6 le ae a! le - PRACTICAL ANATOMY OF THE RABBET 4 * AN ELEMENTARY LABORATORY TEXTBOOK IN MAMMALIAN ANATOMY ay at sabe e BY Al BENSLEY, Ph.D. Associate PW essor of Zoology in the University of Toronto ee | it +¢ os , y wu ie TORONTO: THE UNIVERSITY PRESS PHILADELPHIA: P. BLAKISTON’S SON & CO. ‘ 1910 ‘* a SMT H ne ON j At | CopyRIGuHT, CaNnapDA, 1910, By Tue UNIVERSITY PRESS PREFACE. The object of the present book is to set forth the chief facts of mammalian structure in an elementary, practical form; further, to use the anatomy of a typical mammal as a means of applying the more useful definitions of human anatomy and, so far as the limitations of a single type permit, also the broader conceptions of morphological zoology. On the practical side, its chief aim is to place before the student the materials necessary for a practical study of the type, rather than a descriptive account of its organization, though in some cases, notably in the treatment of the skeleton, it has been possible to follow a descrip- tive method without departing from the original plan. The inclusion of a section devoted to certain general aspects of the structure of the rabbit will, it is hoped, encourage the student to prose- cute his practical study with a more liberal point of view. As to the subject-matter of this section, its selection has been a matter of no little difficulty, and, doubtless, in many respects it might have been improved upon. One must feel, however, that the first question is not one of detail, but of general principle. Progress depends to a considerable extent on the ability to attack small problems with a large spirit. At the present time a vast amount of effort is being devoted to the planning of laboratory courses, and with increasing specialization it becomes more than ever the duty of the instructor to see that the student does not leave the laboratory, provided with a mass of detailed information, but with general conceptions as crude as when he entered it. As a laboratory type the rabbit has been made familiar to students through various zoological textbooks and especially through the “Zootomy’’ of Parker. The use of the animal, however, so far as one may judge, has not been as extensive as its general convenience would seem to warrant. It may, therefore, be of some value to direct atten- tion to this form by providing more facilities for its study. It is un- fortunate, in many respects, that no recent and adequate account of the anatomy of the rabbit is available, as is the case with other mammals used for laboratory study, the classic ‘“‘ Anatomie des Kaninchens’’ of Krause, published in 1884, being still the common source of information. Mammalian dissection is probably of most value to two classes of students, namely, medical or premedical students using it as an intro- duction to human anatomy, histology, or physiology, and zoological students using it as an introduction, or as part of the laboratory prac- tice of vertebrate zoology. For the latter class two aspects of the subject are especially worthy of consideration. One is the more or less detailed study devoted to a single type; the other, the study of a specialized type, the latter point being of more importance if the sub- iV PREFACE. ject is being used as an introductory one. In many respects the con- tinuous study of a single animal is a good corrective for the rough general kind of dissection as suggested by the zoological textbooks, and may be made to share the well-known merit of human anatomy as a laboratory discipline. Again, for the student who afterwards is con- cerned with vertebrate evolution, the study of a specialized type, such as a mammal, gives him, at the outset, something definite and concrete on which to base his conceptions of sequence. Primitive structure is of great value as a means of explanation, but the question, now as hereto- fere, is whether or not the study of primitive animals as a preliminary step represents the correct procedure from a laboratory standpoint. The more the student becomes interested in tracing sequence, the more he will be convinced of the necessity of stating his problem before he begins to solve it. The practical outlines on which the present book is based have been used for several years and in different forms in the laboratories of the University of Toronto. It would be difficult to make due acknowledg- ments to those colleagues and students who at one time or another have assisted in its preparation. Indeed, our chief obligation is to Professor Ramsay Wright, who, in establishing laboratory courses of this kind, has laid the foundation on which we have tried to build. B. A. BENSLEY. University of Toronto, January 3rd, 1910. CONTENTS HEN CIRER@ OUI IU@ING ae 8. soc RCS Sead nick Se Mana eS hia) 8 cy Wa, Woh SEN ie 1 Part I. A GENERAL CONSIDERATION OF THE : SLRUCIURE OF DHE RABBIE: ID GVSTONS “AND iene MOSS Hany camel uhs vals: ah b sh ee ene ak Soom cocleages Siieee heal eae 3 ENGERERE TATION ORS. RUCTURE 34 10 .\/c50s als se Bigs ie sateen ehas ee eine 4 BOGE OCTCAT se OSURION sons oe esne alk cites Fada So pen) ela 6 Gene Are wANAT OMG 2 Nac ole oF ital ool ae SD ee 8 Eoitivelial TMsswes oo fy 5 ess» cc 22s 2 ieee ee eae 9 Connective Tissues: ..........4... .. eee ee, ee 12 Muscular -Tissues....0.......002. 2 ee es 19 Nervous Tissues)... .. 2...) Se eee Pail TERMINOLOGY 3.0.6.3... . ee ee 23 THE GENERAL FEATURES AND GROUND PLAN OF THE ORGAN SYSTEMS 25 The Skeletal Systeme: 74. ype en a geo 2 30 The Nervous. SyStenige 245). eee Ng ae, 34 The Digestive System 7... ......... 7) se, he es 40 The Respiratory System..'...... ee 28 43 The Vaseular System." 0.0.0.0... a 43 The Urinogenital System ......)) 7p ee ee 46 The serous Cavities... .......0eene)) 49 REGIONAT SECTIONS)... 0... sip. os od I we Sa 2 Party II]. OSTEOLOGY OF THESRABBIA. GENERAL DIVISIONS OF THE SKELETON. ..- 38. Gee. a . 0 Seve ee 68 THE VERTEBRAL COLUMN/i oa :....... SEL es ee 68 SEE RBS. . 5. So. Sao eeS”. os enn enemete coer lL. 73 SEMEL ODERINTIM , . 2.2.0 hill MM. Plslere Gila done Me MRS 5: nie Ae. 74 THE OMEEETON OF THEPEIEAD we)... 2 . sale an ect. age tee co fe MimesSkiull asta W holet wea Sa ee. lt 15) RWhe«Bones offre Sk ilie |) ae ee OS |. 85 The Hyoid Apparatus. . TR oe a. . ae! 97 THE SKELETON OF THE Aeon Tite eS ee ae. 98 THE SKELETON OF THE POSTERIOR LIMB vi CoNnTENTS. Part III. DISSECTION OF THE RABBIT. 1 EXTERNAL. PRATURES ¢ 5 sic oie he ek ener eee reas te eel tone 110 See) ABDOMINAL: WAT Lc a etec ce ane ect renege Ome Oe vangs Sau 113 3. THE STOMACH AND OPLEEN........ ati Maca WP one, SNS SO end ee 116 ARTE LIVER: «cep. ac Oe i ee ence SE 120 Re RITE LNTESTINES © 's couse abe ek ot neh eee ee rae ete 122 Gs) RHE URINOGENITAL SYSDEM ese. eine as tke eee eae IPA 7. Tue ABDOMINAL AORTA, INFERIOR VENA CAVA, AND SYMPATHETIC AT RAINES skh eee See Ra Tea ec 133 S. /Pam ANTERIOR LIMBS... 629 Socrates 135 Our hae POSTERIOR LIMB: .hncoild Gece a le eee ee Co eee 144 1OMGbE KIEAD AND NECK: 5.56) ee hee eee noe ee ne tS [ete SIE MIO RA Xia se ooo 0 Sage ee cen eee ee 172 12. THE VERTEBRAL AND OCCIPITAL MUSCULATURE............... 180 135 MHEVCENTDRATENIERVOUS OVSTEM), . 5). cf) ssn less oeereteie tomate 184 APPENDIX. DIRECTIONS FOR THE PRESERVATION OF MATERIAL.... 194 INTRODUCTION: Salaboratory exercise, the anatomical study of an animal is largely a matter of applying a certain practical method of exposition, the student’s attention being concentrated on those facts which may be made out by direct observation. For this reason, and also because continuity is a prime consideration, various important aspects of structure are of necessity left in the background. It is to be con- sidered, namely, that in studying the structure of any organism, the final object is not simply to determine in what its structure consists, that is, its anatomy in a restricted sense, but also to understand what the latter signifies when considered either as functional mechanism or, in general, asa product of the various factors underlying it. Every organism reflects in its structure the operation of a variety of influences, and consequently one cannot form an adequate conception of animal or- ganization without considering it from various points of view. In many respects, as indicated below, the interpretation of structure is not simply a matter of what is to be found in a given form, but also of what the latter represents in comparison with others. Assuming, as in the present case, that the student is principally occupied with the routine of a type dissection, the question of how far he may go afield in the consideration of accessory facts is one which must be determined by his own inclinations. His first need, one which the present book endeavours to fill, is to understand the sources of informafion. Part I, therefore, the subject matter of which has been selected especially with reference to the student who has had no previous experience in the biological sciences, is designed to indicate some of the possibilities in this connection, and also to serve in other ways as an adjunct to the practical account of the structure of the rabbit as outlined in Parts II and III. With the introduction obtained in this way by using the rabbit as an object lesson the student should be able to extend his informa- tion independently, using for this purpose special textbooks in the respective subjects. Ee Aisles le A GENERAL CONSIDERATION OF THE STRUCTURE OF THE RABBIT. DIVISIONS AND METHODS. Biology, the science or study of living organisms, includes several related sciences, the chief of which are Anatomy, the study of organized structure; Physiology, the study of function; and Embryology, the study of development. Anatomy, or Comparative Anatomy, the latter referring to the comparative study of organisms, and Embryology are also considered as divisions, or as practical methods, of Morphology, the general science of the evolution of form. The term “Anatomy” was originally applied to the dissection or study of the human body, and is still considered as referring more especially to the latter. Even in the early stages of biological science, however, the use of the term was extended to organisms generally; and afterwards, chiefly as a result of the introduction of the microscope as a new method of examining structure, it attained its present compre- hensiveness as a term applying to the study of structure generally. It has been found convenient, especially in human anatomy, to distinguish as Gross Anatomy, the study of that kind of structure which is displayed by dissection, or is revealed by naked-eye appearances, and as Microscopic Anatomy, the study of finer structure through the ap- plication of the microscope; or, again, to distinguish as Special or Des- criptive Anatomy, the study of the particular features of the organs of the body, and as General Anatomy, the study of its more fundamental composition. General Anatomy is practically equivalent to Histology, the latter considering the body from the point of view of the structure and arrangement of its cells and tissues. These distinctions are of interest in the present case chiefly as defining more exactly the practical method and the kind of structure to be considered. Thus, dissection is to be recognized as a method of displaying structure of a gross and special kind. It consists in the orderly exposure and displacement of organs with the object of observing their features and relations to surrounding parts. The plan is essentially one of analysis, since conceptions of structure are based on the recogni- tion of differences, the latter being estimated by various features, such as form, color, texture, or position. On the other hand, because of the class of structure with which it deals, dissection is also to be recognized as a preliminary method in comparison with various others involving the use of the microscope. A ANATOMY OF THE RABBI’. THE INTERPRETAGION OF SERUCT URE. Gross structure is, in a sense, only the outward expression of the finer microscopic structure underlying it, the latter being the true basis of the body. This refers not so much to the individual features of organs as to the relation existing between their appearance as gross objects and their tissue composition. Since this relation is more fully discussed below under the head of general anatomy, it need only be mentioned here as an element in the interpretation of structure as viewed from the gross standpoint. All structure, however, may be con- sidered from two points of view—physiological, and morphological. The former is more easily understood. All parts of the body are constructed on a basis of use or function; and although the various processes and activi- ties of the body are more properly considered under the head of physio- logy, a thorough conception of the anatomy of an organ is usually to be gained only by a consideration of its particular role in the general economy. The morphological aspect of structure concerns various features of form and arrangement which, although they have been developed on a basis of utility, cannot be explained directly on that basis, because the factors controlling them lie outside of the body of the individual, and are such as have operated only through a long series of gradually changing conditions in the evolution of its type. As applied to a particular animal, the morphological method consists in explaining its adult structure by reference either to its embryonic development, or to the equivalent conditions in lower existing, or perhaps fossil, forms. The recognized principle of embryology is that known as the Law of Re- capitulation. It is based on the general observation that the definitive structure of an organism is attained through a series of embryonic stages, in which it not only develops from a simple or ground type to a more complex condition, but also reflects in passing the features of lower, and presumably its own ancestral, forms. That of comparative anatomy depends on the comparison of higher, specialized animals with lower, or generalized ones, the latter being assumed, in one feature or another, to have remained in a backward or primitive state of specialization, and therefore to reflect in such features.a low grade of structure of a kind possessed by the ancestors of existing higher forms. These relations form a basis for the comparison of the embryonic development of or- ganisms with the evolution or history of the groups which they represent, the former being distinguished as ontogeny, the latter as phylogeny. The interpretation of the adult structure of an organism is a matter of distinguishing its more general features from its more special ones, the former being in all cases those to which the ontogenetic and phylogenetic principles are especially applicable. All characters of animals have an evolutionary basis, the general INTERPRETATION OF STRUCTURE. Sal nature of which is easily understood, although the process by which they have been developed is still a matter of uncertainty. In comparison with one another, animals present certain resemblances and differences— diagnostic features, which are used as a basis for classifying them into major and minor groups. In many cases characters of resemblance have been shown to be secondary, and are hence described as convergent. In some of these the resemblances are of a gross type, and the structures are described as analogous; in other cases they are exact or homoplastic. Asa rule, however, characters of resemblance are broad marks of affinity, comparable to those seen on a small scale in human families, or in human races, and determined as in the latter cases by heredity. The chief basis of comparison of animals with one another is the general assumption that structures which are similar or identical are homogenous—of com- mon origin. On the other hand, their differences are chiefly marks of diver- gence in evolution. Although it is conceivable that many of the internal features of animals are the result of a general progressive development, more conspicuous in comparison with those of primitive types, the majority of their differences are such as have resulted from adaptive modifications of structure, by which they have become differ- ently adjusted to the particular conditions of their accepted habitats. Adaptation is one great factor in modifying animal form, producing first divergences, as between one type and its contemporaries; although such features may afterwards become settled in particular groups, and thus appear for these as primitive, general, or group- -characters. Adaptation, in other words, is not a matter of present conditions only, of fixed environment, or an environment of a general or special kind. The rabbit as a gnawing animal or rodent, for example, is also an air- breathing, walking vertebrate, and shares these larger and also more ancient features with many other vertebrates of otherwise different kinds. It is customary to include under the term specialization all those features in which an organism may be shown to be more highly modified in comparison with another type. If the latter is an ancestral type, ora lower form exhibiting ancestral features, its more primitive features are said to be prototypal, because they indicate the form from which the higher modification has been derived. Such comparisons not only reveal the fact that different animals are specialized in different degrees, but also show that a given form may be greatly specialized in some respects and primitive in others. Moreover, it is to be considered that animals are at the present time, as they have been in the past, more or less changeable, or plastic types. Some of the most interesting features which they exhibit depend on the circumstance that the adjustment of structure which is rendered neces- sary by the opposing effects of heredity and specialization 1 is not exact or immediate. Thus, it is not difficult to find in any specialized animal, in addition to those organs which are functional or in full development, others which are retrogressive in character and reduced in size. It is also to be assumed, although difficult of proof, except where the inter- mediate member of a known series is being considered, that there are also organs which are sub-functional or progressive. 6 ANATOMY OF THE RABBIT. ZOOLOGICAL, -POSIRION: It will be evident from the foregoing statement that every specialized animal possesses in its organization a vast’ assemblage of features which, if referred to their proper categories, are found to represent many grades of morphological value. In so far as the adult structure of a particular form is concerned, it is possible to consider’them anatomically without discrimination; but, on the other hand, if they are to be explained, it is necessary to proceed on a basis of function, embryonic development, or evolution. The study of an animal as a type or representative of a group, however, concerns only in a general way the features common to its various members, since the majority of features present in any animal are of minor importance, and as such are significant chiefly as indicating the developments which may take place inside it. The question of what an animal is actually representative is a matter of comparison with other forms, in other words, of its zoological position. This is expressed through the medium of classification, the latter being arranged to indicate, so far as is possible, the relationships of organisms to one another. In this connection the following statement of the zoological position of the rabbit may be found useful; and it may also be considered as illustrating, through the comparison of this animal with allied forms, some of the more general characters of animals as outlined above. The domestic rabbit is represented by several races, of which the common variously-colored forms, long-haired Angoras, Lop-Ear Rabbits, and Belgian ‘“‘Hares”’ are more familiar. They are all descen- dents of the wild rabbit (Oryctolagus cuniculus, Lepus cuniculus) of Europe. The latter is thought to have belonged originally to the countries bordering the western portion of the Mediterranean, but its distribution has been greatly extended northward and to other continents through human agency. The family Leporidae contains a large number of closely related species formerly included in the single genus Lepus. They are variously known as hares and rabbits, but the latter designation is considered to apply more exactly to the European rabbit and its domesticated races, the others, with one or two exceptions, being more properly described as hares. The more familiar species include the North American Cotton- Tail (Sylvilagus floridanus, Lepus sylvaticus), and the Prairie Hare or Jack-Rabbit (Lepus campesiris); the European Common_ Rabbit (Oryctolagus cuniculus), and Hare (Lepus europeus). The two European species differ in several well marked features, which form the basis of the accepted distinctions between hares and rabbits. The rabbit is dis- tinguished by its shorter ears and less elongated hind limbs; also by its burrowing habits, and by the circumstance that the young are born in a blind and naked condition. The hare is more nearly a running or cursorial type, and is distinguished by its longer ears—which, moreover, are tipped with black—longer hind limbs and prominent eyes. Unlike the rabbit it does not burrow, but inhabits only an open “form,” and the young when born are clothed with hair and able to see. ZOOLOGICAL, POSITION. fi The various species constituting this family are distinguished by several features, including the imperfect development of the clavicle, longer ears and limbs, and the presence of a distinct although greatly reduced tail, from the Picas or Tailless Hares (Ochodontide) of the mountain- ous districts of Central Asia and of North America (Rocky Mountains). The two families are allied, however, in the possession of a common feature, namély, the presence in the upper jaw of a second pair of incisor teeth. This feature distinguishes the sub-order Duplicidentata from that of the Simplicidentata, the latter containing the majority of rodents and embracing all forms with a single pair of upper incisors. The mammalian order Rodentia, to which the family belongs, con- tains a very large assemblage of forms—the Squirrels, Marmots, Cavies, Beavers, Mice, and Porcupines being among the more familiar. This order is distinguished by the modification of the anterior incisors in both upper and lower jaws to form chisel-like cutting organs, the teeth having their enamel layer disposed chiefly if not wholly on their front surfaces, so that they remain in a permanently sharp condition. This modification is associated with an extensive obliteration of intermediate teeth, com- prising posterior incisors, canines, and anterior premolars; also with elaboration and often great complexity of the remaining premolar and molar teeth, the lower jaw, and, indeed, the parts of the skull generally. Characteristic of these animals is the extension, both for- ward and backward, of the jaw-musculature. The articulation of the lower jaw exhibits an elongated articular process fitting into a corresponding -longitudinal fossa on the skull, the jaw being able to move forward and backward in addition to vertically and from side to side. The teeth are further arcuate in shape, and are provided with open roots, so that their growth is not limited, as it is in the majority of mammals. The rodents are in many particulars primitive types. For example, they tend to retain the five-toed (pentadactyl), plantigrade foot, characteristic of primitive mammalia and, indeed, of terrestrial vertebrates, and exhibit also unelaborated cerebral hemispheres in the brain. In other respects, however, as in the rodent characters above-mentioned and in the elaboration of the intestine, especially the caecum, they exhibit the characters of highly specialized herbivores. Like all higher or placental mammalia (Placentalia), the rabbit is viviparous, the young being retained through a period of gestation in the maternal uterus, to the wall of which they are attached by a vascular con- nection, the placenta. In this feature the placental mammalia differ from the marsupial mammalia (Marsupialia) of Australia and South America, the latter being viviparous, but, with one exception, without placenta; also from the egg-laying mammalia (Monotremata) of Aus- tralia, the latter being oviparous, like the majority of the lower, reptilian forms. These three sub-classes of mammals are united, however, by the common features of the class Mammalia. They are warm-blooded animals, provided with a complete double circulation, and with a hairy investment for the surface of the body. In all, the young are nourished for a time after birth through the secretion of modified cutaneous, milk- producing, or mammary glands. 