The Viscera of the Domestic Mammals

The Viscera of the Domestic Mammals

R.Nickel A.Schummer E.Seiferle The Viscera of the Domestic Mammals Second revised edition by August Schummer, Richard Nickel and Wolfgang Otto Sack With 559 illustrations, some in color, in the text and on 13 plates 1979 Springer-Verlag Berlin Heidelberg GmbH 1979

This work is an authorized translation and revision of R. NICKEL, A. SCHUMMER, E. SEIFERLE, (Ed.), Lehrbuch der Anatomie der Haustiere (Textbook of the Anatomy of Domestic Animals), Volume II: Eingeweide (The Viscera of the Domestic Mammals) by A. SCHUMMER and R. NICKEL, 4th Edition, 1979. Verlag Paul Parey, Berlin und Hamburg, Germany. RICHARD NICKEL t, Dr. med. vet., Professor and Head of the Department of Anatomy, Tieraerztliche Hochschule Hannover, D-3000 Hannover, Germany AUGUST SCHUMMER t, Dr. med. vet., Professor and Head of the Department of Veterinary Anatomy, Justus-Liebig-Universitaet Giessen, D-6300 Giessen, Germany EUGEN SEIFERLE, Dr. med. vet., Dr. med. veto h.c., Professor and Head of the Department of Veterinary Anatomy, Universitaet Zurich, CH-8057 Zurich, Switzerland WOLFGANG OTIO SACK, D.V.M., Ph. D., Dr. med. vet., Professor of Anatomy, New York State Veterinary College, Cornell University, Ithaca, N.Y. 14850 USA. Synopsis of the English edition: Textbook of the Anatomy of the Domestic Animals Volume 1: Locomotor System of the Domestic Mammals. By R. NICKEL, A. SCHUMMER, E. SEIFERLE, J. FREWEIN and K.-H. WILLE. Translation from the German. Approx. 560 pages, with about 517 illustrations in the text and on Il colour plates. In preparation. Volume II: The Viscera of the Domestic Mammals. By A. SCHUMMER, R. NICKEL and W. O. SACK. 2nd revised edition. Translated and revised from the 4th German edition. 1979. 446 pages, with a total of 559 illustrations in the text and on 13 colour plates. Volume III: Circulatory System, Skin and Skin Organs of the Domestic Mammals. By A. SCHUMMER, H. WILKENS, B. VOLLMERHAUS and K.-H. HABERMEHL. Translated from the German by W. G. SILLER and P. L. A. Wight. 1980. Approx. 662 pages, with a total of about 439 illustrations, about 172 in colour. In preparation. Volume IV: Nervous System, Sensory Organs, Endocrine Glands of the Domestic Mammals. By E. SEIFERLE. Translation from the German. Approx. 442 pages, with a total of about 250 illustrations, about 95 in colour, in the text and on 10 colour plates. In preparatlon. Volume V: Anatomy of the Domestic Birds. By A. SCHUMMER. Translated from the German by W. G. SILLER and P. A. L. WIGHT. 1977. 214 pages, with 141 illustrations in the text and on 7 colour plates. Synopsis of the German edition: Lehrbuch der Anatomie der Haustiere Volume 1: Bewegungsapparat. By R. NICKEL, A. SCHUMMER, E. SEIFERLE, J. FREWEIN and K.-H. WILLE. 4th revised edition. 1977. 560 pages, with a total of 517 illustrations in the text and on II colour plates. Volume II: Eingeweide. By A. SCHUMMER and R. NICKEL. 4th edition. 1979.446 pages, with a total of 559 illustrations in the text and on 13 colour plates. Volume III: Kreislaufsystem, Haut und Hautorgane. By A. SCHUMMER, H. WILKENS, B. VOLLMERHAUS and K.-H. HABERMEHL. 1976. 662 pages, with a total of 439 illustrations, 172 in colour. Volume IV: Nervensystem, Sinnesorgane, Endokrine Driisen. By E. SEIFERLE. 1975.442 pages, with a total of 250 illustrations, 95 in colour in the text and on 10 colour plates. Volume V: Anatomie der Hausvogel. By A. SCHUMMER. 1973. 215 pages, with a total of 141 illustrations in the text and on 7 colour plates. CIP-Kurztitelaufnahme der Deutschen Bibliothek Nickel, Richard: The viscera of the dom estic mammals / R. Nickel ; A. Schummer ; E. Seiferle. Transl. and revision by Wolfgang Otto Sack. - 2., rev. ed. / by August Schummer and Richard Nickel. 1 st. Ed. 1973 ISBN 978-1-4757-6816-9 Dt. Ausg. u.d.t.: Nickel, Richard: Lehrhuch der Anatomie der Haustiere. Bd. 2. Eingeweide. ISBN 978-1-4757-6816-9 ISBN 978-1-4757-6814-5 (ebook) DOI 10.1007/978-1-4757-6814-5 NE: Schummer, August:; Seiferle, Eugen: This work is subject to copyright. AU rights are reserved, whether the whole or part of the material is concerned, specifically those rights of translation, reprinting re-use of illustrations, recitation, broadcasting, reproduction by photocopying machine or similar means, and storage in data banks. Under 54,1 of the German Copyright Law where single copies are made for other than privat use, a fee is payable to the publisher according to 54,2 of the German Copyright Law. The amount of the fee is to be determined by agreement with the publisher. 1973, 1979 by Springer-Verlag Berlin Heidelberg Originally published by Springer-Verlag in 1979 Softcover reprint of the hardcover 2nd edition 1979 ISBN 978-1-4757-6816-9

