Netter’s Atlas of Human Embryology This page intentionally left blank Netter’s Atlas of Human Embryology Larry R. Cochard, PhD Associate Professor Northwestern University The Feinberg School of Medicine Chicago, Illinois Illustrations by Frank H. Netter, MD Contributing Illustrators John A. Craig, MD Carlos A. G. Machado, MD Copyright © 2012 by Saunders, an imprint of Elsevier Inc. NETTER’S ATLAS OF HUMAN EMBRYOLOGY, Updated Edition All rights reserved. No part of this book may be reproduced in any form or by any electronic or mechanical means, including information storage and retrieval systems, without permission in writing from the publisher. Permission for Netter art figures may be sought directly from Elsevier’s Health Science Licensing Department in Philadelphia, PA, USA: phone 1-800-523-1649, ext. 3276 or (215) 239-3276; or e-mail [email protected] ISBN: 978-1-4557-3977-6 eBook ISBN: 978-1-4557-3978-3 Library of Congress Catalog No: 2001132799 Printed in the United States of America First Printing, 2002 NOTICE Every effort has been taken to confirm the accuracy of the information presented and to describe generally accepted practices. Neither the publisher nor the authors can be held responsible for errors or for any consequences arising from the use of the information contained herein, and make no warranty, expressed or implied, with respect to the contents of the publication. Last digit is the print number: 9 8 7 6 5 4 3 2 1 To Dr. David Langebartel As my teacher and mentor at the University of Wisconsin—Madison, he stressed the relationship between embryology and adult anatomy, and he did so with energy, authority, and a considerable amount of humor. And to the memory of Dr. Leslie B. Arey He was a colleague at the beginning of my career at Northwestern. It was a privilege and a very humbling experience for a young, green anatomist to teach with the 20th-century master of embryology, anatomy, and histology. This page intentionally left blank Preface This book is intended for first-year medical format. Embryological pictures can also be students, dental students, and other difficult and frustrating for students because beginning students of embryology. As an of the three-dimensional complexity of the atlas, it is a showcase for the incomparable embryo and the unfamiliar structures and artwork of Dr. Frank H. Netter. The Netter relationships. To address this problem, paintings in this Atlas were published in the book consists of more than just labeled The Netter Collection of Medical images. It contains tables, schematics, Illustrations, Dr. Netter’s series of systemic concepts, descriptive captions, summaries, monographs that integrate anatomy, chapter glossaries, and concise text at the embryology, physiology, pathology, bottom of each page that address all of the functional anatomy, and clinical anatomy. major events and processes of normal and They were also published in the Clinical abnormal development. Histological Symposia that address particular topics. As principles are briefly covered to help the necessary, new images were created by uninitiated understand the many references John A. Craig, MD, and Carlos Machado, to embryonic tissues in this book. MD. Plates were selected to match the Little was known about the genetic and scope of material that is suitable for molecular basis of development when Dr. beginning students and arranged in a Netter drew most of his illustrations, and logical sequence. an atlas is not the ideal medium to convey The theme throughout this book is an this type of information. I believe it is emphasis on morphological patterns in important, though, to introduce the subject the embryo and how they relate to the and to include examples of the control of organization and function of structures in development. Illustrations from the Atlas the adult. Another important focus is the are used to introduce cellular, molecular, embryological basis of congenital birth and genetic concepts such as induction, defects. Descriptive embryology can be apoptosis, growth factors, and genetic an educational goal, but the study of patterning and determination. These embryology is more effective, rewarding, are by necessity selective and include and relevant when it is placed in a major events (e.g., limb development, biological or clinical context that goes segmentation of the head) or processes that beyond the embryo itself. The focus have broad significance in development on morphological themes in prenatal (e.g., the interactions between epithelia and development makes it easier to learn adult connective tissue in organ development). anatomy and to understand an abnormality If nothing else, this material will serve in a patient. In keeping with this idea, this to remind students of the complexity of Atlas contains some Netter plates of adult development and the dynamic events at anatomy. These include parts of the body the cellular and molecular level. where complex anatomy has embryonic The terminology tables at the end of relevance. They also provide context to each chapter are also selective. The terms help show the relationships between include major structures, potentially primordia and derivatives. confusing structures, and histological or Like anatomy, embryology is a very anatomical terms that provide context. The visual subject that lends itself to an atlas glossary is also an opportunity to include Preface terms that did not make it into a chapter or to congenital defects. Chapter 9 is on the head and elaborate on important ones. At the risk of some neck region. overlap, I decided to have a terminology section This annotated Atlas can serve as a bridge at the end of each chapter instead of at the end of between the material presented in the classroom the book. This makes it a more effective learning and the detail found in textbooks. It can be useful tool, as students use this Atlas in their studies for board exam review, and to that end, there is rather than an isolated reference feature. an appendix that summarizes all of the major Chapter 1 is an overview of the major congenital anomalies and their embryonic basis. developmental periods, events, and processes and More than anything, this Atlas is about the art of ends with a section on the mechanisms of Dr. Netter. The clarity, realism, and beauty of his abnormal development and the classification of illustrations make the study of embryology more anomalies. Chapter 2 addresses gastrulation, the enlightening and enjoyable. vertebrate body plan, and the placenta. Chapters 3 through 8 are organized by systems and include Larry R. Cochard, PhD viii Frank H. Netter, MD Frank H. Netter was born in 1906 in New York City. He studied art at the Art Student’s League and the National Academy of Design before entering medical school at New York University, where he received his medical degree in 1931. During his student years, Dr. Netter’s notebook sketches attracted the attention of the medical faculty and other physicians, allowing him to augment his income by illustrating articles and textbooks. He continued illustrating as a sideline after establishing a surgical practice in 1933, but he ultimately opted to give up his practice in favor of a full-time commitment to art. After service in the United States Army during World War II, Dr. Netter began his long collaboration with the CIBA Pharmaceutical Company (now Novartis Pharmaceuticals). This 45-year partnership resulted in the production of the extraordinary collection of medical art so familiar to physicians and other medical professionals worldwide. Icon Learning Systems acquired the Netter Collection in July 2000 and continues to update Dr. Netter’s original paintings and to add newly commissioned paintings by artists trained in the style of Dr. Netter. Dr. Netter’s works are among the finest examples of the use of illustration in the teaching of medical concepts. The 13-book Netter Collection of Medical Illustrations, which includes the greater part of the more than 20,000 paintings created by Dr. Netter, became and remains one of the most famous medical works ever published. The Netter Atlas of Human Anatomy, first published in 1989, presents the anatomical paintings from the Netter Collection. Now translated into 11 languages, it is the anatomy atlas of choice among medical and health professions students the world over. The Netter illustrations are appreciated not only for their aesthetic qualities, but more important, for their intellectual content. As Dr. Netter wrote in 1949, “… clarification of a subject is the aim and goal of illustration. No matter how beautifully painted, how delicately and subtly rendered a subject may be, it is of little value as a medical illustration if it does not serve to make clear some medical point.” Dr. Netter’s planning, conception, point of view, and approach are what inform his paintings and what make them so intellectually valuable. Frank H. Netter, MD, physician and artist, died in 1991.