8 ANATOMY OF THE RaBsir. Many of the more general features of the rabbit are such as are not recognized by group designation, but yet are shared with other terrestrial vertebrates, including mammals, reptiles, birds, and, in part, amphibians. This refers to the development of the lungs and associated respiratory tracts, both the true respiratory tracts and the accessory respiratory passages traversing the skull; further, the loss of the branchial or fish-type of respiration and the new disposition of the branchial structures; the development of a tri-segmented type of limb with a full complement of muscles, and originally a pentadactyl, plantigrade foot; for support of the body and for locomotion. The regional differentiation of the vertebral column, especially the mobility of the neck, the free occipital articulation, and the definition of the sacrum, the latter associa- ted with the elaboration of the pelvic girdle, are all features of general significance in the terrestrial vertebrates. Finally, the rabbit agrees with other members of the phylum Chordata in the possession of a ground-plan underlying the most general features of its crgans, and the position, arrangement, and plan of development of its organ-systems. All Vertebrata or back-boned animals possess an axial skeleton formed by the segmented vertebral column. Ina very com- prehensive way they possess as chordates a still more fundamental axial support, the notochord, the latter being an embryonic structure except in the lowest chordates. In a more restricted sense, as Craniota, they possess an organized head region with differentiated brain, special sense organs, and enclosing primary skull. They possess a series of branchial (branchiomeric) structures,appearing either in the adult condition, as in fishes, or as part of the underlying plan in the embryonic condition; and they add to their general features in the arrangement of the organ-systems the further feature of transverse segmentation (metamerism) of a considerable portion of the body. | GENERAL ANATOMY. Although in every respect a continuous structure, the body is differ- entiated into a large number of parts, or organs, the latter being more or less individual in form, composition, or function. Organs are arranged for the most part in systems, each of which is concerned with some general or fundamental function, to which several organs may contribute. In a more general way the body may be considered as an assemblage of tissues. The latter are layers or aggregations of similarly differen- tiated cells. They are of several different kinds, and are variously associated in the formation of organs. Being structures of an inter- mediate position they may be considered either as organ companents or as products of specialized cells. As a body-unit a cell consists of a small mass of living protoplasm, containing a central body, the nucleus, and surrounded or enclosed on its free border by a cell-membrane. The nucleus is a highly organized body, having an important function in the reproduction of the cell and EPITHELIAL STRUCTURES. 9 also in its general activity of metabolism. It contains a characteristic formed material, chromatin, and frequently also a minute spherical body, the nucleolus. The chief features of atypical cell are illustrated in the accompanying figure (1) of the developing ovum, the latter being a single cell, noteworthy for its large size, and also one in which the external form is not greatly modified, as it is in the majority of the cells of the body. Its enclosing membrane, the zona pellucida, by which in its natural position in the. ovary it 1s separated from the surrounding follicular cells, is considered to belong in part to the latter. As fundamental living matter, protoplasm possesses certain properties on which the functions of the body ultimately depend. Considered collectively, these functions are not so well illustrated in the higher or multicellular organisms, in which particular functions are assigned to particular cells,-as in the lower unicellular organisms, in which all func- tions are discharged by a single cell. In simple or protozoan animals the protoplasm is seen to be capable of ingesting food-materials, of discharging waste, of changing its form, and of reacting in one way or another to stimuli arising outside of the body. Moreover, the protozoan cell is capable of giving rise to new cells by division of its substance into two parts, which process originates in the nucleus, and is associated at some stage, usually at least, with union or conjugation of parent cells. All the cells of the body of a multi- cellular organism are products of a single cell, the fertilized egg, but the latter is a product of fusion of two primary elements, the spermatozoon of the male parent and the ovum ee ik Developing ovum of the rabbit- a rom a section of the ovary: chr., chromatin; of the female. The fertilized egg n.m., nuclear membrane; Dates cells of _the does not exhibit the functions of a esis) eee Tallhat: ce esti) Te one-celled body, but possesses the potential of these functions, and the latter appear, to a large extent individually, in the differentiation of its division products into special- ized tissue-elements. The primary tissues of the body are of four kinds—epithelial, con- nective, muscular, and nervous. To these—the fixed tissues—are to be added the fluid substances, blood and lymph, in which the cell elements, the red and white corpuscles, or in the latter case the white elements alone, are suspended in a fluid medium. 1. EPITHELIAL TISSUES. Epithelial tissues are distinguished chiefly as surface investments, such as those of the exterior of ‘the body, the interior of the alimentary canal, the lungs, the respiratory and accessory respiratory tracts, and the 10 ANATOMY OF THE RABBIT. ducts of the urinogenital organs. In all epithelia the cellular feature is a prominent one, and it is largely for this reason that as lining membranes they are not conspicuous in gross structure. They are noteworthy, however, for their products, the hairs and the various kinds of secreting organs or glands. The epithelium of the skin (Fig. 2, ep.) is known as the epidermis or scarf-skin. Although composed of several layers of cells, it forms an exceedingly thin membrane, extending over the surface of the body and connecting at certain points with the epithelia of the internal surfaces. It is supported by a layer of connective tissue which forms the true skin or corium. The epithelium of the internal surfaces forms the chief portion of the mucous membranes, and in the greater portion of the alimen- tary tract the epithelial layer is asso- ciated with a thin layer of smooth muscle to form a mucous _ tunic (Fig. 15, t.ms.)- The coating of hairs on the surface of the body, the presence of which is a notable mammalian feature, is a protective investment arising from the epidermis. A hair is produced by the modification of the central por- tion of an ingrowth of the epidermis, termed the hair follicle (Fig. 2, f.). The latter contains at its base a small elevation of the underlying vascular connective tissue, the hair papilla, through which the structure is nourished. On the general surface of the body the hair follicles are arranged in groups (Fig. 4), and on the lips certain large follicles are set apart for the production of the greatly enlarged sensory hairs or vibrissae. Connected with the hair follicles are thin strands of smooth rg ee muscle, the arrectores pilorum (Fig. 2, gots, & Fem, a section of the upper lip ofa ap) They are placed,in the broad the hair (a.p.!, is that of the adjacent fol. angles formed by the inclined follicles licle); c¢.t., connective tissue of the corium; 32 an o ep., epithelium; f., hair follicle; s. hair With the corium, and their contrac- Shalt ise see Oe, Gmereiaiad tion throws the hair into a more nearly erect position. Epithelial glands are ingrowths of the general layer, the cells of which become greatly modified as secreting structures. The lumen or cavity of the gland, in most cases greatly complicated through the division of the gland substance, is connected with the general surface by a duct which thus serves to carry away its secretion. In some cases the con- nection of a gland with the epithelial surface is embryonic, and in the EPITHELIAL STRUCTURES. 11 adult condition the gland is found separated from the epithelium from which it was originally formed. This condition is represented by the thyreoid and thymus glands of the neck and thorax respectively. In other cases the secreting element is a single cell, the latter thus repre- senting a unicellular gland lying directly in the general layer. The mucus-secreting goblet cells of the intestinal wall are structures of this nature. The majority of multicellular glands conform to one of two types, namely, the tubular gland, in which the secreting portions are of uniform calibre, and the acinous or alveolar gland, in which the secreting portions are sacculated (Fig. 3, A). Both types occur in simple, little branched and greatly branched conditions. Cutaneous glands of two types are commonly present in mammals in association with the hairs, namely, sudoriparous or sweat-glands, which are glands of the tubular type, and seba- ceous glands, which are of the acinous type. In the rabbit, glands are absent from the general surface, but are found in special situations, as, for example, in con- nection with the hair follicles of the lips, the internal surface of the ear, and the external genital organs. The inguinal glands comprise both tubular and acinous portions. The mammary glands of the female are greatly modified cutaneous glands of an acinous type. Apart from the mucus-secreting cells of the general epithelium, the glands of the alimentary canal comprise the important but less elaborated glands of the wall, such as the gastric glands of the wall of the stomach, and the greatly elaborated, out- standing glands which lie beyond the wall and are connected with the interior of the canal only through their ducts. The latter comprise the oral glands, the liver, and the pancreas. The oral glands include chiefly the submaxillary, parotid, sublingual, and Fic. 3. A, Diagram of a multi- infraorbital glands—conspicuous structures cellular gland: al. aye oeeina te in the dissection of the surrounding por- tubular type. _B, The pancreatic duct tions of the head and neck. Their ducts 322, 2ssociated portion ot ic al communicate with the cavity of the mouth. There is a close association between the epithelia of the surface of the body and the nervous tissues, arising from the circumstance that they are derivatives of a common embryonic layer, the ectoderm. In the adult we may distinguish as sensory epithelia special aggregations of cells lying either in a deep or superficial position, and associated more or less closely with the central nervous system. They comprise the olfactory epithelium of the nasal cavity, some of the cells of which are true nerve cells, the 2 ANATOMY OF THE RABBIT. gustatory epithelium of the tongue, and the auditory epithelium of the membranous labyrinth of the ear. The retina—the nervous portion of the eye—is a modified portion of the central nervous system. As linings of surfaces, the ordinary epithelia may be distinguished from certain special coverings of internal spaces, the endothelia and mesothelia. The latter consist microscopically of thin pavement-like cells. They differ from epithelia in origin, being formed, not in con- nection with originally free surfaces, but in relation to spaces of the mesoderm or intermediate layer of the body. Endothelia form the linings of bloodvessels and lymph canals, while mesothelia are the chief layers of the smooth, moist serous membranes which line the peritoneal, pleural, and pericardial cavities. 2. CONNECTIVE TISSUES. The connective tissues form the supporting elements of the body. As ordinary connective tissues they serve to connect organs or parts of organs together, and as skeletal tissues they provide the rigid framework or skeleton from which all soft parts of the body are suspended. They are distinguished by the presence of two main components—the cell basis, and the intercellular substance or matrix. The cellular portion is formative, and is much more conspicuous in the embryonic than in the adult condition. All ae hea connective tissues are products of an em- bryonic tissue, the mesenchyme (Fig. 20, ms.), which consists of branched cells con- nected together by their outstanding pro- cesses. Through the activity of the cells there is formed an intercellular material | consisting either of a homogeneous matrix, mem, OF more frequently a matrix containing formed elements of a supportive nature. Fic. 4. From a section of the skin of the side of the body of an adult rabbit. ™* about 10; showing the grouping of the hair-follicles: co., A. ORDINARY CONNECTIVE TISSUES. corium; ep., epithelium; m.c.m., papa eC coats name eS In the adult condition the ordinary connective tissues, with few exceptions, consist of the cell basis with two kinds of fibrous elements, the white, and yellow fibres (Fig. 5). White fibres are elements of great strength. They ane ime peiameed fibres which do not occur independently, but are joined in a parallel fashion with one another, forming in this way fibre bundles of larger or smaller size. The yellow fibres are of greater diameter. They branch and anastomose, but are not asso- ciated to form bundles. They also differ from white fibres in being highly elastic. The tissue produced in this way is known as fibrous connective tissue. It occurs in several forms according to the relative CONNECTIVE TIssuE STRUCTURES. IS concentration of the two kinds of fibres or the admixture of other materials. The commonest kind of fibrous tissue in the adult is that described as areolar. It is characteristic of the sub- cutaneous tissue (Fig. 4) which connects the skin with the body; but areolar tissue occurs also in various positions where it has a similar function of joining structures loosely together. Subcutaneous connective tissue is a white material, the peculiar appearance and properties of which are due to the fact that the two kinds of fibrous elements are arranged in a loose felt-like network (Fig. 5). When stretched it is found to yield up to a certain point, beyond which it is tough and resistant. It tends to regain its original shape when the tension is removed. _ Fic. 5. Areolar connective tissue Fibrous connective tissue may be Poe aaraiouines ce den ee. greatly modified through the concentration eet aio diecedbe, Obetther one of thetnbmmmselemicnts. Con- centration of white fibres is most common. This condition is illustrated in the thick connective tissue layer forming the true skin or corium (Fig. 4), but is more conspicuous in the glisten- ing white tendons (Fig. 6) by which muscles are attached to bone surfaces, in the ligaments uniting bones with one another, and in the thin, broad aponeuroses which serve for muscular attachment. The structures known anatomically as fasciae are special sheets of connective tissue covering chiefly individual muscles or muscle groups. Fat or adipose tissue is a soft form of con- nective tissue in which the cells are greatly enlarged and contain fatty material in the form of globules. It tends to occur in certain situations and in particular association with the bloodvessels, but otherwise is found in situations where areolar connective tissue might be expected to occur. Special fat masses, sometimes distinguishable by their darker coloration, occur at the side of the neck in the rabbit. In the foetus (cf. Plate VI) large masses of vascular connective tissue are found in this situation, and are probably similar in origin to the storing or hibernating glands of certain other mammals. Coloration or pigmentation of certain portions ee see of the tendon of origin of of the body, especially of the skin and hairs,“and ~ the biceps muscle: mf,, muscle fibres ending on the of the retina, ciliary body, and iris of the eye, is due tendon; t., fibrous con- 17 iss } ten- partly to the presence of special connective tissue aa Ge ne Se 14 ANATOMY OF THE RABBIT. cells, chromatophores, and partly to the presence of pigment granules in epithelium. The absence of pigmentation in animals belonging to species normally colored—albinism—is indicated by the white coloration of the hair and by the pink coloration of the eyes, the latter being due to the circumstance that the bloodvessels of the vascular tunic are not concealed by pigment. B. SKELETAL TISSUES. The skeletal tissues are solid forms of connective tissue which, on account of their more permanent shape, are better adapted to form a support for the body. They are —...m. of two kinds—cartilage and bone. ED Peete Simple or hyaline cartilage eo @&GF (Fig. 7) is asemi-solid and some- ; eset ee © what resilient material of a bluish ae @ ES or pearly coloration. It consists of aoe aes “1 a homogeneous matrix in which S. | = — the icellsare imbedded ahevecelts are distributed singly, or more often in groups of two to four, Fic. 7. Hyaline cartilage: c.c., cartilage each group being contained rhoy. cell; 1., lacuna; m., matrix. small oval space, the cartilage lacuna. The size of the spaces, and also their distance apart, is subject to great variation. The addition of fibrous elements, usually of white fibres, produces a modification known as fibrocartilage. White fibro- cartilage occurs in certain situations, as in the symphysis of the pelvis, or in connection with the interarticular menisci and at the capsular margins of the joints. In the adult skeleton cartilage is present only in small amount. It forms the articular surfaces of joints, the ventral portigns or eee costal cartilages of the ribs, and a pigs portion of the nasal septum; it is aul also found uniting the basal bones nt of the skull. In the embryo, how- ae ever, it forms the entire skeleton, malice: with the exception of a small por- tion which, as described below, is formed of membrane bone. In the Fic. 8. Intramembranous bone; from a 5 ground section of the rabbit's femur: c.l., course of development, except in circumferential lamellae; h.c., Haversian the situations indicated, the carti- canal; h.J., Haversian lamellae; i.l., inter- - stitial lamellae; lac., lacuna. lage is replaced by bone. Bone is a compact, resistant, but yet somewhat elastic tissue, possess- ing much greater strength than cartilage, and therefore forming a more perfect skeletal support. As indicated below, its appearance as a tissue differs somewhat according to its mode of formation. The more typical structure (intramembranous bone) is illustrated in the accompanying figure (8) of a ground transverse section of the dried shaft of the femur. eae a SKELETAL STRUCTURES. 15 The bone materials are deposited in layers or lamellae, which are com- parable to highly modified white fibres of fibrous connective tissue." The lamellae enclose between them greatly branched spaces, the lacunae, in which in the natural condition the bone-cells are accommodated. On account of certain differences in development, the lamellae are disposed in different ways. Some are disposed concentrically around longitudinal spaces, the Haversian canals, forming in this way the so-called Haversian systems. The canals are occupied i in the natural condition by bloodvessels. Others of them, the interstitial lamellae, are disposed in a somewhat parallel fashicn between the Haversian systems; while a third series, the circumferential lamellae, are disposed in a parallel fashion with respect to the periphery or the internal surface of the bone. In the natural condition the bone is enclosed except on its articular surfaces by a layer of con- nective tissue, the periosteum. During the period of growth this membrane contains large numbers of bone-forming cells, the osteoblasts, through the activity of which the deposition of the bone lamellae is accomplished. EO It is only in a few cases that the bones of the skeleton may be looked upon as solid structures— fa as a rule they consist of a fairly thin shell of hard | or compact bone surrounding a central mass of spongy or cancellous bone. This arrangement is ho ee one of great mechanical strength, combined with if || lightness, and at the same time provision is made | in the interior of the bone for bloodvessels and marrow spaces. Thus in a long bone (Fig. 9) the central portion or shaft consists of a cylinder of com- pact bone surrounding an extensive space, the marrow-cavity. The ends or extremities consist each of a thin shell of compact bone continuing that of the shaft and surrounding a mass of can- cellous tissue. In the short, flat, or irregular bones of the skeleton no continuous marrow cavity 1s formed. In point of origin bones are of two chief kinds Fic. 9. Divided femur —membrane or derm bones, and cartilage or cnins a ere replacing bones, butia third groupmic Tepsesemted) ySpr ty eee vishysia! by the tendon or sesamoid bones which are _ s., shaft. developed in the tendons of muscles. The mem- brane bones are few in number. They comprise the roofing and facial bones of the skull, most of which are distinguished by their flattened, tabulate character. They afte formed in connective tissue membranes, and although they sometimes contain cartilage they are not formed on a cartilage ‘basis. Moreover, they are elements lying in a superficial position w ith respect to the skeleton proper. They owe this condition, and also their designation as derm bones, to the circum- stance that they represent surface plates which in lower ‘vertebrates are associated with the skin. 16 ANATOMY OF THE RABBIT. Cartilage bones, on the other hand, are the characteristic elements of the skeleton. They are known as replacing bones because they are formed on a cartilage basis, the latter, as indicated above, representing the primary cartilage skeleton of the embryo. In the course of develop- ment, except in certain regions where the cartilage persists throughout life, the cartilage material is replaced by bone, which tends to surround and also invade it. The way in which bones are formed on a cartilage basis serves to explain many peculiarities of the adult skeleton. In the embryonic condition the cartilage rudiments are associated to form a complete but primitive skeleton. In many cases the replacement of these elements by bone is not direct, certain readjustments being necessary, both for pur- poses of growth and on account of the much more special functional requirements of the adult skeleton. In the embryonic condition the cartilage rudiments are enclosed by a connective tissue sheath, equivalent to the periosteum of a bone, but described as the perichondrium. The osteoblasts of this layer are concerned with the formation of bone material, both in the interior (endochondral bone) and on the surface (intramembranous bone). The forma- tion of endochondral bone is preceded by certain changes which take place in the interior of the cartilage. In the latter, in certain areas, known as the centres of ossification, the matrix becomes partly dissolved, the cells en- larged and ultimately broken down. These changes are associated with a Fic. 10. Outline sketch of the proximal deposition of calcareous material, or end of the femur of a youn animal “<. calcification, by which the portion of the femur. The accessory epiphyses are for cartilage undergoing transformation is the great (tr.ma.). lesser (tr.mi.), and Oo S S) g third (tr.t.) trochanters. temporarily strengthened. Into this area the active cells of the perichondrium are carried through the agency of vascular ingrowths, the periosteal buds, and the result of their presence is the deposition of bone material in association with the remaining portions of the matrix. This condition is partly illustrated in the distal epiphysis of the humerus shown in Fig. 12, the figure being that of a vertical section of the elbow-joint of a four-day-old rabbit. In the long bones the formation of the first or main centres of ossifica- tion takes place in the shaft, and there are formed afterwards accessory or epiphysial centres for the extremities. A divided extremity, such as the proximal end of the femur (Fig. 10), may possess several such centres —a principal one for the chief epiphysis or actual extremity of the bone and several subsidiary centres for its outstanding processes. In the shaft the formation of endochondral bone is of short duration. Through the activity of the osteoblasts lying directly in the perichondrium, or later the periosteum, a process of formation of intramembranous bones goes on during the whole period of growth, and the result of the peripheral tr. t SKELETAL STRUCTURES. 