With Carter in India, the death of Gray, so swiftly after that of the younger Parker, might have spelled catastrophe. Certainly, at St George's there was a GRAY'S ANATOMY Historical introduction e7 were not as yet perfected, and in any case could not provide the bold simplicity of line required for a book like Gray's, which depended so heavily on clear illustration and clear lettering. Recognizing the inferior- ity of half-tone illustrations by comparison with Carter's wood-engraved originals, Pick and Howden courageously decided to jettison the second-rate half-tones altogether. Most of the next edition's illustrations were either Carter's, or old supplementary illustrations inspired by his work, or newly commissioned wood engravings or line drawings, intended 'to harmonize with Carter's original figures'. They successfully emulated Carter's verve. Having fewer pages and lighter paper, the 1905 (sixteenth edition) weighed less than its predecessor, at 4 lb 11 oz/2.1 kg. Typographically, the new edition was superb. Howden took over as sole editor in 1909 (seventeenth edition) and immediately stamped his personality on Gray's. He excised 'Surgical' from the title, changing it to Anatomy Descriptive and Applied, and removed Carter's name altogether. He also instigated the beginnings of an editorial board of experts for Gray's, by adding to the title page 'Notes on Applied Anatomy' by AJ Jex-Blake and W Fedde Fedden, both St George's men. For the first time, the number of illustrations exceeded one thousand. Howden was responsible for the significant innovation of a short historical note on Henry Gray himself, nearly 60 years after his death, which included a portrait photograph (1918, twentieth edition). THE NOMENCLATURE CONTROVERSY Howden's era, and that of his successor TB Johnston (of Guy's), was overshadowed by a cloud of international controversy concerning ana- tomical terminology. European anatomists were endeavouring to stand- ardize anatomical terms, often using Latinate constructions, a move resisted in Britain and the United States. Gray's became mired in these debates for over 20 years. The attempt to be fair to all sides by using multiple terms doubtless generated much confusion amongst students, until a working compromise was at last arrived at in 1955 (thirty-second edition, 1958). Johnston oversaw the second retitling of the book (in 1938, twenty- seventh edition): it was now, officially, Gray's Anatomy, finally ending the fiction that it had ever been known as anything else. Gray's suffered from paper shortages and printing difficulties in World War II, but suc- cessive editions nevertheless continued to grow in size and weight, while illustrations were replaced and added as the text was revised. Between Howden's first sole effort (1909, seventeenth edition) and Johnston's last edition (1958, thirty-second edition), Gray's expanded by over 300 pages – from 1296 to 1604 pages, and almost 300 addi- tional illustrations brought the total to over 1300. Johnston also intro- duced X-ray plates (1938) and, in 1958 (thirty-second edition), electron micrographs by AS Fitton-Jackson, one of the first occasions on which a woman was credited with a contribution to Gray's. Johnston felt com- pelled to mention that she was 'a blood relative of Henry Gray himself', perhaps by way of mitigation. AFTER WORLD WAR II The editions of Gray's issued in the decades immediately following the Second World War give the impression of intellectual stagnation. Steady expansion continued in an almost formulaic fashion, with the insertion of additional detail. The central historical importance of innovation in the success of Gray's seems to have been lost sight of by its publishers and editors – Johnston (1930–1958, twenty-fourth to thirty-second editions), J Whillis (co-editor with Johnston, 1938–1954), DV Davies (1958–1967, thirty-second to thirty-fourth editions) and F Davies (co-editor with DV Davies 1958–1962, thirty-second to thirty-third editions). Gray's had become so pre-eminent that perhaps complacency crept in, or editors were too daunted or too busy to confront the 'massive undertaking' of a root and branch revision (Tansey 1995). The unexpected deaths of three major figures associated with Gray's in this era, James Whillis, Francis Davies and David Vaughan Davies – each of whom had been ready to take the editorial reins – may have contributed to retarding the process. The work became somewhat dull. KEY EDITION: 1973 DV Davies had recognized the need for modernization, but his unex- pected death left the work to other hands. Two Professors of Anatomy at Guy's, Roger Warwick and Peter Williams, the latter of whom had been involved as an indexer for Gray's for several years, regarded it as an honour to fulfill Davies's intentions. Their thirty-fifth edition of 1973 was a significant departure from tradition. Over 780 pages (of 1471) were