Dedicated with Admiration and Gratitude to PAUL MARTIN PAUL MARTIN (1861-1937), Professor, Dr. med. vet., Dr. phil. h.c., Dr. med. vet. h.c. Veterinary Anatomist from 1886 to 1901 in Zurich, and from 1901 to 1928 in Giessen

Preface to the Second English Edition Since the publication of this book in 1973, its principal author, Professor AuGUST ScHUMMER, former Head of the Veterinary Anatomy Department, Justus Liebig-Universitat, Giessen, West Germany, has passed away. Over the years an amiable understanding as between colleagues dedicated to the same scientific discipline developed between us. This was especially true during the years when I was translating this volume, which was perhaps his most cherished work. This understanding seems to have been flavored with paternal feelings on his part. The loss of this man, whose advice was often sought and whose help was freely given, and of this kind of friendship is deeply regretted by the writer, as it is by many others who knew him well. During the last six years the remaining two volumes of the originally planned five-volume set of the Lehrbuch der Anatomie der Haustiere (Textbook of the Anatomy of the Domestic Animals) was published. Volume V (Anatomy of the Domestic Birds) - also principally ScHUMMER's contribution - has since 1977 been available in English. An English edition of Volume III (Circulatory System and Integument) will be published next year. Volumes I and IV (Musculoskeletal and Nervous Systems) will follow in due course. The second edition of this translation, in addition to numerous editorial, typographical and some nomenclatorial corrections, includes two substantive changes. A section on the erection of the ruminant penis has been added, and the pathways by which the spermatozoa leave the testis for the ducts in the epididymis have been clarified. The writer is grateful, particularly to Professor RoBERT HABEL, former Head of the Veterinary Anatomy Department at the New York State College of Veterinary Medicine, Cornell University; to his colleagues in the same department; and to the many reviewers of the first edition for their valuable suggestions, all of which have been considered in the revision of this book for the second edition. In keeping with the goal of providing as complete and as modern a gross-anatomical bibliography as possible, about fifty recent references to the veterinary anatomical and appropriate clinical literature have been added. Ithaca, New York, January 1979 w. 0. SACK Preface to the First English Edition Soon after the first two volumes of the Lehrbuch der A natomie der H austiere by R. NICKEL, A. ScHOMMER, and E. SEIFERLE were published in German, inquiries were made by persons in various countries about the possibility of having this textbook translated into other languages. Therefore, the publisher decided to produce an English edition limited at first to Volume II. The concept and plan of the original German work by AUGUST ScHUMMER and the late RICHARD NICKEL has been preserved in this translation, and what was said in the prefaces to the first and second German editions about the purpose and. scope of the book applies