17, deposition of bone lamellae is, as described above, that the transverse diameter of the bone is greatly increased. The enlargement of the marrow cavity, with which this is associated, is produced by the absorp- tion of bone from the interior. In young animals both the epiphysial centres and the masses of cartil- age in which they are formed are sharply marked off from the body of the bone (cf. Figs. 12-13). This is largely because the formation of the epiphysial centres tends to lag behind that of the main centres, and thus the cartilage extremities of the bones are evident long after the forma- tion of the shaft is under way. In the epiphysial centres the bone formation is endochondral. The bone masses which they form are distin- guished as epiphyses. During the period of growth they are connected with the body of the bone by plates of epiphysial cartilage, into which the surrounding perichondrium extends as an ossification ridge. In this region bone formation takes place, with the result that the whole structure is greatly increased in length. After the period of growth, the duration of which differs in different bones, the epiphyses become firmly coossified with the body of the bone, although the lines of junction or epiphysial lines may still be visible. Thus in the distal extremities of the radius and ulna, in the proximal extremities of the fibula, or in the bodies of the lumbar vertebrae, the epiphysial lines appear even in old animals. In the foregoing figure (9) of the divided femur it will be seen that the position of the epiphysial lines is indicated by bands of compact tissue. Finally, in thoroughly macerated bones of young animals, the epiphyses are usually found to be readily separable from the bones. ‘bo. _ Ina comparison of the adult skeleton RYGUHiT AATRE Cecio teeetion of the with the more primitive embryonic _ skull in a three-day-old rabbit: b.o., skeleton, several differences in the arrange- cee te ee oe ment of the elements are evident. | Thus oe aes. eee ee many bones, notwithstanding their pos- — supraoccipital. session of several centres of ossification, are to be looked upon as individual either in the cartilage or in the bone condition. In other cases, as in the basal portion of the skull, separate bone elements are produced in a mass of cartilage primarily continuous. These either remain distinct throughout life, or, as in the occipital region, (Fig. 11), become fused together to form compound or composite bones. In still other cases, as in the vertebrae, the apparently single elements of the adult condition are the products not only of originally distinct bones, but also of primarily separate cartilage masses. The bones of the skeleton are united or articulated with one another by connective tissue in the form of ligaments, by cartilage, or in some cases by both together, 7.e., by fibrocartilage. Ligamentous union, distinguished as syndesmosis, is the most general type of articulation. $.0. 2 en. 7 _eX.0.. 3 18 ANATOMY OF THE RABBIT. Cartilage union or synchondrosis occurs in certain situations, as in the basal region of the skull. Union by fibrocartilage or symphysis is characteristic of the articulation of the two sides of the pelvis (symphysis pubis). The articulations of bones are of two types—immovable articulations or synarthroses, and movable articulations, diarthroses, or joints. In the former, motion is either absent or at least greatly restricted. In the latter, it is definitely provided for through the presence of joint-structures. Thus in a joint (Fig.12) the apposed surfaces of the bones are accurately modelled in relation to one another, and are moreover covered by a layer of cartilage, the latter forming a joint cushion. Between the two surfaces is a space, the cavity of the joint, containing a viscid material, the synovia, which serves for lubrication. The space is enclosed by a connective tissue capsule. The strength of the joint depends largely on the enclosing capsule, but it is usually greatly increased by the presence of accessory ligaments. In the more complex joints, such as that of the knee Fic. 12. Section of the elbow-joint of a four-day-old rabbit c., capsule; e.b., endochondral bone in the distal epiphysis of the humerus; e.a., extensor muscles of the forearm; e.m., extensors of the hand; f.a., flexors of the forearm; f.m., flexors of the hand; h., humerus; ol., olecranon; r., radius; s.c., Synovial cavity; u., ulna. Fic. 18. Section of the knee-joint of a four-day-old rabbit: c., capsule; f., distal epiphysis of femur; 1.p., patellar ligament (insertion of quadriceps femoris); p-.v., (Fig. 13); interarticular cartilages (men- popliteal vessels and nerves; t., proximal isci) are enclosed between the bone {Rpbysis of tibia: +.,ligament of the surfaces, and the latter are connected ligament; xx , anterior cruciate ligament. directly by short ligamentous cords. The various ligaments of a joint permit free motionof the bones, but only up to a certain point. Several differences are observable in joints according to the form of the apposed surface and the kind of motion provided for. Thus in the ball-and-socket joint or enarthrosis, exemplified by those of the shoulder and hip, a bone is able to move in various directions about its base of attachment, although actually, in the limbs, this motion is almost restricted to an anteroposterior direction. In the ginglymus or hinge- joint, as exemplified by the distal articulations of the limb, motion is MuscuLar Tissuks AND MUSCLEs. 19 restricted to a single plane. The gliding joint or arthrodia is one in which a slight degree of motion is made possible by one surface slipping over the other; it is exemplified in the accessory articulations of the vertebral arches. 3. MUSCULAR TISSUES. Muscular tissues are the active portions of the individual muscles of the skeleton and of the muscle coats of visceral organs. Their chief feature consists in the elongation of the cells to form fibres. These fibres may be considered to possess the contractile properties of proto- plasm, but with the contraction limited to one direction. Except ina few cases the fibres are arranged in a parallel fashion, so that the line of contraction of the muscle or muscle layer is the same as that of each of its fibres. The result of contraction in both is the shortening of the longitudinal axis and the increase of the transverse axis. Two chief types of muscle fibres occur in the body—the smooth or unstriated fibres, which are characteristic of the involuntary muscles or muscle ccats of the visceral organs, or of the skin, and the striated fibres which compcse the individual or voluntary muscles of the skeleton Smooth fibres (Fig. 14 B) are elongated, spindle-like cells, the substance of which is longitudinally striated, but possesses no _ transverse markings. The single nucleus of the cell occupies a central position, The muscles which they form are dis- tinguished as involuntary because their operation is not under the con- trol. of the will, their connections being with the sympathetic nervous system. The striated fibres (Fig. 14 A) are very much larger, cylindrical structures, the substance of which possesses characteristic transverse striations. Each fibre is enclosed by a loosely attached membrane, the sarcolemma, on the inner surface of which many nuclei occur.* The presence of these nuclei indicates ia that the fibre is not a single cell but Fic. 14. A, Striated (skeletal) muscle of the a syncytium, 72., an association of * =e as amraae ypc fom te cells unseparated by cell boundaries. The muscles formed by such fibres are under the control of the will, their connections being directly with the central nervous system. They comprise not only the typical muscles of the skeleton, but also the special muscles connecting the skeleton with the skin. The muscular substance of the heart differs both from striated and smooth muscle in being composed of branched anastomosing fibres, *The position of the nuclei is characteristic of the so-called white muscles. In the semitendinosus of the rabbit, which is a red muscle, the nuclei occur between the fibril bundles of the interior of the fibre. 20 ANATOMY OF THE RABBIT. which apparently form a continuous network. Like striated muscle, it possesses characteristic transverse markings, but, like involuntary muscle, it is under the control of the sympathetic nervous system. As gross structures the voluntary muscles are functional units, each of which has a particular action according to the movement per- mitted by the parts of the skeleton to which it is attached. They present a longitudinal striation which is roughly referred to as the direc- tion of the fibres, and which is of great value in identification. The striation is due to the circumstance that the fibres are arranged in parallel groups or muscle bundles, each of which is surrounded and separated from the adjacent bundles by a connective tissue enclosure, the perimysium. A muscle is typically spindle-shaped, consisting of a middle fleshy portion, termed the belly of the muscle, and of tapering ends which provide for attachment. The attachment is effected by a strong band of fibrous connective tissue, the muscle tendon (Fig. 6). In contraction it is usually seen that one end, thus designated as the origin of the muscle, remains stationary, while the! other, known as the insertion, assumes a_ position nearer the origin, carrying with it either the structure which is to be moved, or another portion determined by the point of leverage or the position of the joint. The action of the muscle is estimated in a direct line between origin and insertion, although the effect of the contraction, on account of the position of the joint, may be otherwise. From these more typical conditions, however, many modifica- tions occur. Se Sage SAS! Ria Some muscles, such as those of the (antrum pylacuaie: heletemaen of dees aodominal’wall, are disposed’ im» the rabbit: m.m., muscularis mucosae; t.m.c. form of flattened sheets, the ends of circular layer of the muscular tunic; t.m.1., z z longitudinal layer. of the muscular tunic; which are attached by broad, thin pe, me ee ee ee Sheets offeounective tissue, the apons euroses. In unipennate muscles the fibres are attached obliquely to the side of the tendon, or in bipennate muscles to both sides, like the vane of a feather. In the so-called biceps, triceps and quadriceps muscles of the limbs, the origin is divided into two, three or four portions. Furthermore, the recognition of origin and insertion depends on usual but not invariable relations. The exact effect of muscle contraction also depends as a rule on the synchronous action of other muscles. A muscle like the diaphragm does not possess an insertion after the fashion of ordinary muscles; and in some cases, asin the intrinsic muscle of the tongue or the so-called orbicular or sphincter muscles, both origin and insertion may be absent. Involuntary muscle is distinguished by its white or greyish coloration and by its smooth or homogeneous appearance. It forms characteristic Og ig et NeERvous TISSUES. yay layers in connection with visceral organs or with the skin, and is thus much less individual than the voluntary muscles in its relations to particular parts. It forms the nruscular portion (muscularis mucosae) of the mucous tunic of the alimentary canal, and also a separate muscular tunic lying in the outer portion of its wall (Fig. 15). In the muscular tunic the fibres are arranged in both circular and longitudinal directions. Involuntary muscle also forms a small constituent of many organs, such as certain glands, in which contractility is not a chief function. It forms a large constituent of the wall of the urinogenital tubes, particularly the bladder and the uterus. In association with elastic connective tissue it is an important constituent of the walls of the bloodvessels. Although there is an underlying community of structure in the walls of the bloodvessels, the two chief types of vessels, arteries and veins, present conspicuous differences, both in functional behaviour, and in their appearance in the dead animal. The differences are largely the result of differences in the relative amounts of the above-mentioned constituents. The arteries are thick-walled, elastic tubes, which, under the force of blood from the heart, first become greatly expanded, and then gradually con- tract, so that the blood is forced into the smaller capillary vessels. The veins on the other hand are thinner- walled, less elastic vessels, through which the blood is forced largely through the pressure from behind. In the dead animal the arteries appear white, flat or collapsed, and . empty. The veinson the other hand /sa ni amureaoemee io) a” appear large and dark on account of dendrites; | c.g., chromatophile granules ; their distension with blood. a ee 4. NERvouS TISSUES. Nervous tissues form the basis of the central nervous system and of the outlying nerves and ganglia. They comprise two kinds of elements —nerve cells and nerve fibres. In the central nervous system these elements are imbedded in a mass of neutral tissue, the neuroglia. Nerve cells are characteristic of the central nervous system and of the spinal and sympathetic ganglia. They differ greatly in form, but typically each consists of a cell-body (Fig. 16) bearing two kinds of processes—a fibre-process, the neuraxis or neurite, and a series of branched protoplasmic processes, the dendrites. The cell-body is distinguished by the presence in its interior of granular masses, the chromatophile or tigroid bodies. The latter extend into the dendrites, but not into the neuraxis. The dendrites may be greatly elaborated, and may be present to a considerable number. The neuraxis isa nerve fibre process. Since it continues as the central portion or axial cord of a nerve fibre, it may traverse a relatively enormous distance on its way to a peripheral organ. ANATOMY OF THE RABBIT. A nerve fibre consists of a central core, the axial cord, enclosed, except in the case of those of the olfactory nerve, by certain membranes. Two kinds of fibres are distinguished—medullated fibres, and non-medullated fibres. The former are characteristic of the peripheral nerves. In these (Fig. 17) the axial cord is surrounded by a comparatively thick membrane of fatty material, the medullary or myelin sheath. The latter is continuous except at certain points, the nodes of Ranvier, where the axial cord appears free except for an external investment of the whole fibre, the neurilemma. In the non-medullated nerve fibres the myelin sheath is lacking. This type of fibre is chiefly distributed in connection with the sympathetic system. A nerve is an association of nerve fibres, the latter being disposed in a parallel fashion and united together into bundles of larger or smaller size by connective tissue, which also forms a general peripheral investment, the epineurium. The dead-white coloration of a nerve is due to the Fic. 17. Portions of two pore id neve fibres fatty materials ol the myelin sheaths; but wemves from the sciatic of the are commonly found imbedded in a fatty con- rabbit: a.c., axial cord; a m.S., myelin sheath, nective stained black with osmic tissue which is associated with the a J wediemma: Cpimeirium and 1$/also of white coloranon: n.r., node of Ranvier. Nerve fibres, and also nerves, are distinguished functionally as afferent and efferent. They are organs of conduction, which carry impulses either from the peripheral parts of the body to the central nervous system, or in the opposite direction. Sensory nerves are afferent, while motor nerves are efferent. Nerves, however, usually contain both afferent and efferent fibres and are hence described as mixed. In the distribution of afferent and efferent fibres there is a marked difference between the external or somatic por- tions of the body and the internal or visceral portions. Consequently, both somatic and visceral kinds of afferent and efferent fibres are con- veniently distinguished. On account of certain diff- erences in coloration, the cellular and white fibrous constituents of the central 5.m.p f.m.a Fic. 18. Section of the spinal cord of the rabbit: c., central canal; f.m.a., anterior median fissure; s.m.p., posterior median sulcus; f.a., f.1., and f.p., anterior, lateral, and posterior funiculi of the cord; c.g.a. and c.g.p., anterior and posterior grey columns (horns of grey matter); r.a., and r.p., anterior and posterior nerve roots; s.a., white substance. nervous system produce characteristic patterns where either one is . cheer 3 CR TEND Sin BR lA Eye ee ee ie TERMINOLOGY. 23 concentrated. Thus the cellular material is greyish, and is hence distinguished as the grey substance, while white fibrous material pro- duces when concentrated an opaque white appearance similar to that seen in the peripheral nerves, and is hence described as white substance. In the spinal cord (Fig. 18) the grey substance is disposed as a central core, the white substance asa peripheral investment. A similar relation is found in the basal portion of the brain, but the characteristic pattern in the cerebral hemispheres and in the cerebellum is one in which the grey substance forms a peripheral, investing, or cortical layer. TERMINOLOGY. In special or descriptive anatomy it is necessary to employ an extensive system of terminology in order that the various structures of the body may be individually designated, classified, and referred to their respective positions. The terms used for this purpose may be classified into four groups, as follows: (1) general terms—those included in the names of parts, but applicable in themselves to similar structures (arteries, merves, etc.) in various parts of the body; (2). specific terms or names of parts; (3) regional terms—those defining areas (topographic); and (4) terms of orientation. Although few in number, the terms of orientation may be rezarded as the most generally useful terms of descriptive anatomy. This is because they are based on very general relations of the body, and are therefore of wide application. For this reason they are here selected for definition to the exclusion of others of a more restricting or individualizing kind. In all vertebrates we may recognize a longitudinal axis, corresponding, in general, to the line formed by the vertebral column. In the usual or prone position of the body this axis is horizontal. The uppermost surface is then described as dorsal, the lowermost surface as ventral, the sides of the body as lateral. Any position forwards, with respect to the long axis, is anterior in comparison with any position backwards, which is posterior. In relation to the long axis it is convenient to recognize a median vertical plane of section, which is one dividing the body into right and left halves; also transversal planes, which are planes situated at right angles to the median plane and to the long axis, and sagittal planes. The latter are vertical planes parallel to, and also including (as mid- sagittal), the median vertical plane. The median vertical plane is the plane of bilateral symmetry, each half of the body as thus defined being in a general way the reverse counterpart of the other. Structures situated in part in the median plane are unpaired, and are described as median, while structures situated wholly outside of the plane are paired, right and left, or dextral and sinistral. In relation to the median plane and to the sides of the body, structures are described as medial when nearer the former, and as lateral when nearer the sides of the body. The term intermediate is applied 24 ANATOMY OF THE RABBIT. especially to a position between medial and lateral, but this restriction is perhaps not justifiable. In considering the extent of bilateral symmetry, it is necessary to bear in mind that, although a fundamental feature in vertebrates, it is not perfectly retained in the adult condition. Symmetry is destroyed by the migration of an unpaired structure from a median to a lateral position, as is seen, for example, in the case of the abdominal portion of the ali- mentary canal; or, again, by the reduction or disappearance of structures belonging to one side of the body, as, for example, in the case of the mammalian aortic arch. Referring to centre and circumference, either in the body as a whole, or in particular parts, the terms deep and superficial, central and periph- eral, or internal and external may be applied. It may be observed, however, that the terms internal and external are sometimes used in the sense of medial and lateral, both in descriptive language and in the names of parts. In comparison with the terms medial (medialis) and median (medi- anus) the term middle (medius) may be used to designate the position of a structure lying between two others, the latter being otherwise desig- nated, for example, as anterior and posterior, or one in the median plane. The limbs being more or less independent structures, it is proper to apply to them certain terms otherwise applicable to the main portion of the body. The chief terms not properly applied elsewhere are proximal, meaning nearer the centre or base of attachment, and distal, toward the extremity. In the middle segment of the fore limb the respective positions of the bones (radius and ulna) are indicated as radial and ulnar. The terms tibial and fibular are also applicable, although with less reason, to the corresponding segment of the hind limb. The dorsal and ventral surfaces of the fore foot are described respectively as dorsal and volar, those of the hind foot as dorsal and plantar. In determining the identity of structures in a quadrupedal mammal, considerable difficulty may at first arise from the fact that descriptive terms, such as those just defined, are frequently included in the names of parts, the latter being, at the same time, terms applied in the first instance to the human body, in which the recognized relations are some- what different. In comparison with that of a quadrupedal vertebrate the human body occupies a vertical or erect position, and is to be con- sidered as having been rotated upward through ninety degrees on the posterior limbs. The latter accordingly occupy for the most part their original position, and the human arm indeed largely reassumes this position when allowed to hang freely at the side of the body. As in all cases, the face retains its forward direction. Thus the terms anterior and posterior as used in human anatomy mean dorsal and ventral, provided they refer to parts of the body, such as the entire trunk region, which have been affected by rotation. The terms superior and inferior as applied to man are similarly anterior and posterior as applied to a lower form. Since it is unwise to change the form of the official terms of human anatomy, it becomes necessary to interpret all such terms when used for a quadrupedal mammal according to the relations exhibited by GENERAL PLAN OF ORGANIZATION. 