newly written, almost a third style. This could be a reflection of Holmes's own self-regard, but there may be some truth in it. There can be no doubt that, as Editor of seven subsequent editions of Gray's Anatomy (third to ninth editions, 1864– 1880), Holmes added new material, and had to correct and compress passages, but it is also possible that, back in 1857, Gray's original manuscript had been left in a poor state for Holmes to sort out. In other works, Gray's writing style was lucid, but he always seems to have paid a copyist to transcribe his work prior to submission. The original manu- script of Gray's Anatomy, sadly, has not survived, so it is impossible to be sure how much of the finished version had actually been written by Holmes. It may be that Gray's glittering career, or perhaps the patronage that unquestionably advanced it, created jealousies among his colleagues, or that there was something in Gray's manner that precluded affection, or that created resentments among clever social inferiors like Carter and Holmes, especially if they felt their contributions to his brilliant career were not given adequate credit. Whatever the explanation, no reference to Gray's life or death appeared in Gray's Anatomy itself until the twen- tieth century (Howden et al 1918). A SUCCESSION OF EDITORS Holmes expanded areas of the book that Gray himself had developed in the second edition (1860), notably in 'general' anatomy (histology) and 'development' (embryology). In Holmes's time as Editor, the volume grew from 788 pages in 1864 to 960 in 1880 (ninth edition), with the histological section paginated separately in roman numerals at the front of the book. Extra illustrations were added, mainly from other published sources. The connections with Gray and Carter, and with St George's, were maintained with the appointment of the next editor, T. Pickering Pick, who had been a student at St George's in Gray's time. From 1883 (tenth edition) onwards, Pick kept up with current research, rewrote and inte- grated the histology and embryology into the volume, dropped Holmes from the title page, removed Gray's preface to the first edition, and added bold subheadings, which certainly improved the appearance and accessibility of the text. Pick said he had 'tried to keep before himself the fact that the work is intended for students of anatomy rather than for the Scientific Anatomist' (thirteenth edition, 1893). Pick also introduced colour printing (in 1887, eleventh edition) and experimented with the addition of illustrations using the new printing method of half-tone dots: for colour (which worked) and for new black- and-white illustrations (which did not). Half-tone shades of grey com- pared poorly with Carter's wood engravings, still sharp and clear by comparison. What Henry Vandyke Carter made of these changes is a rich topic for speculation. He returned to England in 1888, having retired from the Indian Medical Service, full of honours – Deputy Surgeon General, and in 1890, he was made Honorary Surgeon to Queen Victoria. Carter had continued researching throughout his clinical medical career in India, and became one of India's foremost bacteriologists/tropical disease specialists before there was really a name for either discipline. Carter made some important discoveries, including the fungal cause of mycetoma, which he described and named. He was also a key figure in confirming scientifically in India some major international discoveries, such as Hansen's discovery of the cause of leprosy, Koch's discovery of the organism causing tuberculosis, and Laveran's discovery of the organ- ism that causes malaria. Carter married late in life, and his wife was left with two young children when he died in Scarborough in 1897, aged 65. Like Gray, he received no obituary in the book. When Pick was joined on the title page by Robert Howden (a profes- sional anatomist from the University of Durham) in 1901 (fifteenth edition), the volume was still easily recognizable as the book Gray and Carter had created. Although many of Carter's illustrations had been revised or replaced, many others still remained. Sadly, though, an entire section (embryology) was again separately paginated, as its revision had taken longer than anticipated. Gray's had grown, seemingly inexorably, and was now quite thick and heavy: 1244 pages, weighing 5 lb 8 oz/2.5 kg. Both co-editors, and perhaps also its publisher, were dis- satisfied with it. KEY EDITION: 1905 Serious decisions were taken well in advance of the next edition, which turned out to be Pick's last with Howden. Published 50 years after Gray had first suggested the idea to Carter, the 1905 (sixteenth) edition was a landmark one. The period 1880–1930 was a difficult time for anatomical illustra- tion, because the new techniques of photo-lithography and half-tone Historical introduction e8 had developed a distinct