VIII Preface equally to this first English edition. The work deals with the body cavities, digestive system and teeth, spleen, and with the respiratory and urogenital systems of the dog, cat, pig, ox, sheep, goat, and horse. Each organ system is described in a general and comparative chapter, which is followed by shorter special chapters for the carnivores, pig, ruminants, and horse. In agreement with the original authors, substantive changes were made in several instances to take into account the results of recent research and to eliminate conflicts between views commonly held by German anatomists and those outside of Europe, but foremost to profit by the advances in Nomina anatomica veterinaria* (NAV), a uniform international nomenclature, which came into existence while this translation was in progress. This nomenclature lists a single, usually descriptive term for homologous structures in all domestic mammals, and wherever possible for the same structure in man; and thus has the potential of simplifying student instruction and promoting interdisciplinary understanding. The work of the International Committee on Veterinary Anatomical Nomenclature in many instances included re-evaluations of existing anatomical concepts; and it was these that necessitated most of the changes in the present work. The nomenclature conforms, with very few exceptions, to the second edition of the NA V. In keeping with the textbook character of the present work, most of the official Latin terms have been translated to accepted English equivalents. Only where the Latin differed greatly from the English, or where it enhanced understanding, has the official term been added parenthetically. In this edition, a bibliography has been added at the end of each of the major sections, and an attempt has been made to compile citations of the more recent studies dealing with the gross anatomy of the viscera of the domestic mammals. Dr. K. H. WILLE of the Department of Veterinary Anatomy, Justus-Liebig-Universitaet Giessen has gathered most of the predominantly German references. The work on the English edition was begun at the Ontario Veterinary College, Guelph, Ontario, Canada, and completed at the New York State Veterinary College, Cornell University, Ithaca, New York. At the latter institution generous financial support was received for the project through Dr. G. C. PoPPENSIEK, Dean of the Veterinary Faculty, from the General Research Support Grant of the National Institutes of Health, U.S. Department of Health, Education and Welfare. This made it possible to defray typing and editorial expenses and to obtain the valuable assistance of Mrs. ANTOINETTE M. WILKINSON, Ph.D., who greatly enhanced the idiomatic quality of the manuscript. It is a pleasure to acknowledge the interest and continuous support given this project by Prof. RoBERT E. HABEL, Head of the Department of Anatomy, New York State Veterinary College, and the friendly help received from the other members of his department. Thanks are due to Prof. STEPHEN J. RoBERTS, former Chairman of the Department of Large Animal Medicine, Obstetrics and Surgery, New York State Veterinary College, for his advice, often sought, on clinical questions; to Prof. C.]. G. WENSING, Department of Anatomy, Faculty of Veterinary Medicine at the University in Utrecht, Holland, for reading and commenting on the section on the Descent of the Testis and for contributing a figure; and to the many persons who in the past several years helped with typing, editing, and proof reading. The PAUL PAREY Publishing Company, particularly its co-owner Dr. FRIEDRICH GEORGI, is to be commended for undertaking to publish this English edition, and for its traditionally excellent and careful production. It is hoped that this work will be as well received as the original German edition, and that it will contribute to filling the need for a modern, comprehensive textbook of Veterinary Anatomy. Ithaca, New York, January 1973 W. 0. SACK * Schaller, 0., R. E. Habel and J. Frewein, Editors. Nomina anatomica veterinaria, 2nd. ed. Vienna, International Committee on Veterinary Anatomical Nomenclature, 1972.