25 man. On the other hand, in ordinary description little advantage is to be gained from adherence to this principle. The terms anterior and posterior apply with greater force to a lower vertebrate than to man, while the terms superior and inferior are only of interest in the latter. In this case the rule here followed is to use the terms anterior and pos- terior for descriptive purposes without reference to the human relation. The same applies to the terms of direction—upward, downward, forward, and backward. It may be pointed out, however, that it has become the practice with some to employ in place of anterior and posterior, such terms as cephalic and caudal, thereby eliminating one of the difficulties; or similarly to use the termination—ad, in connection with words sig- nifying position, for the purpose of indicating direction (e.g., dorsad, cephalad, laterad, for dorsalward, etc.). Reference may also be made here to the fact that the human structures to which identifying names are applied frequently fail in one way or another to correspond to structures in a lower form. Composite structures to which individualizing names are applied, for example, may be represented by independent parts. Also, structures which are similar in form or function may be convergent. Finally, although it is essen- tial to endeavor to apply all terms as accurately as possible, it will be remembered that a terminology primarily arranged for one type cannot be exactly applied to another without considerable qualification. THE GENERAL FEATURES AND GROUND PLAN OF THE ORGAN SYSTEMS. In order to arrive at a proper understanding of the special features of organs, or of the relations to one another of organ groups, it will be found useful in many cases to consider them in the light of their deriva- tion from a general or ground form. A brief statement of what may be accomplished in this way and of the methods involved has already been given above (p. 4) in discussing the general interpretation of structure and the zoological position of the type, so that in the following pages only the actual facts of organization will be considered. In this connection it will be ponemiperedl that the subject is a very broad one, and in the present case it will not be possible to do more than select in the various organ systems the more important features, the con- sideration of which will be of direct assistance in the practical study of the type. Before taking up the individual organ systems, however, it will be advisable to refer in the first place to the plan of classification, and secondly to explain the more fundamental features of vertebrate organization as something underlying the disposition of the systems themselves. It will also be convenient to summarize the chief features of the embryonic organ systems, since it is by reference to embryonic conditions that the general features of the body are most readily recog- nized. 26 ANATOMY OF THE RABBIT. '" Classification of the Organ Systems.—The term organ system is employed in descriptive anatomy to designate a group of organs which cooperate in a general function. In many respects the systems repre- sent primitive functions, and it is therefore largely on account of the independent elaboration of these that the systems may be recognized also on a structural basis as groups of organs allied in origin and de- velopment. The exact number of systems recognized depends on certain arbitrary distinctions, the following being those usually distinguished: (1) The integumentary system, comprising the skin, and its ap- pendages, namely, the hairs and the general cutaneous, mammary, and inguinal glands. (2) The skeletal system, comprising the cartilage and bone elements of the skeleton, with their connections. (3) The muscular system, comprising all contractile structures of the body. Since, however, the involuntary muscles are arranged as muscle lavers in connection with visceral organs, the muscular system is usually considered as including only the individual or voluntary muscles of the skeleton and skin. (4) The nervous system, comprising the central nervous system (the brain and spinal cord) and the peripheral nervous system, the latter consisting of the paired cranial and spinal nerves with their associated ganglia. A special portion of the peripheral nervous system is set apart as the sympathetic nervous system. The latter consists of a pair of ganglionated sympathetic trunks lying along the ventral surface of the vertebral column, and of two series of ganglia, prevertebral and peripheral, connecting the trunks with the visceral organs. (5) The digestive system, comprising the digestive tube and its out- standing glandular appendages—the oral glands, the liver, and the pancreas. (6) The respiratory system, comprising the lungs, and respiratory passages, namely, the bronchi, the trachea and the larynx. With this system may also be included the accessory respiratory passages formed by the nasal fossae. (7) The vascular system, coniprising the organs of circulation, namely, the heart, arteries, capillary vessels,and veins. The lymph- conducting canals are also portions of the circulatory system, but since they are largely independent of the bloodvessels, they are usually con- sidered as forming with their associated lymph glands a_ separate lymphatic system. (8) The urinogenital system, comprising the reproductive and excretory organs, together with their common ducts—the urethra of the male and the vestibulum of the female—and the associated bulbourethral gland. The reproductive organs comprise, in the male, the central organs or testes, and the deferent ducts, both of which are paired, the unpaired seminal vesicle, and the paired prostatic and paraprostatic glands. In the female, the reproductive organs comprise the paired ovaries, uterine tubes and uteri, together with the unpaired vagina. The excretory GENERAL PIAN OF ORGANIZATION. 27, organs of both sexes comprise the paired kidneys and ureters and the unpaired urinary bladder. Certain organs of the body are not included in this classification: (1) The thymus and thyreoid glands are connected with the digestive tube in the embryonic condition, and for this reason are sometimes included with the digestive system, although in the adult they occur as in- dependent structures. (2) The suprarenal body is situated close to the kidney of either side, but is independent of the latter, both in the adult condition and in point of development. (3) The special (olfactory, optic and auditory) sense-organs of the head are highly elaborated structures, the relations of which are partly with the central nervous system. General Organization.—In the rabbit, as in all vertebrates, the general plan of organization involves three chief features, as follows: (1) axial orientation—the arrangement of the chief organ-systems longitudinally about a more or less central, axial support; (2) meta- merism—the arrangement of a con- siderable portion of the body on a segmented or metameric plan, according to which structures are repeated in a serial fashion along the axis; (3) branchiomerism — the arrangement of a small anterior portion of the body on a serial but branchial plan, the latter depending not on the succession of true metameres but of visceral arches. The extent to which the general disposition of the organ-systems is dependent on a fundamental plan will be evident from a comparison of the accompanying figure (20) of a trans- Fic. 19. Rabbit-embryo of 10} days (48 verse section of the rabbit-embryo, or mm.):m., mandibular; h., hyoid; 1 and 2, first of the schematic section of a general- and second branchial arches; a.l., anterior . = : > Hb bad: Meee memes’, Reepaeee ized vertebrates (Big. 21), with the Keibels Normentafeln, V.: Fig. 12.) , actual sections of the rabbit-foetus as given in the plates, more especially the abdominal section of Plate VIII “and the thoracic section of Plate Vil. It will be considered more fully below in connection with the general features of the organs. Metamerism (Fig. 19, me.) is characteristic of a dorsolateral portion of the body, identifiable in the embryo as that including the dorsal and intermediate portions of the middle layer or mesoderm Ghigs 20, dis and n.). In the adult it determines a number of features of serial arrangement, including the succession of the vertebrae and ribs, the divisions of the related dorsal musculature and its extensions to the ribs and abdominal wall, and indirectly the succession of the spinal nerves and their branches, of the parietal branches of the aorta, and the parietal roots of the inferior caval vein, as well as of the tributaries of the azygos vein. 28 ANATOMY OF THE RABBIT. Branchiomerism is an adult feature of lower aquatic vertebrates, such as fishes, where it appears as a succession of true. gill (or branchial) structures, which support gill filaments as functional respiratory organs. In higher terrestrial animals it appears as an embryonic feature (Fig. 19, m.h..1, 2) and is to be considered both as a determinant of adult form and asa mark of aquatic ancestry. As in lower vertebrates, it underlies not only structures of branchial significance (branchial arches in the restricted sense) (Fig. 19, 1, 2), but also modified branchial structures, including the first or mandibular arch (m.), and the second or hyoid arch (n.). So great is the modification of these structures in passing from the embryonic ? s ae : 7 ; EAS) Gee BA NG TDS ie Fic. 20. Transverse section of a rabbit-embryo of about 103 days, showing the arrangement of the organ-systems: ao., aorta; ch., not- ochord; coe., coelomic cavity; d.m., dorsal mesoderm (myotomic and scierotomic divisions); e., primitive alimentary canal (enteron); ect., ectoderm; 1.b., limb-bud; ms., mesenchyme; my., external portion of a myotome; n., nephrotome of embryonic kidney; inter- mediate mass of mesoderm; sp. and so., splanchnic (visceral) and somatic (parietal) portions of the ventral mesoderm. to the adult condition that the recognition of the ground-plan is perhaps here of less general importance. It, however, determines the position and relations of certain skeletal structures, including the auditory ossicles, the hyoid, and in part the laryngeal cartilages—a point of some value in the classification of the parts of the head skeleton. It also determines the succession of certain soft structures, including the fifth, seventh, ninth and tenth cranial nerves; also the chief arterial vessels of the heart, which are more fully referred to below. GENERAL PLAN OF ORGANIZATION. 29 Embryonic Plan of the Organ Systems.—In the individual organ- systems the main features of the general plan, as estimated on embryonic development, may be outlined as follows: 1. The formation of an axial skeletal support, consisting primarily of a strand of cellular tissue, the notochord, and secondarily of a seg- mented cartilaginous, afterwards bony, vertebra! column. 2. The formation of (a) a primary cartilage skull (chondrocranium) as a support for the brain and capsules of the special sense organs (neurocranium or cerebral cranium); and (b) a series of cartilaginous visceral arches (splanchnocranium or visceral cranium). 3. The formation of the chief skeletal muscle in a dorsolateral position along the axis. 4. The formation of the central nervous system as a tube of nervous matter (neural tube), lying on the dorsal side of the axial support, and differentiated into a general posterior portion, the spinal cord, and an anterior expanded portion, the brain. 5. The formation of the digestive tube as a median structure, lying directly beneath the axial support, and of special glandular appendages arising from the epithelium of its wall. 6. The formation of the lungs as paired outgrowths of the ventral wall of the digestive tube, afterwards connected with the outside of the body by accessory respiratory tracts traversing the head. 7. The formation of the circulatory system primarily on an aquatic plan. This involves the formation of (a) the heart in a ventral position to the digestive tube and immediately behind the gills; (b) a ventral aorta, passing forward to the gills, and dividing into a paired series of branchial aortic arches; (c) a dorsal aorta, combining the aortic arches, and passing backward along the ventral surface of the axial support; and (d) a series of paired veins returning the blood from various parts of the body to the heart. 9. The formation of the reproductive organs or gonads in association with the dorsal lining of the coelomic cavity, and their connection with the outside of the body by modified kidney ducts. 10. The formation of the kidneys, either as embryonic or permanent structures, from an intermediate mass of tissue, lying in general between the dorsal musculature and the lining of the coelomic cavity (ef. position of embryonic kidney in Fig. 20, n.). 11. The formation, in the ventral portion of the body, of an extensive space, the coelomic cavity or coelom, afterwards differentiated into pericardial, pleural and peritoneal portions. 30 ANATOMY OF THE RABBIT. THE SKELEDAL SVs DEM That portion of the skeleton which in the adult is designated as axial includes the vertebral column, together with the associated ribs and sternum, and the skeleton of the head. The axial relation belongs in the first instance to the bodies of the vertebrae and to a small portion of the base of the skull. A line drawn through the centres of the vertebral bodies indicates the position of the primary axial support, the notochord. Siiltve Fic. 21. Schematic representation of the chief organ-systems of a generalized vertebrate as seen in a transverse section of the abdominal region: Integument—int. Skeleton—v., vertebral body; a.v., vertebral arch; c.v., vertebral canal. Muscular system—s.m., skeletal muscle; v.m., visceral muscle. ; Nervous system—m.s., spinal cord, with the central canal, and the dorsal (pos- terior) and ventral (anterior) roots of the spinal nerves; g.r.p., ganglion of the posterior root; r.c., ramus communicans to sympathetic trunk; r.m.a. and r.m.p.. anterior and posterior rami of a spinal nerve; t.s., sympathetic trunk. Digestive system—i., intestine. Vascular system—ao., aorta. Urinogenital system—k., kidney; go., gonad (ovary or testis). Serous cavity—c.p., general coelom, pleuroperitoneal, or peritoneal cavity; p.v. and p.p., visceral and parietal parts of the serous tunic—visceral and parietal peritoneum; mes., mesentery. The arches of the vertebrae are identified with the general functions of support of the related musculature and enclosure of the spinal cord. In lower vertebrates the line formed by the vertebral column and the base of the skull is for the most part straight. In a mammal the vertebral *The majority of the features mentioned below are illustrated in the plate figures of regional sections of the rabbit-foetus which tollow, and for this reason the references are not indicated except in special cases. ca THE SKELETAL SYSTEM. 31 column presents marked dorsolumbar and cervical flexures. The axis of the skull proper, the basicranial axis, identifiable as a line passing through the centres of the basioccipital, basisphenoid and presphenoid bones, may differ by a considerable angle from that of the related cervical vertebrae. Further, the basicranial line, if continued forward in the rabbit, would pass through the dorsal wall of the skull immediately in front of the orbits (See Plate IJ, or Fig. 33 of the divided skull). It accordingly differs by a considerable angle from the basal line of the face, or basifacial axis. The skull consists primarily in the embryo of a cartilage trough, the extent of which is roughly definable as the area occupied by the occipital, anterior and posterior sphenoidal, and ethmoidal portions (cf. Plates III-V). Asa cartilage skull it is designated as the chondro- cranium, and after its conversion into bone as the osteocranium. It is no more than an enclosure for the brain, except that it has associated with it the cartilage capsules of the nasal, visual, and auditory organs, and, in the case of the first and last of these, the capsules are incorporated with the skull proper. Thus, the primary skull is designated as the neuro- cranium or cerebral cranium, to distinguish it from a second portion of the head skeleton, the splanchnocranium or visceral cranium, which includes the series of visceral arches suspended from the ventral surface of the neurocranium. The addition to the primary head skeleton of a large number of membrane bones results in more or less confusion of the original divisions, since the membrane portions of the visceral cranium are, with the exception of the mandible, united by suture with those of the cerebral cranium, while the true cartilage or cartilage bone portions of the former, occurring as the auditory ossicles, the hyoid and larynx (in part) (Plate II), although highly modified, remain in a more or less independent relation. The skull, or cranium—using that term in a general sense—may be described as consisting of a cranial portion (the cranium proper) and of a facial portion, the latter including as visceral structures the upper jaw and the mandible, and as cerebral structures the parts of the turbinated bones and the associated secondary respiratory tract formed by the nasal fossae. In a mammal the investing membrane bones of the face are largely associated with a great extension of the nasal capsules, or that portion of the primary skull from which the turbinated cartilages are formed. Thus, as illustrated in the transverse section of Plate III, the nasoturbinals and maxilloturbinals are formed on lateral extensions of the primary septum (mesethmoid cartilage), but are supported more directly, and in the adult wholly, by the investing membrane bones (nasals, maxillae and premaxillae). The elements of the head skeleton may be classified as follows: 1. The CEREBRAL CRANIUM (cranium cerebrale or neurocranium), including: (a) The primary cartilage skull (chondrocranium), enclosing the brain, and containing in its wall the olfactory and auditory capsules (embryonic) ; 62 ANATOMY OF THE RABBIT. (b) The secondary bone skull (osteocranium), replacing (a) and comprising the occipital, sphenoid, ethmoid, inferior turbinal, and periotic bones; (c) The associated derm elements, comprising the interparietal, parietal, frontal, nasal, vomerine, lacrimal, tympanic,* and squamosal bones. 2. The VISCERAL CRANIUM (cranium viscerale or splanchno- cranium), including; (a) The primary mandibular and hyoidf visceral arches (embry- onic) ; (b) The secondary elements, replacing (a)—the malleus, incus, and stapes of the auditory chain; the hyoid bone and its connections with the skull; (c) The associated derm elements of the face and palate, com- prising the premaxillary, maxillary, zygomatic, mandibular, palatine and vestigial pterygoid bones. The anterior and posterior portions of the appendicular skeleton include each a proximal or girdle portion and a distal portion, the free extremity. In each the free extremity is divisible into proximal, middle and distal segments. The proximal joint of the limb, either of the shoulder or hip, is an enarthrosis, although it will be seen by examination of the rabbit that the glenoid cavity of the shoulder is not greatly elaborated as a concave surface. The distal joints of the limbs conform to the type of the ginglymus, and with minor exceptions the movements are restricted to one, namely, the sagittal, plane. In each limb they are described as movements of flexion and extension, since the general effect of movement is to bend or straighten particular parts on one another. Similar motions are observable in the limb as a whole, but are not spoken of as of flexion and extension, since the shoulder and hip joints are of a totally different type. In the limb as a whole, the chief motions are of pre- and post-traction, but it is convenient to recognize also movements of ad- duction (toward the body) and of abduction (away from the body). In both cases the axis of the upper arm, or of the thigh, is assumed to be parallel to the median vertical plane. In mammals, as in other terrestrial vertebrates, the anterior and posterior limbs are equivalent part for part. They are said to be serially homologous or homoplastic. There is, however, particularly in the mammalia, a pronounced difference between the anterior and posterior limbs in the respective positions of the different segments. Consequently, in studying the related musculature, it is advisable to consider not only the functional relations of muscles, but also the equival- *The identification of the tympanic as a derm element has been questioned. +The thyreoid cartilage of the larynx and its connection with the hyoid (greater cornu) are modified branchial arches, but the structure as a whole is not included with the head skeleton. THE SKELETAL System. 