character of its own in the interval), and sold extremely well there (Williams and Warwick 1973). The influence of the Warwick and Williams edition was forceful and long-lasting, and set a new pattern for the following quarter-century. As has transpired several times before, wittingly or unwittingly, a new editor was being prepared for the future: Dr Susan Standring (of Guy's), who created the new bibliography for the 1973 edition of Gray's, went on to serve on the editorial board, and has served as Editor-in-Chief for the last two editions before this one (2005–2008, thirty-ninth and fortieth editions). Both editions are important for dif- ferent reasons. For the thirty-ninth edition, the entire content of Gray's was reorgan- ized, from systematic to regional anatomy. This great sea-change was not just organizational but historic, because, since its outset, Gray's had prioritized bodily systems, with subsidiary emphasis on how the systems interweave in the regions of the body. Professor Standring explained that this regional change of emphasis had long been asked for by readers and users of Gray's, and that new imaging techniques in our era have raised the clinical importance of local anatomy (Standring 2005). The change was facilitated by an enormous collective effort on the part of the editorial team and the illustrators. The subsequent and current editions consolidate that momentous change. (See Table 1.) Table 1 Gray's Anatomy Editions Edition Date Author/Editor(s) Publisher Title 1st 1858 Henry Gray JW Parker & Son Anatomy Descriptive and Surgical The drawings by Henry Vandyke Carter. The dissections jointly by the author and Dr Carter 2nd 1860 Henry Gray JW Parker & Son 3rd 1864 T Holmes Longman 4th 1866 T Holmes Longman 5th 1869 T Holmes Longman 6th 1872 T Holmes Longman 7th 1875 T Holmes Longman 8th 1877 T Holmes Longman 9th 1880 T Holmes Longman 10th 1883 TP Pick Longman 11th 1887 TP Pick Longman 12th 1890 TP Pick Longman 13th 1893 TP Pick Longman Gray's preface removed 14th 1897 TP Pick Longman 15th 1901 TP Pick & R Howden Longman 16th 1905 TP Pick & R Howden Longman 17th 1909 Robert Howden Longman Anatomy Descriptive and Applied Notes on applied anatomy by AJ Jex-Blake & W Fedde Fedden 18th 1913 Robert Howden & Blake & Fedden Longman 19th 1916 Robert Howden & Blake & Fedden Longman 20th 1918 Robert Howden & Blake & Fedden Longman First edition ever to feature a photograph and obituary of Henry Gray 21st 1920 Robert Howden Longman Notes on applied anatomy by AJ Jex-Blake & John Clay 22nd 1923 Robert Howden Longman Notes on applied anatomy by John Clay & John D Lickley 23rd 1926 Robert Howden Longman 24th 1930 TB Johnston Longman 25th 1932 TB Johnston Longman 26th 1935 TB Johnston Longman 27th 1938 TB Johnston & J Whillis Longman Gray's Anatomy 28th 1942 TB Johnston & J Whillis Longman 29th 1946 TB Johnston & J Whillis Longman 30th 1949 TB Johnston & J Whillis Longman 31st 1954 TB Johnston & J Whillis Longman 32nd 1958 TB Johnston & DV Davies & F Davies Longman 33rd 1962 DV Davies & F Davies Longman 34th 1967 DV Davies & RE Coupland Longman 35th 1973 Peter L Williams & Roger Warwick Longman With a separate volume: Functional Neuroanatomy of Man – being the neurology section of Gray's Anatomy. 35th edition, 1975 36th 1980 Roger Warwick & Peter L Williams Churchill Livingstone 37th 1989 Peter L Williams Churchill Livingstone 38th 1995 Peter L Williams & Editorial Board Churchill Livingstone 39th 2005 Susan Standring & Editorial Board Elsevier The Anatomical Basis of Clinical Practice 40th 2008 Susan Standring & Editorial Board Elsevier The Anatomical Basis of Clinical Practice 41st 2015 Susan Standring & Editorial Board Elsevier The Anatomical Basis of Clinical Practice of the illustrations were newly commissioned, and the illustration cap- tions were freshly written throughout. With a complete re-typesetting of the text in larger double-column pages, a new index and the innova- tion of a bibliography, this edition of Gray's looked and felt quite unlike its 1967 (thirty-fourth edition) predecessor, and much more like its modern incarnation. This 1973 edition departed from earlier volumes in other significant ways. The editors made explicit their intention to try to counter the impetus towards specialization and compartmentalization in twentieth- century medicine, by embracing and attempting to reintegrate the com- plexity of the available knowledge. Warwick and Williams openly renounced the pose of omniscience adopted by many textbooks, believ- ing it important to accept and mention areas of ignorance or uncer- tainty. They shared with the reader the difficulty of keeping abreast in the sea of research, and accepted with a refreshing humility the impos- sibility of fulfilling their own ambitious programme. Warwick and Williams's 1973 edition had much in common with Gray and Carter's first edition. It was bold and innovative