Preface to the Second German Edition The favorable reception given again the second volume of our Textbook of the Anatomy of Domestic Animals and the need for a second edition of this volume after the relatively short interval of six years, strengthens our conviction that we have produced a book useful especially to students of veterinary medicine. The second edition differs from the first only in that minor corrections were made in the text and in some references to figures. In this connection we thank Dr. W. 0. SACK, Associate Professor, New York State Veterinary College at Cornell University, for his valuable suggestions. We also thank Dr. K. H. WILLE, Assistant at the Department of Veterinary Anatomy, Justus-Liebig-Universitat Giessen, for his suggestions and technical assistance during the preparation of the manuscript for this edition. No changes have been made in nomenclature*. It was decided to delay such a revision until the work of the International Commission on Veterinary Anatomical Nomenclature is completed and full agreement has been reached on the terms to be used in the future. Unfortunately, our friend and colleague, RICHARD NICKEL, did not live to see the new edition of the second volume which he helped to write with much enthusiasm and great skill. Again, sincere thanks are due to the PAUL PAREY Publishing Company for their excellent production of this edition of the second volume. We hope this book serves the purpose intended by the authors and meets the needs of the readers. Zurich and Giessen, April 1967 EuGEN SEIFERLE AuGUST ScHUMMER '' In the present first English edition the nomenclature conforms to the NAV (See also footnote to Preface to the First English Edition).

Preface to the First German Edition The second volume of our Textbook of the Anatomy of Domestic Animals follows the first volume after a longer interval than anticipated, because of the extremely timeconsuming preparation, especially in connection with the illustrations, and the everincreasing administrative burdens of the authors. As announced in the preface to the first volume, this volume, which is written by A. ScHUMMER and R. NICKEL, deals with the viscera of the digestive, respiratory and urogenital systems. In keeping with the basic plan of the work, each organ system and its principal functions are described first in a general and comparative chapter, followed by more detailed descriptions in shorter special chapters for the carnivores, pig, ruminants, and horse. The viscera are given especially extensive treatment, both in text and illustrations, because an accurate knowledge of the viscera of domestic mammals is fundamental to all branches of Veterinary Medicine. For good reason, an anatomy textbook is judged also by the quality and instructiveness of its illustrations. The 480 new illustrations presented here, many of which were carefully selected to show topographical relationships, are testimony to the skill and perception of our medical artists Miss VALERIE GuBE of the Department of Veterinary Anatomy, Justus Liebig Universitaet Giessen, and WALTER HEINEMANN and GERHARD KAPITZKE of the Department of Anatomy, Tieraerztliche Hochschule Hannover. We are also grateful to Dr. DIETMAR HEGNER for drawing several illustrations. We thank especially our faithful co-workers Dr. BERND VoLLMERHAUS, assistant at the Department of Veterinary Anatomy, Justus-Liebig-Universitat Giessen, and Dr. HELMUT WILKENS, prosector at the Department of Veterinary Anatomy, Tierarztliche Hochschule Hannover, who with great skill saw to the labeling of the illustrations, prepared the legends, and helped in many ways with the preparation of the manuscript. We are indebted to Dr. habil. KARLHEINZ HABERMEHL, Dr. KLAUS LoEFFLER, Dr. RuDOLF ScHWARZ, and HEINZ KoLBE for their assistance in reading the proofs. Sincere thanks are due to the PAUL PAREY Publishing Company, especially to Mr. FRIEDRICH GEOGRI, co-owner of the company, for his interest in the work, his understanding of the authors' intentions, and for the personal attention to this volume also, giving it their traditionally excellent and careful production. We hope that the second volume of the Textbook of the Anatomy of Domestic Animals will find as wide acceptance as the first. Zurich, Hannover, and Giessen, in the Fall of 1959. EuGEN SEIFERLE RICHARD NICKEL AuGUST ScHUMMER