33 ence of the bone surfaces. This may be estimated either from the rotations of the limbs in embryonic development, or by reference to an ideal or neutral condition as indicated in the accompanying diagram (Fig. 22). The neutral type (A) may be defined as one in which the proximal segment (humerus or femur) is situated at right angles to the median vertical plane, the middle segment directed downward parallel to the median plane, and the distal segment again at right angles to it. In this condition the bones of the middle segment are parallel, with the radius, or the tibia, according to the particular limb, anterior in position. The first digit is also anterior. The entire anterior surface is indicated in the accompanying diagram by shaded lines. The angles b. and ec. are “extension”’ angles; e. and f..“‘flexion’’ angles; a. is an “abduction”’ angle, while d. is an “adduction” angle. In both limbs of a mammal, the entire appendage is rotated downward to a position more or less underneath the body. In the anterior limb (B) the proximal segment is rotated backward, the middle and distal segments forward. The two divisions thus come to differ from one € Vala A 1 Fic. 22. Schematic representation of the respective positions of the segments in the mammalian limbs: A, neutral; B, anterior limb; C, posterior limb. Ex- planation in text: tr.p., transverse plane another to the extent of two right angles. Also, in the front limb, the in- teresting condition is observable that the radius and ulna are crossed on one another, the position of the former being chiefly anterior, although its proximal end is lateral, and its distal end, like the first digit, medial. In the rabbit, as in most mammals, the hand is thus fixed in a prone position, while in man the hand may be rotated to a supine position in which the radius is wholly lateral, or in which the two bones are parallel. In the posterior limb (C) all three segments are rotated forward. Con- sequently the bones of the middle segment retain their original parallel position with reference to one another. The extension angles of the knee and ankle are anterior, the flexion angles posterior. They differ from the corresponding angles of the anterior limb only at the middle joint, but here the difference amounts to two right angles. It will be observed also that the distal or ankle joint of the posterior limb retains a primitive condition, in which the foot is placed approximately at right angles to the leg; in other words, it is of a plantigrade type, one in which 4 2) 34 ANATOMY OF THE RABBIT. the sole of the foot rests on the ground. Thus the muscles described as flexors in the foot pass over the heel. They are functionally flexors of the toes, but extensors of the foot as a whole. Those described as extensors, lie for the most part on the anterior surface of the tibia, and are turned from their original course to the dorsal surface of the foot. They are functionally extensors of the toes, but flexors of the foot as a whole. THE NERVOUS SYSTEM. In the central nervous system, the more general features relate to the form of the whole structure as a neural tube, containing a central cavity, the neurocoele. This is differentiated into two portions, namely, a less modified portion, the spinal cord,or spinal medulla, containing as its cavity the central canal, and a greatly expanded portion, the brain, or encephalon, which is divided into a series of paired and unpaired seg- ments, and contains in its interior a corresponding series of divisions of the original cavity, known as its ventricles. Although the internal structure of the brain is a matter of the dis- position of its nerve-centres and fibre-tracts, it is customary to estimate its primary divisions by reference to a general plan, the latter being in most respects one that is common to vertebrates generally and thus appears in a less elaborated condition in the embryo, or in the adults of lower forms. The main features of this plan are indicated in the ac- : companying diagram (Fig. 23). The brain as first formed in the embryo appears as an anterior ex- panded portion of the neural tube, or rather three expansions arranged in a linear series. They are described as the primary cerebral vesicles; or, as primary divisions of the future brain, they are designated in anteroposterior order as the prosencephalon, mesencephalon, and rhombencephalon. The first of the primary divisions, the prosencephalon, or primary fore-brain, becomes divided during development into two portions, namely, an anterior portion, the end-brain or telencephalon, which is largely a paired structure, and a second portion, unpaired, the dience- phalon, or inter-brain. The larger paired portion of the telencephalon is the basis of the cerebral hemispheres. It contains, as divisions of the primary cavity, a pair of cavities, the lateral ventricles. The anterior portion of the telencephalon, moreover, becomes differentiated, so that a small terminal olfactory segment, the rhinencephalon, is more or less perfectly marked off from the rest. In the mammalian brain this part is chiefly identifiable as the paired olfactory bulb, the latter being the anterior portion of the olfactory lobe or olfactory brain, and containing in its interior an extension of the lateral ventricle. The unpaired portion of the prosencephalon is considered as belonging in part to the telencephalon and in part to the diencephalon. Its cavity, the third ventricle, is connected with the lateral ventricles through the interventricular foramen. Its anterior wall is formed by a transverse connection of the cerebral hemispheres, the lamina terminalis. In all Tur Nervous SYSTEM. 35 vertebrates this portion of the brain is remarkable for the manner in which its wall is differentiated. The ventral portion extends downward as a slender funnel-like structure, the infundibulum, the tip of the latter — Fic. 23. Plan of the divisions of the vertebrate brain: A, embryonic; B, adult, projection from dorsal surface; C, adult, sagittal section. Primary divisions—PR, prosencephalon; T, telencephalon; DI, diencephalon; MS, mesencephalon; RH, rhombencephalon; MT, metencephalon; MY, myelence- phalon; S, spinal cord. a.c., cerebral aqueduct; b.o., olfactory bulb; cb., cerebellum; c.m., mummillary body; c.o., optic chiasma; c.p., pineal body; c.q., corpora quadrigemina; ep., epitha- lamus; fi., interventricular foramen; h., hypophysis; h.c., cerebral hemisphere; hyp., hy pothal amus; inf., infundibulum; 1.t., lamina terminalis; p., pons; p.c., chorioid plexus of third ventricle: p.cr., cerebral peduncle; t:, thalamus, also in- dicates position of massa intermedia; v.l., lateral ventricle; v.m.p., posterior medullary velum; v.q., fourth ventricle. being in contact with the pituitary body or hypophysis and its base connected with a small grey elevation, the tuber cinereum. Its cavity is the recessus iafandipuli. Immediately in front of the infundibulum the 36 ANATOMY OF THE RABBIT. ventral portions of the optic tracts join to form the optic chiasma, and immediately behind it the floor is thickened, forming externally a pair of rounded protuberances, the mammillary bodies. In the brain of the rabbit this structure consists superficially of a larger median portion with faint lateral elevations appended to it. Collectively, these structures are considered to form a major division, the hypothalamus, the latter con- sisting of two portions, namely, an optic portion, comprising the in- fundibulum, tuber cinereum, and the optic chiasma, and a mammillary portion, including the mammillary bodies. The more dorsal portion of the diencephalon, containing the major part of the third ventricle, is known as the thalamencephalon. Its lateral walls are greatly thickened, while its roof is extremely thin, especially in its anterior-part. Here the actual roof of the ventricle is formed only of a thin layer of tissue, the epithelial chorioid lamina, but the latter has associated with it a series of vascular ingrowths of the investing pia mater, the latter being described in this relation as the chorioid web (tela chorioidea). The two structures together form a chorioid plexus. This extends downward into the third ventricle, reaching out also into the lateral ventricles. The dorsal portion of the thalamencephalon bears posteriorly the pineal body, the latter together with certain related structures, the habenulae and habenular commissure, forming the epithalamus. The general portion of the thalamencephalon bordering the third ventricle, and broadly connected across the latter by the massa intermedia, is the thalamus. In the brain of the rabbit it will be seen that the thalamus is chiefly indicated externally by a rounded protuberance, the pulvinar. The latter is dorsal in position and is imperfectly marked off from a second protuberance, the lateral geniculate body, lying on its postero- lateral side. To the medial side of this is a third protuberance, the medial geniculate body. The medial and lateral geniculate bodies as thus defined constitute the metathalamus (Fig. 53). The second of the primary divisions, the mesencephalon, or mid-brain, is noteworthy in a mammal as lacking a ventricle. Its cavity is a narrow canal, the cerebral aqueduct, leading from the third ventricle backward to the fourth ventricle, or cavity of the rhombencephalon. Externally, its roof is differentiated into four rounded elevations, the corpora quadrigemina, of which the anterior pair are much larger than the posterior ones. Its floor is chiefly formed by a pair of divergent cords, the cerebral peduncles. The parts of the mesencephalon and prosencephalon together con- stitute the large brain, or cerebrum. The third primary division, the rhombencephalon, or primary hind- brain, is a greatly elaborated portion from which arise the majority of the cranial nerves. The constricted area joining it with the mesencephalon is known as the isthmus rhombencephali. It includes the anterior medullary velum and brachia conjunctiva (Fig.55). The rhombencephalon itself is divisible into two portions, especially well defined in the mammalia, namely, the metencephalon, or hind-brain, and the myelence- phalon, or after-brain. The former includes the small brain, or cerebellum, 37 Tur Nervous SYSTEM. Cerebrum saunpad 1D4Ga43.) Durumasiiponh vaog 40) snupvjoyyrd 7 SNWUWDIDYIVII [NI SNUDIDY J, snuvjoyjod a Fy SUOISIAIp IOUT pue unyyod “wnywrys sngsod (uoyeydoououtys) aqoq A40jp9v {10 (e}eSuU0TqGO eI[Npour Io ‘UTBIq-I9}JV) uoleydaouea Ay (ure4q-puty{) uo[eydaousjay suog | wn)]agas4ay J | Uo[DYd 2IUIUWDIDY J, (uleiq-19}UT) ; uojeydaouerq snuvyeyyodAy yo uorizsod Aapypuuwmn fy snureyeyyodAy jo worsod 2440 ee (ule1q- pus ) uojeydaouajay | — asayd SULaY [DAQIAID (uojeydeouq) NIVUd AHL AO SNOISIAIG AHL AO NV Id (uleiq-pury Areutig) _ 7 (QPISOA [B1G9199 I0119}SOJ) uojeydeousquioyy “[I] (uleiq-piut AUT) (OJOISIA [BIGI190 VIPpI]) uojeydaousse_y “TT (ureiq-o10} ATvUTId) (Q[DISOA [BIGoIIO IOLII}UV ) uo[eydeouesolg ‘J 38 ANATOMY OF THE RABBIT’. and a ventral structure of a commissural nature, the pons. The my- elencephalon is a transitional portion connecting the brain with the spinal cord. The cavity of the rhombencephalon is the fourth ventricle. It is a peculiarly shaped space, the floor and lateral walls of which are very greatly thickened, while the roof is for the most part thin. The roof appears at first sight to be formed largely by the cerebellum, but is in reality formed by two membranes underlying the latter. One of these, the anterior medullary velum, is connected forwards with the mesence- phalon, while the other, the posterior medullary velum, covers a triangular space over which the posterior margin of the cerebellum does not extend. The posterior medullary velum has the same structure as the chorioid plexus of the third ventricle, but is much more poorly developed. Apart from its principal divisions, which, as indicated above, are more or less ccommon to all vertebrates, the brain depends for its external form on the elaboration of certain parts in comparison with others. In the mammalia the cerebral hemispheres and the cerebellum are the chief form- determinants, although the pons and the corpora quadrigemina may also be considered in this connection. It will be seen also that the form of the brain is more or less dependent on the existence at certain places of well-marked flexures (cf. Plate II). The first of these, the cephalic flexure, is in the region of the mesencephalon, the anterior portion of the brain being bent downward; the second, or pontine flexure, is at the fourth ventricle; while the third, or cervical flexure, is at the point where the myelencephalon passes over into the spinal cord. The peripheral nervous system embraces two groups of paired and, for the most part, metamerically arranged nerves, namely, the spinal nerves—those arising from the spinal cord and leaving the vertebral column through the intervertebral foramina; and the cranial or cerebral nerves—those arising from the brain and passing through the foramina of the skull. Of these the spinal nerves are less modified, both in structure and distribution. A spinal nerve (Fig. 21) is a product of two roots, one of which, the radix posterior, is sensory and arises from the dorsal side of the cord, while the other, the radix anterior, is motor and arises from its ventral side. The posterior root bears a root-ganglion. The combined nerve is distributed in three chief parts, of which two, the anterior and posterior rami, are distributed as mixed nerves to the skeletal muscle and the skin, —the anterior ramus being in practically all cases the prominent nerve—— while the third, the ramus communicans, is connected with the sym- pathetic trunk. The sympathetic trunk is formed on either side by a series of ganglia joined together by connecting cords. It is connected not only with the spinal nerves through the rami communicantes, but also with a series of prevertebral (coeliac, superior mesenteric, etc.) ganglia and plexuses from which nerves pass, chiefly in association with the bloodvessels, to the peripheral ganglia and plexuses of various organs. ‘The anterior and posterior rami of the spinal nerves contain both sensory and motor components, and are typical somatic nerves. The ramus communicans is a visceral connection containing similar com- THE NErvous SystEM. 39 ponents, here better described as afferent and efferent. It connects the central nervous system either with the actual viscera, or with visceral structures in a general sense, the chief relations of the sympathetic system being with the smooth muscle of visceral organs, or with that of the bloodvessels in various parts of the body. The cranial nerves are comparable in some respects to the spinal nerves, but in various ways are highly modified structures. The third, fourth and sixth, respectively oculomotor, trochlear and abducent nerves, which are distributed to the muscles of the eye, are considered as belong- ing to the somatic motor division. The second, or optic nerve, and the nervous portion, or retina of the eye, are specialized parts of the central nervous system, arising embryonically as an outgrowth of the latter. The nerves of the remaining special sense-organs, including the first, or olfactory nerve and the eighth, or acoustic nerve, have been con- sidered as parts of an extensive system of nerves, which, in lower aquatic vertebrates, also contains representatives of the branchiomeric series, and is distributed widely to sense-organs lying in the skin. Of the remaining cranial nerves the fifth, seventh, ninth and tenth are branchiomeric. Although the connections of these nerves are not fully considered in the dissection as here outlined, their chief characteristic as branchiomeric structures may be indicated. The fifth, or trigeminal nerve, the nerve of the mandibular arch, arises in two parts, one of which, the portio major, is sensory, the other, the portio minor, motor.. The portio minor unites with the third or mandibular division of the portio major. Thus, the terminal branches of all three divisions, ophthalmic, maxillary, mandibular, are distributed as somatic sensory nerves to the skin of the head, and, in addition, the mandibular nerve distributes visceral motor branches to certain muscles (masticatory group, mylohyoid and digastric) regarded as belonging to this, the first arch. A visceral sensory con- nection with the mouth is considered to be formed by the lingual branch of the mandibular nerve and by the palatine branches of the spheno- palatine ganglion. Both are, however, connected with the central nervous system through the seventh nerve, the former by the chorda tympani, and the latter by the great superficial petrosal. The seventh, or facial nerve is the nerve of the second, or hyoid arch. It is chiefly distributed as a visceral motor nerve to the cutaneous muscles of the head. The ninth, or glossopharyngeal nerve, belonging to the third arch, the tenth, or vagus, belonging to the fourth and succeeding arches in lower forms, and the eleventh, or spinal accessory nerves, the latter apparently related to the vagus as a motor portion, are distributed as visceral motor nerves to the phary ngeal and laryngeal musculature, and as visceral sensory nerves to various ; visceral organs, the ninth nerve supplying the gustatory organs of the tongue. The spinal accessory, moreover, has a characteristic distribution to the cleidomastoid, sterno- mastoid and trapezius muscles of the side of the neck and shoulder. The twelfth, or hypoglossal nerve has the relation of the ventral or motor portion of a spinal nerve, and is distributed as a motor nerve to the muscles of the tongue. 40 ANATOMY OF THE RABBIT. THE DIGESTIVE SYSTEM. The digestive system comprises as its chief portions the digestive tube and the digestive glands. The digestive tube is divisible into several parts, which, with the exception of the caecum and its vermiform process, are arranged in a linear series. The digestive glands comprise the oral glands, the liver, and the pancreas. They are parts of an extensive series of epithelial glands, otherwise contained within the wall of the tube, and for this reason not appearing as gross structures. The parts of the digestive tube may be classified as follows: I. Oral Cavaty: 5. Small Intestine. Oral cavity proper. Duodenum. Vestibulum oris. Mesenterial intestine. Jejunum. 2. Pharynx. eum. Nasal portion. Oral portion. 6. Large Intestine. Laryngeal portion. Caecum. Vermiform process. 3. Oesophagus. Colon. Rectum. 4. Stomach. In its most general features the digestive system is significant as an epithelial tube, in which the food during its passage is subjected to the action of digestive juices provided by the epithelial glands, and is modified, by solution or otherwise, so that it is capable of being absorbed through the epithelial surface. In the form of the digestive tube as seen in a vertebrate, however, a number of gross mechanical features are evident, such as, for example, the increase in capacity, or in absorptive area, through the folding of the mucous membrane, or the expansion of the wall; or again, the presence of a special muscular tunic, and its modifica- tion at certain places, as in the oesophagus, the pyloric limb of the stomach, and the first portion of the colon. Moreover, many features of the abdominal portion of the tube, and, indeed, certain of its recognized divisions, depend on its relation to an extensive serous sac—in a mammal the peritoneal cavity. In this connection it is to be considered that the digestive tube is primarily a median structure. It has this relation in the earlier stages of embryonic development (Fig. 20), and in many of the lower vertebrates it does not deviate to a great extent from a median position. In all higher vertebrates, however, the tube becomes greatly elongated in comparison with the cavity in which it lies, and thus becomes extensively displaced to one side or other of the median plane. This development, while advanced in all mammals, may be said to reach an extreme in the herbivorous mammalia; and in many cases it is further increased by the independent elaboration of the blind intestine or caecum. In the rabbit the combined length of the small and large intestines is approximately eleven times that of the body. THE DIcEsTIVE SYSTEM. 4] In considering the divisions of the digestive tube in the rabbit, the posterior, or post-cephalic portion, comprising the oesophagus and suc- ceeding parts, may be distinguished from the anterior, or cephalic portion, the latter comprising the oral cavity and pharynx. The former is a free portion embracing the digestive tube proper, while the latter is a fixed portion exhibiting a variety of general mammalian features connected with the organization of the head. The oesophagus is a slender but greatly expansible tube leading from the pharynx to the stomach. In its passage backward it traverses the neck and the thorax, and in both regions occupies a median position. In the thorax (Plate VII) it will be observed that it lies between the heart and the dorsal aorta, thus exhibiting the original relation of the digestive tube to the aortic portion of the vascular system. The succeeding portions of the digestive tube are those associated with the peritoneal cavity, and with the exception of the terminal portion, the rectum, are displaced from a median position. Consequently, the divisions which are recognized are based partly on the differential characters of the wall, and partly on the position of structures more especially in relation to the supporting peritoneum. Thus, the chief features of the stomach depend on the expansion of the organ and the rotation of its pyloric end forward and to the right. In the intestinal tract as a whole the chief although by no means most conspicuous feature of position depends on the looping of the entire structure on itself, so that the terminal portion, chiefly the transverse colon, crosses the ventral surface of the duodenum and then turns backward on the dorsal surface of the mesenterial small intestine. The duodenum is sharply marked off from the mesenterial intestine as an extensive loop, containing the major part of the pancreas and its duct, and lying on the right side of the dorsal wall of the abdomen. The mesenterial intestine is a greatly convoluted portion, lying chiefly on the left side of the abdominal cavity, and loosely supported by the broad, frill-like mesentery. From the pylorus to the sacculus rotundus there is no abrupt change in the character of the wall, although the first portion of the mesenterial intestine, that designated as the jejunum, and the duodenum may be considered together as a more vascular portion with thicker walls in comparison with the second portion, the ileum, in which the wall is less vascular and more transparent. The main portion of the large intestine, the colon, although greatly specialized, may be considered to consist as in manof ascending, trans- verse, and descending parts, that is to say, the ascending colon lies on the right side of the body and passes in a general way from its point of origin on the caecum forward to a point where it becomes flexed to the leit as the transverse colon; the latter crosses the body and is flexed backward as the descending colon. In the rabbit, however, that portion definable as the ascending colon is greatly elongated, and is composed of five principal limbs, united by flexures. Two of these, in dissection from the ventral surface, are concealed by the base of the superior mesenteric artery, since they lie on its right side. The descending colon is also only nom- inally related to the left side of the body wall, its supporting peritoneum, the descending mesocolon, being closely connected with the mesoduodenum 42 ANATOMY. OF THE RABBIT. of the ascending limb of the duodenal loop. The course of the caecum as it lies in the body is comparable to two turns of a left-hand spiral, its blind termination, the vermiform process, being dorsal in position and directed for the most part backward. It may be observed at this point that in their vascular supply the more typical divisions, namely, the transverse and descending colons, have arterial branches, respectively, the middle and left colic arteries, comparable to those of man; while on the other hand the right colic relation, on account of the great complexity of its parts, is represented by a large number of vessels, branches of a common ileo- caecocolic trunk. . The form of the anterior,or cephalic portion of the digestive tube (Plate II) depends on its fixed relation with respect to the enclosing parts of the head-skeleton. In the rabbit, as in mammals generally, the oral cavity is divisible into two portions, of which one is the oral cavity proper, while the other, the vestibulum oris, is a space enclosed between the alveolar process of the jaws and the teeth on the one hand and the cheeks and lips on the other. As in other vertebrates, the tongue is a muscular structure projecting upward and forward into the oral cavity from its base of attachment on the hyoid apparatus, but its greater elaboration, as well as the differentiation of special processes, the circumvallate and foliate papillae, for the accommodation of the gustatory organs, are features of mammalian significance. The roof of the oral cavity is formed by an extensive palatal surface, comprising the hard palate, and the membranous, or soft palate. These structures also form the floor of the accessory respiratory tracts of the nose, the posterior aperture being thus carried backward to a point more directly above the aper- ture ‘of the larynx. As accessory structures the teeth present two mammalian features: they are heterodont, or differentiated according to particular regions, and the adult series, excepting those designated as molars, are permanent teeth, replacing deciduous, or milk teeth of the young animal. The condition is thus described as diphyodont in comparison with that in lower vertebrates, where there is usually a multiple tooth change, new teeth being developed as required (polyphyodont type). Moreover, in the rabbit, as in all mammalia, the number is restricted, so that, considering the differentiation of the teeth, it is possible to express their relations by a dental formula. In the mammalia generally the teeth are differen- tiated into incisors, canines, premolars “and molars, oe in placental mammals the full dental formula is indicated as i. 3, c. 4, pm. 7, m. 3. In the rabbit as in other rodents, however, the a sition is greatly modified by the elaboration of two pairs of incisors and the corresponding obliteration of intermediate teeth, the place of the latter being occupied by an extensive gap, or diastema in mbites no teeth occur. The dental formula onthe rabbit isa) =e; 2) pm. m0 It will also be observed in this animal that the absence of the ae teeth allows the lips to be approximated behind the incisors, and since the lips are in this region also provided with hairs on their internal surfaces, the oral cavity is separated almost completely from a small space enclosing the incisor teeth. This adaptation, however, is not so perfectly developed in the rabbit as in certain others of the rodent order. * Ne THE RESPIRATORY AND VASCULAR SYSTEMS. 43 The chief features of the pharynx depend on its relation as a common or general portion of the digestive tube with the tubes of the respiratory system. It is divisible into an oral portion, representing the direct connection of the oral cavity with the oesophagus, a dorsal or nasal portion, connected with the nasal fossae, and a ventral or laryngeal portion, containing the aperture of the lary aD: < THE RESPIRATORY SYSTEM. In all air-breathing vertebrates the lungs arise embryonically as ventral outgrowths of the digestive tube, and are secondarily con- nected with the outside of the body through special perforations of the anterior portion of the head and through the oral cavity. As indicated above, this connection in a mammal is represented by an extensive nasal cavity bearing on its lateral walls the olfactory sense- organs. It is distinguished as an accessory respiratory tract from the true respiratory tract formed by the trachea and its terminal divisions, the bronchi. The respiratory system as represented by the lungs and related tubes, is nominally ventral to the oesophagus, but this relation is chiefly true of the trachea. Inthe thorax (Plate VII) the bronchi are, in general, interposed between the oesophagus and the heart, the lungs being expanded laterally into the paired pleural cavities. Tie VASCULAR SYoDEM In the rabbit, as in all vertebrates, the vascular system embraces a central, muscular organ of propulsion, the heart, and a series of branched tubes, the bloodvessels, the latter being of three different kinds: (a) thick- walled, elastic, distributing vessels—arteries; (b) microscopic terminal canals in the peripheral organs—capillaries ; and (c) thin-walled collecting vessels—-veins. The chief mammalian feature in this system consists in the division of the heart into two portions (Plate VII), each consisting of a receiving chamber, or atrium, and a driving chamber, or ventricle, and the arrange- ment of their vascular connections in such a way that two complete circulations are established. One of these is the long, or systemic circula- tion. It is concerned with the distribution of blood to the various parts of the body, with the exception of the lungs. It is established by the left ventricle, the aorta, the carotid and subclavian branches of its arch, and the parietal and visceral branches of its thoracic and abdominal portions. The blood is collected from the anterior portions of the body through paired internal and external jugular and subclavian veins, com- municating with the right atrium of the heart through paired superior cavals; from the posterior portions of the body through the unpaired and also asymmetrical inferior caval vein, the latter passing forward on the right of the median plane and entering the posterior end of the right atrium. The second, short, or pulmonary circulation, is concerned with 44 ANATOMY OF THE RABBIT. the distribution of the blood to the lungs for purposes of aeration. It is established by the right ventricle, the pulmonary artery and its paired branches, and by the “capillaries ah the lungs. The blood is delivered to the left atrium through several pulmonary veins. A similar division of the circulatory organs occurs as a homoplastic modification in birds, which, it will be observed, are also warm-blooded vertebrates. Many of the peculiar features of the mammalian circulation which at first sight do not appear to be general, but are so in reality, depend on the circumstance that the complete partition of the organs is a final stage of a general progressive development, observable in air-breathing verte- brates, in which the lungs and their vascular connections become perfected for pulmonary respiration. On the other hand, the vascular system as it appears in the embryo, more especially its aortic portion, is arranged according to the type of branchial respiration as found in fishes. In this condition the blood is sent forward from the heart through a ventral aorta. The latter is connected with a series of paired branchial aortic arches, traversing the rudimentary gill structures, and thus passing upward around the sides of the primitivé pharynx. The dorsal aorta is formed by the junction of the branchial aortic arches, and passes backward as a main distributing vessel on the ventral side of the axial support. The heart itself is formed primarily on two-chambered plan, similar to that in fishes, where all the blood is received by single atrium and is delivered forward to the gills by a single ventricle. The definitive condition of the chief Fic. 24. Plan of the branchial aortic arterial vessels is arrived at by an extensive arches. The adult mammalian vessels are indicated in black (systemic) “or modification of the branchial plan. AS shaded (pulmonary). 1-6, primary : A ‘i os ‘i arches; ao., aorta; a.p., pulmonary indicated in the accompanylng diagram artery; c.e., external carotid; c.i., in- : 9 Dio 2 . ternal carotid; d.a., ductus arteriosus (Fig. 24) the arch condition is retained by (Botalli); i., innominate artery: s.d., the aorta and by the pulmonary artery, Ge Weeke she Bey aC URe and it is interesting to note also that the primary connections of these vessels, repre- sented in the embryo by an open canal, the ductus arteriosus (Botalli), is indicated in the adult condition by a short fibrous cord, the arterial ligament. The adult aortic arch represents only the left one of a pair, and since that of the right is only represented imperfectly by the innominate artery and the base of the subclavian of that side, a condition of asymmetry results, which is mainly expressed by the sinistral position of the arch with reference to the ‘oesophagus (Plate VII). By comparison with the embryonic plan, it is seen that the primitive features of the heart and the arterial vessels include the ventral position of the heart itself, the equivalence of the two atria and of the two ventricles—these structures being partitioned internally but THE VASCULAR SYSTEM. 45 imperfectly divided externally—the forward position of the first portion of the aorta, and the position of the aorta as a median vertebral trunk. The vascular system is noteworthy for several departures from the condition of symmetry, one of these having already been mentioned. In addition, it is seen that ina mammal,as in terrestrial vertebrates generally, the base of the pulmonary artery is rotated in a spiral fashion about the base of the aorta, so that from its beginning on the right ventricle it passes across the ventral surface of the base of the aorta, and divides on the dorsal side of the latter into its two main branches. Moreover, the separation of the ventricular portion of the heart into two chambers is associated with an enormous increase in the muscularity of the wall in the left ventricle, or, in other words, in that portion which is concerned with the larger, systemic circulation. The inferior caval vein (Plate VIII), a highly specialized vessel, is asymmetrical, since from its begin- ning in the pelvic cavity to its termination on the right atrium it lies wholly to the right of the median plane. The azygos vein of the thorax (Plate VII), a vessel uniting the majority of the paired intercostal veins, and interesting as a remnant of the primitive circulation, is also asymmetrical, since the trunk lies to the right of the bodies of the vertebrae, and is connected at its base with the right superior caval vein. In general, the blood which is distributed to the various parts of the body passes through but one set of capillary vessels, and is then returned through the systemic veins to the heart. In all vertebrates, however, a special portion of the systemic venous circulation is set aside as the hepatic portal system, distinguished by the possession of a second series of capillary vessels ramifying in the liver. Thus, in the rabbit, the blood distributed to the stomach, spleen, and intestine through the coeliac, superior and inferior mesenteric arteries, is collected into a main intestinal vessel, the portal vein, and the latter, approaching the liver through the lesser omentum, divides in that organ into a series of portal capillaries. The portal capillaries, like the systemic capillaries proceed- ing from the hepatic artery, unite in the tributaries of the hepatic veins. In lower vertebrates, although not in the mammalia, a second system of venous capillaries occurs in connection with the kidneys, and is known as the renal portal system. The lymphatic system, both in its functional relation and in origin, is an appendage of the venous portion of the vascular system. The system is an important one, of which, unfortunately, little may be seen by ordinary dissection, the structures which appear in this way being the lymph glands, or lymph nodes, centres of cell formation, occurring in the course of the conducting vessels. These as superficial structures are found either singly, asin the head and neck, or more or less grouped, as in the axillary and inguinal spaces. As deep structures they are conspicuous in the intestinal mesenteries, and in the walls of the digestive tube, occurring in the latter chiefly as continuous masses of lymph follicles, as, for example, in the walls of the sacculus rotundus, the vermiform process, or the tonsil; or, again, as aggregated lymph follicles (Peyer’s patches) at various points in the intestinal wall. The conducting portion of the system comprises an extensive series 46 ANATOMY OF THE RABBIT. of canals, beginning as lymphatic capillaries in peripheral organs, and ending as lymphatic trunks which empty into the great veins. The lymphatic trunks of the anterior portion of the body are designated from their association with the corresponding veins as jugular and subclavian. They enter the venous system on either side at the point of junction of the internal and external jugular veins or of the common jugular and subclavian (Fig. 52). The lymphatic vessels of the posterior portion of the body, including the intestine, unite to form a common canal, the thoracic duct. The latter lies for the most part between the aorta and the verte- bral column, and traverses the thorax in this position to enter the venous system at the same point as the jugular and subclavian trunks of the left side. The lymphatic capillaries are terminal, absorptive vessels, differing from blood capillaries both in the character of their walls and in their relations to other portions of the system, since they are not interposed as in the vascular system between vessels of a larger order. The lym- phatic vessels connecting the capillaries with the lymphatic trunks form extensive plexuses, in connection with which the lymph nodes are dis- tributed. THE URINOGENITAL SYSTEM. The urinogenital system comprises two primary systems—reproductive and urinary—differing widely in their central organs, but associated to a certain extent by having common ducts. In the rabbit, as indicated in the accompanying diagram (Fig. 25), this association extends only to the presence in the two sexes of a urinogenital canal, or urinogenital sinus connecting both urinary and genital structures with the outside of the body. This canal is designated in the male as the urethra, but in the female as the vestibulum, since the structure known from the human relation as the female urethra is only a urinary canal leading from the bladder, and in man is not associated with the reproductive ducts. In general, however, the relations of the urinary and reproductive organs involves two chief features. First, in primitive vertebrates, the urinary and genital ducts open into the posterior end of the digestive tube, the latter forming in this relation a common canal, the cloaca. In terrestrial vertebrates, the urinary bladder is developed as a ventral outgrowth of the digestive tube, and, exeept in amphibians, both sets of ducts undergo a migration from their original position on to the wall of its canal, the latter being thus transformed into a urinogenital sinus. This development reaches its extreme in the higher mammalia, where the urinogenital sinus is completely separated from the digestive tube, and where the urinary ducts are also transferred from a posterior or hypo- cystic position on the wall of the urinogenital sinus to an anterior or epicystic position on the dorsal wall of the bladder. Secondly, there is a more fundamental association between the reproductive and excretory organs, depending on the circumstance that the former are primarily in the vertebrates organs connected with the THE URINOGENITAL SYSTEM. 47 lining of the coelom and discharging their products into the cavity, while the kidneys are primarily tubular structures communicating also with this cavity, so that they become modified in part as reproductive ducts. The coelomic connections of the kidney tubules are with few exceptions only seen in the embryonic condition, since the definitive kidneys in all vertebrates are structures in which the tubules are connected with the vascular system, and have either lost, or, as in the specialized permanent kidneys of the mammalia, have not developed, the coelomic apertures. The embryonic development of a mammal includes the formation not only of a final kidney or metanephros, but also of two embryonic structures, one of which, the first kidney or pronephros, is embryonic in all vertebrates, while the second, designated as the mesonephros, or embryonic kidney, is one occurring in the adult condition of intermediate Fic. 25. Diagrams of the male (A) and female (B) urinogenital systems of the rabbit: b., urinary bladder; k., kidney; ur., ureter; r, rectum. (A)—c.p., crus penis; d.d., ductus deferens; ep., epididymis; g., gubernaculum; t., testis; v.s., seminal vesicle; u.v., male urethra. (B)—c.c., crus clitoridis; o0., ovary; t.u., uterine tube; u.m., female urethra; ut., uterus; va., vagina; vs., vestibulum. forms. It is from the ducts of the primary kidneys that the reproductive ducts arise. Thus, in the female of most vertebrates the oviduct opens by an expanded funnel into the coelomic cavity, usually at some distance from the ovary; and, although in the rabbit the structure is said to arise as a secondary dev clopment, it exhibits here as in other mammals the feature of a coelomic opening, coupled with that of close association with 48 ANATOMY OF THE RABBIT. the ovary, so that the products of the latter, while nominally discharged into the coelom, are actually received directly into the uterine tube. The oviduct itself is phylogenetically the oldest of the urinogenital ducts, since it is identifiable in the embryo as the duct of the pronephros. In the rabbit, as in all mammals, the testis is connected with the urethra through the canal formed by the epididymis and ductus deferens, these structures comprising in a modified form a portion of the mesonephros and its duct. Since the excretory functions are fully provided for by the development of a permanent kidney and its duct, the ureter, the ducts of the embryonic kidney have in the adult no urinary connections; but in intermediate vertebrates, in which the mesonephros occurs as an adult kidney, the interesting condition is observable in the male that the duct of this structure serves both urinary and reproductive. functions. It will be seen in the rabbit that the female genital ducts, apart from the common urinogenital sinus, include an unpaired portion, the vagina, and a paired portion, comprising the uteri and uterine tubes. In all vertebrates up to and including the monotrematous mammalia the oviducts open separately either into the cloaca, or into the urino- genital sinus. In the higher mammalia, however, a process of fusion is observable, extending from the partial coalescence of the vaginae, as in marsupial mammals, to the complete coalescence of both vaginae and uteri, as in man. Thus, there are recognized in the mammalia the types of (a) unpaired uterus—uterus simplex; (b) semi-divided uterus—uterus bicornis; and (c) completely divided uterus, such as that of the rabbit—uterus duplex. In the respective positions of the central organs, as exemplified by the rabbit, there are several features of general significance. Thus, the kidneys (Plate VIII) are paired structures lying on the dorsal wall of the abdominal cavity, where they are interposed between the peritoneum and the dorsal musculature. They are covered by peritoneum only on their ventral surfaces. As indicated above, the permanent kidneys of a mammal are highly specialized structures, but they show in these rela- tions features common to lower forms and dependent in both cases on the primary position of the intermediate mass, from which in the em- bryo the kidneys are formed. The gonads are primarily associated with the dorsal lining of the coelomic cavity, although certain observations on the development of these structures in the lower vertebrates appear to indicate that their elements are assembled from other parts of the embryo. In some of the mammalia, as in all lower vertebrates, the testis occupies an abdominal position in the adult condition; but usually it undergoes an extensive mi- gration, passing from the abdominal cavity into a special sac of peritoneum enclosed by the scrotum. This change—described as the descent of the testis—is effected through the agency of a muscular cord, the guber- naculum. In many cases, as in man, the cavity enclosing the testis is completely separated from the abdominal cavity, but in the rabbit a more primitive condition is retained in which the sac of the testis is widely open to the abdominal cavity,and the organ thus passes freely THE SEROUS CAVITIES. 