Contents Introduction....... Body Cavities....... Thoracic Cavity and Pleura. Abdominal Cavity, Pelvic Cavity, and Peritoneum. Omenta and Mesenteries........... Peritoneal Folds Associated with the Urogenital Organs Bibliography: Body Cavities, Omenta, and Mesenteries.. Digestive System Mouth and Pharynx, General and Comparative. Oral Cavity. Lips.. Cheeks.. Gums.. Hard Palate Tongue Lingual Muscles. Hyoid Muscles. Sublingual Floor of Oral Cavity. Salivary Glands... Parotid Gland Mandibular Gland. Sublingual Glands. Pharynx.... Soft Palate.... Lymphatic Organs of the Pharynx (Tonsils) Deglutition........... Mouth and Pharynx of the Carnivores Oral Cavity... Salivary Glands. Pharynx... Tonsils.... Mouth and Pharynx of the Pig Oral Cavity... Salivary Glands. Pharynx.... Tonsils.... Mouth and Pharynx of the Ruminants. Oral Cavity... Salivary Glands. Pharynx... Tonsils.... Mouth and Pharynx of the Horse Oral Cavity... Salivary Glands. Pharynx.... Tonsils.... Bibliography: Mouth and Pharynx Page 1 2 4 6 11 17 18 21 21 23 25 25 25 27 31 32 36 39 41 44 44 44 52 52 56 57 57 58 59 60 60 60 62 62 63 64 64 66 68 69 69 69 71 73 73 74

XII Contents Teeth, General and Comparative Replacement of Teeth Types of Teeth... Dental Formula... Morphology of Teeth. The Teeth of the Carnivores The Teeth of the Pig.... The Teeth of the Ruminants The Teeth of the Horse Bibliography: Teeth.. The Alimentary Canal, General and Comparative Esophagus Stomach... Intestines... Small Intestine Large Intestine Anal Canal. Liver.... Pancreas.... The Alimentary Canal of the Carnivores Esophagus Stomach. Intestines Liver.. Pancreas. The Alimentary Canal of the Pig Esophagus Stomach. Intestines Liver.. Pancreas. The Alimentary Canal of the Ruminants Esophagus.... Ruminant Stomach......... Structure and Interior of the Ruminant Stomach Omenta Intestines Liver.. Pancreas. The Alimentary Canal of the Horse Esophagus Stomach... Intestines... Small Intestine Large Intestine Liver.... Pancreas.... Bibliography: Esophagus and Stomach Intestines..... Liver and Pancreas... Page 75 77 77 78 79 81 85 88 93 97 99 99 101 107 108 109 110 114 119 122 122 122 127 134 136 137 137 137 139 145 146 147 147 148 159 166 168 176 179 180 180 181 185 185 188 194 197 198 200 202 General and Comparative.. The Spleen of the Carnivores The Spleen of the Pig... Spleen 204 206 207

Contents XIII Page The Spleen of the Ruminants. The Spleen of the Horse Bibliography: Spleen Respiratory System General and Comparative. Nose.... Apex of the Nose.. Nasal Cavity.... Incisive Duct, Vomeronasal Organ, and Lateral Nasal Gland Nasopharynx.. Paranasal Sinuses.... Larynx.... Cartilages of the Larynx Ligaments and Articulations of the Larynx. Muscles of the Larynx.......... Laryngeal Cavity and its Lining...... Movements of the Larynx and its Cartilages Trachea.... Lungs.... The Respiratory Organs of the Carnivores The Respiratory Organs of the Pig The Respiratory Organs of the Ruminants The Respiratory Organs of the Horse Bibliography: Respiratory System Urogenital System Urinary Organs, General and Comparative Kidneys.. Renal Pelvis.. Ureter... Urinary Bladder Urethra... The Urinary Organs of the Carnivores The Urinary Organs of the Pig... The Urinary Organs of the Ruminants. The Urinary Organs of the Horse.. Comparative Anatomy of the Kidney Bibliography: Urinary Organs 208 208 209 211 211 213 216 219 221 223 225 225 230 234 235 236 238 240 247 254 261 271 279 282 282 287 288 288 290 291 294 295 298 301 302 Male Genital Organs, General and Comparative General Organization Testis..... Epididymis..... Ductus Deferens Coverings of the Testis and of the Spermatic Cord. Descent of the Testis Accessory Genital Glands............ Penis and Urethra............... Blood Vessels, Lymphatics, and Innervation of the Male Genital Organs Male Genital Organs of the Carnivores. Testis, Spermatic Cord, and Coverings Accessory Genital Glands. Penis.... 304 304 304 308 309 310 312 317 318 322 324 324 325 325