49 from one cavity to the other. It is also observable in this animal that the gubernaculum is represented in the adult as a short thick cord connecting the end of the testis with the wall of the enclosing sac (Fig. 26). In the female the round ligament of the uterus is a structure similar in a general way to the gubernaculum, and in the rabbit will be seen to be inserted in a depression of the body-wall resembling both in form and position a rudimentary vaginal process. The ligament is continued, however, beyond this point, ending in the wall of the urinogenital aper- ture. The presence of this structure is an indication that the ovary, as well as testis, is subject to change in position. In the adult condition, however, the ovary occupies approximately an original position on the dorsal wall of the abdominal cavity ; and it will be observed in the rabbit that the position of the structure is one also defined by the points of origin of the spermatic arteries and veins of both sexes, with the ex- ception, however, of the left spermatic vein in the male. THE SEROUS CAVITIES. The organs collectively described as visceral are those associated with the serous cavities. They belong to several systems, but present the common feature of being projected into the membranous linings of these cavities so that they are more completely invested by them. The serous sacs are extensive body-spaces, derivatives of a primary body cavity or coelom. They are usually considered as containing the visceral organs, but the condition is more accurately described as one in which the visceral organs encroach, chiefly from a dorsal position, on the enclosing membranes. The latter are thus divided into two portions, one of which is distributed as a parietal or peripheral layer, forming the enclosure of the sac, while the other is disposed as a visceral layer on the surface of the visceral organs. The serous sacs are enclosed by thin, moist, serous membranes, consisting chiefly of mesothelium, which give to the visceral organs their characteristic appearance. In lower vertebrates, where the diaphragm is absent or imperiectly developed, the coelom is divided into two chief portions — the pericardial cavity, enclcsing the heart, and the pleuroperitoneal cavity, lodging the remaining visceral organs, including in terrestrial vertebrates the lungs. In the mammalia the pleurcperitoneal cavity is completely divided into two portions by the diaphragm, the smaller pleural portion being again divided into right and left pleural cavities through the presence of certain structures filling the median portion of the thorax. There are thus recognizable in a mammal four large serous spaces, namely, the pericardial, peritoneal, and paired pleural cavities. The pericardial cavity, the smallest of these spaces, is situated between the paired pleural cavities. Its enclosing membrane, the peri- cardium, forms a capacious sac for the heart, and is reflected directly over the surface of the latter as a thin membrane, the epicardium. The pleural cavities are those lodging the lungs, the latter being pro- jected into them froma medial position. The lining membrane or pleura 5 50 ANATOMY OF THE RABBIT. is divided into three chief portions—the pulmonary pleura, investing the greater part of each organ, the costal pleura, lining the internal surface of the thorax, and the diaphragmatic pleura, covering the anterior surface of the diaphragm. The latter is broadly connected with the pulmonary pleura through the pulmonary ligament. The peritoneal cavity, the largest of the serous spaces, comprises in a mammal a general portion, the abdominal cavity, and its posterior ex- tension into the pelvis—in the male also into the sac of the testis. The general relations of the cavity to the abdominal organs is indicated dia- grammatically in Fig. 21. Its lining membrane, the peritoneum, is divisible into two portions, the parietal peritoneum, lining the abdominal wall, and the visceral peritoneum, investing the visceral organs. Of the latter the kidneys encroach only to a minor extent on the serous lining, so that they are covered by peritoneum only on their ventral sur- faces. The digestive tube, on the other hand, is removed to such an extent from the abdominal wall that the peritoneum forms a complete serous coat, and is connected with the parietal peritoneum of the wall through a thin transparent mem- brane, the mesentery. The latter consists of two plates of peritoneum, enclosing between them a thin layer of connective tissue, the lamina mesenterii propria, for the trans- mission of nerves, bloodvessels and lymph canals. As indicated above, the relations of the abdominal portion of the diges- tive tube are greatly modified by its elongation and displacement from a median position. Thus, while in the embryo the common mesentery is Fic. 26. Diagram showing Ithe relation of recognizable as a continuous median the testis to its investments: ai. inguinal vertical fold, in the adult it follows ring; c.e., caput epididymidis; cr., cremaster , : : muscle: did., ductus deferens; g., guber- the convolutions of the digestive tube, naculum:. (mes, mesorchium:' piv ard and is therefore considered as divided vaginalis propria; p.v., cavity of the vaginal into corresponding parts. In many process; s., integument of the scrotum; ; S.v., spermatic vessels; t., testis. cases the relations of these are greatly complicated by secondary adhesions. In the rabbit the mesoduodenum, mesentery, and descending mesocolon will be recognized as parts in which a more typical arrangement is retained. Moreover, in the anterior portion of the abdominal cavity the peritoneum is concerned not only with the investment of two large visceral structures, the stomach and the liver, but also with the formation Ge a lining for the posterior surface of the diaphragm. Thus the general condition is less simple than in the small and large intestines. The peritoneum, THE SEROUS CAVITIES. Sil passing from the dorsal wall, successively invests the spleen, the stomach, and the liver, and is reflected from the last-named structure to the diaphragm and the ventral body-wall through the coronary, triangular, and falciform ligaments. Its gastric portion is differentiated into the mesogastrium (phrenicosplenic and gastrolienal ligaments), the greater omentum, and the lesser omentum. Similarly, in the posterior part of the body the peritoneum passes from the rectum to the urinary bladder, enclosing also in the female the vagina. It is then reflected to the ventral body-wall as the middle umbilical fold. In the male, as indicated in the accompanying diagram (Fig. 26), the peritoneal relations of the testis are greatly modified by the migration of the organ from an abdominal to a scrotal position. The entire sac lodging the testis is an evaginated portion of the abdominal wall, and since in the rabbit the cavity is widely open throughout life to the abdominal cavity, the lining membrane—that designated as the parietal layer of the tunica vaginalis propria—-is continuous with the parietal peritoneum of the abdomen, and thus represents a permanent vaginal process. Like other structures of the abdominal cavity, the testis itself is covered by peritoneum, the latter being designated as the visceral layer of the tunica vaginalis propria. This investment is ccnnected with the parietal layer by the mesorchium, and in the rabbit it will be observed that the latter is chiefly attached forwards on the dorsal wall of the abdomen, 7.e., in a position indicating the original situation of the testis itself. In the female the ovary is closely associated with the dorsal wall of the abdomen, and its supporting peritoneum, the mesovarium, is in- significant. Its duct in passing backward, however, becomes greatly displaced from a dorsal position, and thus comes to be supported by a broad fold of peritoneum. The latter is considered to consist of two portions, one, the mesosalpinx, being the support of the uterine tube, the other, the mesometrium, that of the uterus. » The entire fold, how- ever, forms a continuous structure, and is known in this relation as the broad ligament. REGIONAL SECTIONS. The following plate-figures (I-VIII) are from characteristic sections of a rabbit-foetus of 56mm., and may be used either in connection with the general features of topography as outlined above, or for the identi- fication of various minor structures appearing in the dissection. Certain points regarding the sections are perhaps worthy of notice. First, in the longitudinal section illustrated in Plates I and II it will be noticed that paired structures frequently appear; this being because of the fact that the section is not exactly median, at least in certain places. Secondly, in using sections of the foetus for gross anatomical features it is necessary to make allowance in some cases for the different propor- tions of organs, and consequent slight differences in position, in the foetal as compared with the adult condition. Finally, many of the features appearing in the original sections are such as could not be repro- duced in the plates, although they are indicated in the accompanying skeleton figures, and may be referred to in this way. “WINUIIIG “77 *SOUTJSOJUT “ST ‘uojeydaouargq * *suUOT}O9S OSIOASURI} BUIPUOdsat *AqIAeS [eIpreotiag “gz ‘useiydeliq “ZT “uojeydaouasayy * “109 JO S[9A9] 9}eOIPUT [TT A-TIT “B{1Oe pure Jivazy ‘cz *(TeIqay19A01d) “uojeydsaouajayy * ‘uinjdas [eseN “FE “APAR [VIN “FZ UOl[sURs OllayuasauI IOlIadNg ‘vgT “uoTeYydaoua]ad Jy “UINTUBIOISEG “Ee ‘BUNT “EZ “yunty orjayyedurAs jo elysuery ‘oT *pioo yJeutdg “AYIABS [VIO “ZE *(sTassoa “SIAT@9g “GT “AYIARO [PIURID ‘ansuol “Te pue pro9 jo aseq) snoriquig “cz “OBIQIJIOA [VOTAIOD “PT ‘Teued [e1qa119 A, “‘Bayoely, “Oe . TOAVT “TZ “OBIGOJIOA OINVIOYUTL “ET ‘ainyemMosnur [eurdg ‘snseydosag “6% *(uort0d or10TAd) yoeurojg ‘QZ “ovIqeayI9A TeqUIN’T “ZT “you JO sosseul-jeyy * ‘purls snuAyy “9z ‘AYIARO [RUIWIOpqY “6 ‘uojeydeaousjay, “TT ‘9JOSNUI SNHWIxXeU snouRyND * ‘I €LV1Id YOU SNOILVNOISAa an HAO dHiSON “ACOG AUMIINY BAHL AO NOILOGS ITVOILYGA NVIGAYW V idl DESIGNATIONS FOR PLATE II. 1. Transverse sinus of dura mater. 32. Maxilla. 2. Dura mater. 33. Hard palate (palatine and maxilla). 3. Pallium of cerebral hemisphere. 34. Presphenoid. 4. Lateral ventricle. 35. Intersphenoidal synchondrosis. 5. Olfactory bulb. 36. Basisphenoid; hypophysial fossa. 6. Olfactory tract. 37. Sphenooccipital synchondrosis. 6a. Divided olfactory nerve in the cribriform 38. Basioccipital. plate. 38a. Supraoccipital. 7. Chorioid plexus of third ventricle. 39. Nasal portion of pharynx. 8. Anterior commissure. 40. Soft palate. 9. Thalamus. 41. Oral portion of pharynx. 10. Optic chiasma. 42. Epiglottis and epiglottic cartilage. 11. Tuber cinereum. 43. Thyreoid cartilage of larynx. 12. Mammillary body. 44, Laryngeal cavity. 13. Superior colliculus. 45, 45a. Cricoid cartilage. 14. Inferior colliculus. 46. Oesophagus. 15. Anterior medullary_velum. 47. Cricothyreoideus muscle. 16. Cerebral peduncle; cephalic flexure. 48. Thyrcoid gland. 17. Isthmus rhombencephali. 49. Sternohyoideus muscie¢. 18. Fourth ventricle. 50. Genioglossus muscle: 19.4Pons; pontine flexure. 51. Geniohyoideus muscle. 20. Cerebellum. 52. Mylohyoideus muscle. 21. Posterior medullary velum. 53. Mandible. 22. Cervical flexure. 54. Occipital musculature. 23. Central canal of spinal cord. 55. Semispinalis capitis. 24. Hypophysis. 56. Rhomboideus minor. 25. Frontal bone. 57. Superior portion of trapezius. ) 58. Atlas. 26. Nasal bone. § 27. Nasal fossa. 59, Epistropheus. 28. Mesethmoid cartilage. 59a. Odontoid process. 29. Cartilage of vomeronasal organ. 60. Third cervical vertebra. 30. Premaxilla. 61. Median vertebral vein. 31. Nasopalatine duct and cartilage. 62. Body of hyoid bone. o ~ De A MEDIAN VERTICAL SECTION OF THE HEAD, . Nasa 2. Leva 3. Nasa . Naso 5. Maxi }. Nasa . Naso 3. Vome DESIGNATIONS FOR PLATE III. bone. or alae nasi muscle. septum. turbinal cartilage. fossa. acrimal duct. Premaxilla. ; Smal upper incisor. . Large upper incisor. 2. Naso 3. Oral valatine ducts. cavity. loturbinal (concha inferior). -ronasal organ and cartilage. . Tongue. . Vibrissae. ). Caninus muscle. . Terminals of superior maxillary nerve. Buccal glands. . Buccinator muscle. . Terminals of inferior alveolar nerve. . Quadratus labii inferioris muscle. 2. Mandible. 23. Lower incisor. . Meckel’s cartilage (primary mandibular arch). . Mentalis muscle. IBGE A TRANSVERSE SECTION OF THE ANTERIOR NASAL REGION. 57 DESIGNATIONS FOR PLATE IV. 1. Superior sagittal sinus of dura mater. 25. Nasal tract; choana. 2. Lateral ventricle. 26. Palatine bone. 3. Cerebral hemisphere. 27. Oral cavity. 4, Pia mater. 28. Palatine nerve. 5. Frontal bone. 29. Sphenopalatine ganglion. 6. Cartilage of orbital wing. 30. Infraorbital vein. 7. Mesethmoid cartilage. 31. Internal maxillary artery 8. Cupula posterior cartilage. 32. Maxillary nerve. 9. Obliquus superior muscle. 33. Maxilla. 10. Ophthalmic vessels and nerves. 34. Zygomatic bone. 11. Levator palpebrae superioris muscle. 35. Submaxillary duct. 12. Rectus medialis muscle. 36. Buccinator muscle. 13. Retractor bulbi muscle. 37. Masseter muscle. 14. Rectus inferior muscle. 38. Parotid duct. 15. Sclera. 39. Facial nerve. 16. Retina and chorioidea. 40. External maxillary artery and vein 17. Vitreous body. (anterior facial vein). 18. Lens. 41. Platysma muscle. 19. Posterior chamber of eye. 42. Inferior labial artery and vein. 20. Anterior chamber. 3. Mandible. 21. Cornea. 44. Genioglossus muscle. 22. Ciliary body and iris. 45. Digastricus muscle. 23. Upper eyelid. 46. Quadratus labii inferioris muscle. 24. Lower eyelid. 47. Geniohyoideus muscle. 58 IV. A TRANSVERSE SECTION OF THE ORBITAL REGION. DESIGNATIONS FOR PLATE V. Longus capitis. Rectus capitis anterior. 3. Oral portion of pharynx. . Thyreohyoideus muscle. 5. Sternohyoideus muscle. 1. Parietal bone. 2. Transverse sinus of dura mater. 3. Superior colliculus. 4. Cerebral aqueduct. 5. Isthmus rhombencephali. 6 7 g YVNbwwbbb IDoPwWNwe ~ Pons: 1. Greater cornu of hyoid. . Trigeminal nerve. 27. Stylohyoideus major muscle. 8. Basilar artery. 28. Lingual artery. 9. Facial nerve. 29. Hypoglossal nerve. 10. Cartilaginous auditory capsule. 30. Tendon of digastricus muscle. 11. Cochlea. 31. External maxillary artery. ) Stylohyoideus minor. Styloglossus. Internal maxillary artery. Tympanic bone. Mandible. Submaxillary gland. 38. Anterior facial vein. 39. Internal carotid artery. 1) Basioccipital bone. Tensor tympani muscle. Tympanic cavity. Malleus. 6. Tributari.s of posterior facial vein. Squamosal bone. 8. Cephalic portion of median vertebral vein 19. Nasal portion of pharynx. 20. Origin of basioclavicularis and levator scapulae major muscles. NOL wl SD Or > “J ] 1 ] ] ] ] ] 60 A TRANSVERSE SECTION OF THE AUDITORY REGION. 61 DESIGNATIONS FOR PLATE VI. 1. Rhomboideus minor. 19. Oesophagus. 2. Superior portion of trapezius. 20. Inferior thyreoid nerve. 2a. Levator scapulae minor. 21. Inferior thyreoid vein. 3. Splenius. 22.° Trachea. 4. Semispinalis capitis. 23. Thyreoid gland. 5. Rectus capitis posterior superficialis. 24. Cardiac branch of vagus (n. depressor) 6. Obliquus capitis major. 25. Sympathetic trunk. 7. Arch of epistropheus. 26. Vagus nerve. S. Ganglion of posterior root. 27. Common carotid artery 9, Longissimus cervicis. 28. Internal jugular vein. 10. Longissimus capitis. 11. Vertebral artery and vein. 12. Longus atlantis. 13. Vertebral body. Transverse process (anterior root). Median vertebral vein. Longus colli. Longus capitis. Fat-body. . Sternohyoideus muscle. . Sternothyreoideus muscle. . Sternomastoideus muscle. . Descending ramus of hypoglossal nerve. External jugular vein. Basioclavicularis muscle. . Levator scapulae major muscle. . Cleidomastoideus. Platysma. ae pa CON Oe WwWwWwWWwWwWwWWW bl TOO PwWNWr Ow} 62 Vi A TRANSVERSE SECTION OF THE ANTERIOR CERVICAL REGION. oo Gs DESIGNATIONS FOR PLATE VII. Semispinalis dorsi. Longissimus dorsi. Iliocostalis. Spinal cord. Ganglion of posterior root and intercostal nerve. Tubercle of rib. Head of rib. Sympathetic trunks. Azygos vein. . Thoracic aorta. . Oesophagus. 12 12a. Right and left vagi. 13. Lung. 14. Bronchi. 15. Branches of pulmonary artery. 16. Pulmonary veins. 17. Right atrium. 18. Tricuspid valve. 19. Right ventricle. 20. Left atrium. 21. Left ventricle. 22. Pericardial cavity. 23. Pulmonary pleura. 64 24, 25. 26. 27 28. 29. 30. <2, 2) 36 9 Or 2 o 9 oO 2 3 4, de 1, Costal pleura. Bone ribs. Costal cartilage. . Sternum. Cutaneus maximus muscle. Inferior portion of trapezius Rhomboideus major. Inferior angle of scapula. . Latissimus dorsi. . Serratus posterior. Intercostales externi and interni Intercostalis internus. Thoracic portion of serratus anterior. Obliquus externus abdominis. . Transversus thoracis. Pectoralis major. Rectus abdominis. Anconaeus longus (caput longum of triceps). Extensor antibrachii parvus. . Anconaeus medialis. . Anconaeus lateralis. . Distal extremity of humerus. Proximal portion of radius. Vi A TRANSVERSE SECTION OF THE THORAX. 65 ee ee ep DESIGNATIONS FOR PLATE VIII. Spinal cord. 18, 18a. Posterior and anterior lobules of left Vertebral canal. lobe of liver. penebre pee. Au 19, 19a. Right lobe of liver. MONO OTS Leen: 20. Obliquus internus abdominis and trans- TH CODD Quadratus lumborum. oom 5 YA PERE peta versus abdominis. Psoas major. 21. Obti Sra des SNE PETER Peoiermingr 21. ligquus externus abdominis. _ aie 99 ns 4 na Sympathetic trunk. 22. Rectus abdominis. Abdominal aorta. 22a. Cutaneus maximus. tre Cobo orks) Inferior caval vein. 23. Middle umbilical fold. Descending mesocolon. 24. Urinary bladder (canal of foetal allantois). Ureter. 25. Umbilical arteries. Renal pelvis. 26. Duodenum. Renal papilla. 27. Pancreas and mesoduodenum. Left kidney. 28. Descending colon. Parietal peritoneum. 29. Parts of mesenterial small intestine. Visceral peritoneum. 20. Caecum. 66 VIII. A TRANSVERSE SECTION OF THE ABDOMEN. PART If OSTEOLOGY*OF THE RABE: For a practical study of the rabbit’s skeleton, a thoroughly cleaned, but otherwise rough, unmounted skeleton will be found most convenient. The skull should be divided with a fine saw at a little to one side of the median plane, or a second skull may be provided for this purpose (cf. Fig. 33). The most useful specimens for reference are: (1) a well- mounted skeleton of the adult animal, showing the natural relations of the bones; and (2) a rough skeleton of a young animal of from one to five weeks, showing the primary composition of cartilage bones. For the special study of the skull { (pp. 85-97) a disarticulated specimen may be employed, but the majority of the features may be made out in the intact or divided skulls. The general account of the skull as given below will be found to cover most of the osteological points noted in the dissection. DIVISIONS OF “GEE SK DEE RON: The skeleton is divisible into two main portions, namely, the axial skeleton and the appendicular skeleton. The former comprises the vertebral column, the ribs, the sternum, and the skeleton of the head; the latter, the supports of the anterior and posterior limbs, and the associated pectoral and pelvic girdles. THE VERTEBRAL COLUMN: The vertebral column (columna vertebralis) is formed of a lnear series of segments, the vertebrae. In accordance with its function asa general support of the body, and also its relations with the nervous system and the spinal musculature, the vertebrae, with minor exceptions, are constructed on the same plan. Those of particular regions also present certain features in common, so that it is possible to classify them into cervical, thoracic, lumbar, sacral, and caudal groups. A typical vertebra—for the characters of which any one of the thoracic or lumbar series may be taken (Fig. 27, D-F)—consists of a basal portion, the vertebral body (corpus vertebrae), and of a dorsal, vertebral arch (arcus vertebrae). The two portions enclose a large aperture, the vertebral foramen (foramen vertebrale). The successive foramina form an almost complete tube, the vertebral canal (canalis vertebralis), for the accommodation of the spinal cord. The body of a vertebra is a cylindrical, or somewhat dorsoventrally compressed, mass of bone, which bears at either end an articular surface for attachment to the adjacent vertebra... The articular surfaces are borne on thin plate-like epiphyses, the epiphysial lines being evident even in older animals, especially in the lumbar region. The dorsal por- 68 THE VERTEBRAL CoLUMN. 69 tion of the body bears on either side the pedicle, or root of the vertebral arch (radix arcus vertebrae), the dorsal surface of the body forming in this way the floor of the vertebral foramen. The dorsal portion of the arch, borne on the pedicle, is distinguished as the lamina. The anterior and posterior margins of the pedicle are notched, each notch psa. Fic. 27. Representative vertebrae: A, atlas, anterior surface; B, epistropheus, lateral surface; C, fifth cervical vertebra, anterior surface; D, fourth dorsal, lateral surface; E, F, second lumbar vertebra, anterior and lateral surfaces. a.a., anterior arch of atlas; a.p., posterior arch of atlas; a.v., vertebral arch; c.v., vertebral body; d., dens epistrophei; f.a.a., anterior articular facet of epistropheus; f.a.s., superior articular pit of atlas; f.a.s.1., superior articular facet of epistropheus; f.c.i., inferior costal demifacet for head of rib: f.c.s., superior costal demifacet; f.c.t., costal facet of transverse process; f.d., fovea dentis; f.i., intervertebral foramen; f.tr., foramen transversarium; f.v., foramen vertebrale; 1., lamina of vertebral arch; m.l., lateral mass of atlas; p.a., accessory process of lumbar vertebra; Pp.a.i. inferior articular process; p.a.s. superior articular process; p.m., mammillary process; Pp.s., spinous process; p.S.a., anterior spinous process; p.t., transverse process; p.tn., trian- gular process; r. radix of vertebral arch; r.a., r.p., anterior and posterior radices of transverse process of cervical vertebra; t.a., t-p., anterior and posterior tubercles of atlas. or incisure being converted, through its association with that of the adjacent vertebra, into a rounded aperture, the intervertebral foramen (foramen intervertebrale), for the passage outward of a spinal nerve. 7 70 ‘ANATOMY OF THE RABBIT. The arch of the vertebra is noteworthy for its projections or processes. On either side is a horizontal plate of bone, the transverse process (pro- cessus transversus), and, dorsally, a median projection, the spinous process (processus spinosus), all three serving for the attachment of the vertebrae to one another by ligaments, and for the attachment of the spinal musculature. Special articular surfaces, borne on low articular pro- cesses (processus articulares), are found on the anterior and posterior margins of the arch. The anterior, or superior articular surfaces are directed for the most part toward the dorsal surface, and are overlapped in the natural condition by the inferior articular surfaces, which are directed toward the ventral surface. A certain amount of movement is permitted by one surface slipping across the other, the mechanism illustrating the arthrodia, or gliding-joint. The cervical vertebrae (vertebrae cervicales) are seven in number. The posterior five are similar, while the anterior two are specially modified in relation to the skull. The posterior vertebrae (Fig. 27, C) are dorso- ventrally compressed, their arches low, and the spinous process short. In the seventh vertebra, however, the spinous process begins to be elon- gated as in the succeeding thoracic vertebrae. In each vertebra the transverse process is perforated by a costo-transverse foramen (foramen transversarium), the latter serving for the passage of the vertebral artery forward to the head. Through the presence of this aperture, the base of the transverse process is divided into two parts, namely, a dorsal, or posterior root (radix posterior), and a ventral, or anterior root (radix anterior). The anterior root is a coalesced rib, and is comparable in its general relations to the normal ribs of the thoracic vertebrae. The first vertebra is the atlas (Fig. 27, A). It is peculiar in lacking the vertebral body, the latter being represented by the odontoid process of the epistropheus (cf. Plate II); also in possessing special articular surfaces, and in having its transverse process greatly flattened in the dorsoventral direction. It consists of a ventral half-ring, the anterior arch (arcus anterior), a dorsal half-ring, the posterior arch (arcus pos- terior), with paired lateral masses (massae laterales) uniting them. The lateral masses also form the bases of the transverse processes. The anterior arch bears on its ventral side a small backwardly-directed process, the anterior tubercle (tuberculum anterius). A similar posterior tubercle (tuberculum posterius) on the dorsal surface of the posterior arch is comparable to the spinous process of an ordinary vertebra. The anterior surface of the atlas bears on either side an extensive concave smooth surface, the superior articular pit (fovea articularis superior), for articulation with the convex occipital condyles of the skull. Its posterior surface bears on either side a somewhat triangular inferior articular facet (facies articularis inferior) for articula- tion with the epistropheus. These surfaces take the place of the arch- articulations of ordinary vertebrae. Through the compression of the transverse process, the costotransverse foramen is converted into a canal. The anterior aperture of this leads by a shallow groove, the sulcus arteriae vertebralis, into a second aperture perforating the posterior arch. THE VERTEBRAL COLUMN. 