XIV Contents Male Genital Organs of the Pig.... Testis, Spermatic Cord, and Coverings Accessory Genital Glands...... Penis.... Male Genital Organs of the Ruminants. Testis, Spermatic Cord, and Coverings Accessory Genital Glands... Penis.... Male Genital Organs of the Horse... Testis, Spermatic Cord, and Coverings Accessory Genital Glands.... Penis.... Bibliography: Male Genital Organs Page 329 329 330 330 333 333 334 336 340 340 341 345 348 Female Genital Or~ans, General and Comparative General Organization of the Female Genital Organs Ovaries.... Tubular Genital Organs Uterine Tube Uterus.. Vagina... Vestibule.. Vulva and Clitoris. Muscles Associated with the Female Genital Organs Blood Vessels, Lymphatics, and Innervation of the Female Genital Organs Postnatal Changes in the Female Genital Organs Placentation and the Gravid Uterus.. Female Genital Organs of the Carnivores. Ovaries Uterine Tube........ Uterus.... Vagina, Vestibule, and Vulva. Female Genital Organs of the Pig Ovaries Uterine Tube........ Uterus.... Vagina, Vestibule, and Vulva. Female Genital Organs of the Ruminants. Ovaries Uterine Tube........ Uterus.... Vagina, Vestibule, and Vulva. Small Ruminants...... Female Genital Organs of the Horse. Ovaries Uterine Tube........ Uterus.... Vagina, Vestibule, and Vulva. Bibliography: Female Genital Organs Index.... 351 351 352 355 356 358 361 362 363 365 365 366 367 369 369 370 371 372 375 375 375 376 376 378 378 379 380 382 384 385 385 386 386 388 389 393

List of Abbreviations (The last letter of the abbreviation is duplicated to indicate the plural.) a. caud. com: cran. dext. dors. duct. ext. for. gl. int. lam. lat. Jig. arteria, artery l.n. lymphonodus, lymph node caudalis, caudal longitud. longitudinalis, longitudinal communis, common m. musculus, muscle cranialis, cranial maj. major dexter, right med. medialis, medial dorsalis, dorsal min. minor ductus, duct n. nervus, nerve externus, external proc. processus, process foramen prof. profundus, deep glandula, gland sin. sinister, left internus, interna supf. superficialis, superficial lamina transv. transversus, transverse lateralis, lateral v. vena, vein ligamentum, ligament ventr. ventralis, ventral References to Figures These appear in parentheses in the text, mostly in this form: (36/a). The number preceding the oblique dash refers to the figure; that which follows the oblique dash in italics refers to the part so labeled in that figure. Thus the notation (36/a) refers to part a in Figure 36. However, the notation (36, 37, 38/a) refers to part a in Figures 36, 37, and 38; whereas the notation (39; 40; 41/a) refers to Figures 39, 40, and part a in Figure 41, that is, part a refers only to Figure 41. Numerals and letters in italics but not preceded by a number, such as (a, 2), refer to parts in the figure whose number was last quoted. Color Plates Figures printed in color are combined on Plates and, with a few exceptions, are numbered consecutively with the text figures. They are distributed as follows: Fig. 132 Fig. 144-147 Fig. 162 and 163 Fig. 292-303 Fig. 351-353 Fig. 505-512 Plate I Plate II and III Plate IV PlateV-IX Plate X-XII Plate XIII next top. 96 next top. 104 next to p. 120 next to p. 224 next to p. 240 next to p. 352