71 The space enclosed by the atlas is divided into a dorsal portion, corresponding to the vertebral foramen of other vertebrae, and a ventral portion which in the natural condition lodges the odontoid process of the epistropheus. The division is effected partly by a small tubercle on the inner side of each lateral mass, and partly by a _ transverse liga- ment which is stretched between them and over the dorsal surface of the odontoid process. On the floor of the ventral portion, a rounded articular surface, the fovea dentis, marks the point of articulation of the anterior articular facet of the odontoid process with the inner surface of the anterior arch. The second vertebra is the epistropheus (Fig. 27, B). It resembles the succeeding cervical vertebrae more closely than does the atlas. It is noteworthy for its great size, for the lateral compression of its arch and spinous process, and for the possession of a stout forwardly-directed odontoid process, or tooth (dens epistrophei). It is articulated with the atlas through an anterior articular facet, borne on the ventral surface of the odontoid process, and by large paired superior articular facets borne on its base. The spinous process of this vertebra and the trans- verse processes of the atlas form together three main points of attach- ment for the occipital musculature. The thoracic vertebrae (vertebrae thoracales) are twelve in number. They are distinguished chiefly by the possession of articular pits for the attachment of ribs (Fig. 27, D). A rib is articulated at two. points, namely, one on the body of the vertebra, the other on the transverse process. The former is marked by a small round depression, the costal pit (fovea costalis), or costal facet. In the last two vertebrae the facet is borne wholly on the vertebral body to which the rib belongs. In the remaining vertebrae a complete articulating surface is formed by two demifacets, one being on the vertebra to which the rib belongs, the other on the vertebra immediately in front. The articulation of a rib with a transverse process is marked by an oval facet, the costal pit of the transverse process (fovea costalis transversalis). It is present only in the first ten of the thoracic vertebrae. In all vertebrae of the thoracic series the spinous processes are well- developed. They increase in length to the third, and then become gradually shorter, although their surfaces are, on the whole, slightly increased in extent. The anterior ten are directed backward, the elev- enth is almost vertical, while the twelfth is directed forward, like those of the succeeding lumbar vertebrae. The lumbar vertebrae (vertebrae lumbales) are seven in number. They are large vertebrae, conspicuous for their extensive surfaces and processes for muscular attachment (Fig. 27, E, F). The transverse processes continue the general line of the ribs of the thoracic region. They are directed forward, as well as outward, and the tip of each is formed by a thin triangular plate (processus triangularis), which represents afused rib. At the posterior side of the base of each is a short, flattened projection, the accessory process (processus accessorius). The spinous process is especially well-developed, and is directed forward. The articular processes are rotated upward, so that their surfaces are directed 72 ‘ANATOMY OF THE RABBIT. more nearly toward, or away from, the median plane, instead of to the dorsal or ventral surface. The anterior articular surfaces are borne on the bases of stout, upwardly-directed mammillary processes (processus mamumillares). The latter are most characteristic of the lumbar verte- brae, but may be seen to arise in the posterior thoracic region as small elevations of the transverse processes. Each of the first three of the lumbar vertebrae bears a median ventral projection, the anterior spinous process (processus spinosus anterior), for the attachment of the lumbar portion of the diaphragm. The sacral vertebrae (vertebrae sacrales) are four in number. In contrast to the true vertebrae—those united by ligament and articular surfaces—of the remaining portions of the vertebral column, they are false vertebrae, united in the young by synchondroses, and in the adult coalesced to form a composite structure, the os sacrum (Fig. 28). The A B Fic. 28 The ossacrum: A, ventral (pelvic) surface; B, dorsal surface; c.v., bodies of coalesced vertebrae; f.a., auricular surface; f.s.a., anterior sacral foramina; f.s.m., median sacral foramina; f.s.p., posterior sacral foramina; p.a.s., superior articular process of first vertebra; p.m., mam- millary process of first vertebra; pr. promontory; p.s., spinous processes. axis of the sacrum forms an obtuse angle with that of the lumbar ver- tebrae, the angle being indicated by a ventral projection, the promon- tory (promontorium), formed by the last lumbar and first sacral ver-. tebrae. The sacrum is the medium through which the vertebral column —in other words, the posterior portion of the trunk—is supported on the posterior limbs. Its anterior dorsal portion bears on either side a roughened area, the auricular surface (facies auricularis), for articulation with the pelvic girdle. This surface is borne for the most part on the transverse process of the first sacral vertebra. THE RIss. 73 The sacrum exhibits many features resulting from its formation through the fusion of originally distinct vertebrae. On the ventral, or pelvic surface (facies pelvina), the lines of junction may be traced either between the bodies, or between the transverse processes. Four pairs of apertures on this surface, the anterior sacral foramina (foramina sacralia anteriora), lead into the intervertebral foramina, and give passage to the sacral spinal nerves. On the dorsal surface (facies dorsalis) a pair of posterior sacral foramina lie in the line of junction of the first and second vertebrae. The spinous processes are evident in all four vertebrae. The combined articular and mammillary processes are conspicuous only in the first two, but are represented in the succeeding two by low, roughened tubercles. In the middle line dorsally the vertebral arches are separated by con- spicuous apertures, the median sacral fora-’ mina. The caudal vertebrae (vertebrae caudales) are sixteen in number. They are segments of small size, increasing slightly to the third, and then gradually decreasing to the end of the column. The arches are complete in the first seven. The transverse processes are vestigial in all except the third. At the end of the column the segments are reduced to slender cylinders of bone representing the vertebral bodies. EE RUBS: The ribs (costae) are twelve in number on either side. Each is composed of a dorsal portion, the costal bone (os costale), or bone-rib, and a ventral portion, the costal cartilage (cartilago costalis) (Fig. 29). From their attachment on the vertebral column the bone-ribs are directed outward, downward, and backward. The costal cartilages are directed for the most part inward, down- ward, and forward. The first costal cartilage forms a pronounced angle with the corres- _ Fic. 29. The sternum and first ponding bone-rib. In the succeeding ribs i>. ventral view: 1-7, the true ¢ ribs; 8, first false rib; c.c., head the angle is gradually replaced by a broad of rib;’ cl.c., neck of rib! cr., S costal cartilage; cr.c., body of curve. trib; ¢.s., body of sternum; m.s., Ribs are classified as true ribs (costae manubrium sterni; 0.c., bone-rib; : z p.x., xiphoid process; t.c., costal verae), and false ribs (costae spuriae). The _ tubercle. former—comprising the anterior seven— are those directly attached to the sternum. The latter—comprising the posterior five—are either indirectly attached, or unattached. The unattached ribs are designated as floating ribs. Generally speaking, the bone-ribs are cylindrical; but the anterior 74 ANATOMY: OF THE RABBIT. five or six are more or less flattened, with their main surfaces respectively medial and lateral. The compression is most marked in old animals. The first rib is extremely short. The succeeding ribs increase in length to the sixth, and then decrease to the twelfth. The arch formed by each rib has its greatest convexity, or angle, at some point toward the dorsal surface. Passing backward, the point of greatest convexity changes from a medial to a lateral position. This, together with the elongation of the more posterior ribs, results in an enormous increase in the posterior extent of the thoracic cavity. The vertebral end, or head of the rib (capitulum costae), is articulated with the body of the vertebra to which it belongs, and also, in the case of the first ten, with the vertebra immediately in front. The articulation with a transverse process is marked by a small smooth elevation, the costal tubercle (tuberculum costae). It is present only in the first nine ribs. Except in the first rib, and in the last four, the tubercle bears a sharp, dorsally-directed process for muscular attachment. The slender portion of the rib intervening between the head and tubercle is the neck (collum costae), the remaining larger portion being distinguished as the body of the rib (corpus costae). The bony thorax is formed by the ribs and the sternum with the assistance of the thoracic vertebrae. It encloses a large space, the thoracic cavity (cavum thoracis). The latter is conical in shape, with the apex directed forward. The dorsoventral diameter of the cavity is considerably greater than the transverse diameter. Apart from the intercostal spaces, the cavity is open at two points: anteriorly, the first thoracic vertebra, the first rib, and the manubrium sterni together enclose a small opening, the superior thoracic aperture (apertura thoracis superior); posteriorly, the seventh and succeeding ribs, together with the posterior thoracic vertebrae and the xiphoid process of the sternum, enclose a much larger opening, the inferior thoracic aperture (apertura thoracis inferior). In the natural condition it is largely closed by the diaphragm. The curved boundary formed by the ribs in this region is the costal arch (arcus costarum), the angle formed at the point of attachment of the xiphoid process being the infrasternal angle (angulus infrasternalis). Through their articulations with the vertebral column, and the nature of the costal cartilages, the ribs are capable of being moved, or rotated, forward. The movement results in an increase of the extent of the thoracic cavity, and is of importance in respiration. THE STERNUM. The sternum consists of a linear series of six segments, the sternebrae. The first segment is the manubrium sterni. It is about twice the length of the middle segments. It is somewhat triangular in section, two of its surfaces being ventrolateral, the third dorsal and directed toward the thoracic cavity. To its anterior tip is attached the sternoclavicular ligament, by which the greatly reduced clavicle is united with the sternum. THE SKELETON OF THE HEAD. 75 The four middle segments are similar in appearance, and form the body (corpus sterni). The sixth segment, described as the xiphoid process (processus xiphoideus), is an elongated strip of bone, to the posterior end of which is attached a broad, thin plate of cartilage. The first costal articulation is situated at about the middle of the manubrium, the remaining six at the points of junction of the segments. Five of them occur singly, while the sixth and seventh costal cartilages are attached together at the point of junction of the last segment of the corpus sterni with the xiphoid process. DEBS SKI EON OE Et Rein AWD: The head-skeleton comprises: (1) the series of elements constituting the skull; and (2) the hyoid bone, with its connections. The skull, or cranium—using that term in a general sense—includes the cranium proper, that portion enclosing the brain and containing in its wall the auditory capsules, and the bones of the face (ossa faciei), the latter in- cluding the series of elements related for the most part to the jaws and palate. The primary relations of the constituents of the head-skeleton have already been indicated above (p. 31). Ame Te SKU No Au Wl Ole rs, The skull is a composite structure, consisting of a large number of elements, which, with the exception of the mandible, are united by synarthroses, so that they produce the effect of a continuous mass. The mandible is a more or less independent structure, articulated with the main body of the skull by a typical joint. The skull is roughly divisible into two portions, namely, an anterior, facial portion, and a posterior, cranial portion. The cranial portion has a somewhat conical shape, its apex being directed forward. It is sepa- rated from the facial portion by a depression on either side of the skull, the orbital cavity (orbita), which serves for the accommodation of the optic bulb. Unlike the remaining special sense-organs, the eye is not included within the skull-wall. The two portions are united both medially and laterally, the lateral connection being established by the zygomatic arch (arcus zygomaticus), which bridges the lateral portion of the orbit. The facial portion has also a somewhat conical shape, its apex being formed by the anterior extremity of the upper jaw and the incisor teeth. Its base is formed in part by the connection with the cranial portion, as already described, and also by the anterior walls of the orbits. The cranial portion exhibits an extensive posterior, nuchal surface (planum nuchale), situated in general at right angles to the cervical portion of the vertebral column and also to the dorsal, lateral, and ventral walls of the skull. This surface includes the external surface of the Occipital bone, with the exception, chiefly, of the basilar portion of the latter. Its dorsal portion forms an area of attachment for the spinal and special occipital musculature. Its ventral portion is perforated by a 76 ANATOMY OF THE RABBIT. large aperture, the foramen magnum occipitale, for the passage of the central nervous system from the cranial cavity into the vertebral canal. On either side of this is a smooth, ridge-like projection, the occipital condyle (condylus occipitalis), for articulation with the superior articular pits of the atlas. At a little distance lateral to the occipital condyle, the nuchal surface is continued downward through the medium of a some- what triangular, pointed jugular process (processus jugularis). This structure is separated from the occipital condyle by a pronounced notch, the posterior boundary of a deep narrow excavation, the jugular fossa (fossa jugularis), which lies between the condyle and the tympanic bulla. The jugular process serves for the attachment of muscles belonging to the tongue, hyoid, and mandible, namely, the styloglossus, stylohyoidei major and minor, and the digastricus, the suspensory ligament of the lesser cornu of the hyoid also being included in the ligament of the stylohyoideus minor. Toward the dorsal margin of the nuchal surface, the dorsal surface of the skull is projected backward as a shield- shaped psa: Fic. 30. Lateral surface of the skull: AS, alisphenoid (ala magna); BO, basioccipital (basilar portion of occipital); BS, basisphenoid (body of posterior sphenoid); F, frontal; I, interparietal; L, lacrimal; M, maxilla; MS, mastoid portion of petrosal (petromastoid); N, nasal; OS, orbitosphenoid (ala parva); P, parietal; PL, palatine; PM, premaxilla; SO, supraoccipital (squamous portion of occipital); SQ, squamosal; T, tympanic; ZY, zygomatic. a.p., piriform aperture of nose; d.i., incisor teeth; d.m.,.molars; d.pm , premclars; f.i., infraorbital foramen; f.mx., maxillary fossa; f.o., optic foramen; f.s., stylomastoid foramen; f.t., temporal fossa; 1.]., lateral lamina of pterygoid process; 1l.m., medial lamina; m.a.e., osseous portion of external acoustic meatus; p.a., alveolar process of maxilla; p.e., ethmoidal portion of orbitosphenoid; p.-f., frontal process of premaxilla; p.j., jugular process of occipital; p.m., mastoid process of mastoid; p.mx., maxillary process of frontal; p.o., orbital process of maxilla; p.o.e., external occipital protuber- ance; p.S., squamosal process of parietal; p.s.a., and p.s.p., anterior and posterior supraorbital processes of frontal; p.z., zygomatic process of squamosal; p.z.m., zygo- matic process of maxilla; s, sphenoorbital process of maxilla; s.m., spina masseterica; sq., Squamosal process of squamosal. promontory. The lateral margin of this projection is the superior nuchal line (linea nuchae superior). It forms a curved ridge, the position of which indicates the dorsal limit of the occipital musculature. The posterior, somewhat tri-radiate tip of the projection is the external occipital protuberance (protuberantia occipitalis externa), an important ee THE SKULL AS A WHOLE. WG median point of attachment for the occipital muscles and the ligamentum nuchae. The ventral wall of the cranial portion is the basal portion (basi- cranium) of the entire skull. As indicated above (p. 31), its axial line, the basicranial axis, continues, in general, that of the bodies of the vertebrae, and its posterior portion is equivalent, morphologically, to vertebral segments. It is formed by a linear series of three bones, namely, the basilar portion of the occipital, the body of the posterior sphenoid, and that of the anterior sphenoid (respectively basioccipital, basisphenoid and presphenoid bones). Its extremely narrow, anterior portion forms the roof of a deep groove which encloses the nasal portion of the pharynx. As viewed from the ventral surface, it is seen to dis- appear in the facial complex at some distance dorsal to the posterior margin of the bony palatine bridge. Laterally, it is separated from the orbit on either side by a vertical plate formed by the palatine bone, and also by two downward projections of the posterior sphenoid, the medial and lateral laminae of the pterygoid process (processus pterygoideus). These structures enclose between them the pterygoid fossa (fossa pterygoidea), the walls of which serve for the attachment of the external and internal pterygoid muscles of the mandible. The lateral wall of the cranial portion forms anteriorly a large part of the boundary of the orbit. The cranial wall of the orbit is partly formed by two upward projections of the basicranium, namely, the lesser wing of the anterior sphenoid, or orbitosphenoid, and the greater wing of the posterior sphenoid, or alisphenoid. The remaining portion is formed by membrane elements, including the frontal bone of the roof of the skull and the squamosal bone, the latter distinguishable as the support of the posterior root of the zygomatic arch. Immediately behind the orbit, the root of the zygomatic arch projects outward and then downward. It is formed by a zygomatic process (processus zygomaticus) of the squamosal bone, and the tip of this forms a vertical plate, which is united by a horizontal suture with the zygomatic bone. On the ventral side of the process, close to the cranial wall, is the mandibular fossa (fossa mandibularis), for articulation with the mandible. On its dorsal side, but more especially on the adjacent portion of the cranial wall, there isa shallow, horizontal groove, lodging in the natural condition the temporalis muscle of the mandible, and therefore representing a greatly reduced temporal fossa (fossa temporalis). In the natural condition the anterior portion of the groove is converted into a foramen through the presence of a stout ligament extending from the posterior supraorbital process to the base of the zygomatic arch. The dorsal boundary of the fossa is formed by a pronounced ridge, the temporal line (linea temporalis), the latter forming also the lateral margin of the roof of the skull in this region. Behind the posterior root of the zygomatic arch, the external surface of the lateral wall is largely occupied by the swollen tympanic bulla (bulla tympani), formed by the tympanic bone. It contains the capac- ious tympanic cavity (cavum tympanicum) and certain skeletal structures of the middle ear, namely, the auditory ossicles (ossicula auditus), the 78 ANATOMY OF THE RABBIT. relations of which are more fully dealt with below (p. 90). The dorsal portion of the tympanic bulla is continuous with a short bony tube which opens at a short distance dorsad by a large oval aperture. This tube is part of a more extensive canal, the external acoustic meatus (meatus acusticus externus) which, in the natural condition, leads down- ward through the base of the external ear to the tympanic membrane. The tympanic bulla does not form the lateral wall of the skull in this region, and is not exposed to the cranial cavity. It is applied closely to the external surface of the periotic or petromastoid bone, which forms the lateral boundary of the cranial cavity, and contains the structures of the internal ear. The external or mastoid portion of this bone appears in the space enclosed between the tympanic bulla and the jugular process of the occi- pital bone, where it is readily distinguishable by its pitted appearance. Its ventral portion bears a slender projection, lying parallel to the jugular process, the mastoid process (processus mastoideus). A series of foramina, lying partly within the orbit, and extending thence posteriorly along the boundary between the lateral and ventral walls to the occiput, put the cranial cavity in communication with the outside, and serve for the passage of nerves and vessels. The first and largest of these, the optic foramen (foramen opticum), occupies the middle portion of the orbit, and trans- mits, in the natural condition, the optic nerve. Following this Fic. 31. Dorsal surface of the skull: F, frontal; , Z : ¥ I, interparietal; L, lacrimal; M. maxilla; .MS, is a vertical slit-like aperture— mastoid, portion of petrosal (petromastoid); N, : is if nasal; P, parietal; SO, supraoccipital (squamous not to be confused with the portion of occipital); SQ, squamosal; ZY, perforations of the external AION alae 1 ‘ f hie eran ‘4 sas f.mx., maxillary fossa; f.t., temporal fossa; amina of the pterygoid process Iinis stags nuchal line; 1.t.,: temporal line; iy : : p.f., frontal process of premaxilla; panix., Fille superior orbital fissure maxillary process of frontal; p.o.e., external (fissura orbitalis superior). It occipital protuberance; ps4. and) ps2, i $ f 5 anterior and posterior supraorbital processes of represents both the superior frontal; p.sc., subcutaneous process of lacrimal : Ari 4 c : p.z., zygomatic process of squamosal; p.z.m., orbital fissure of the normal zygomatic process of maxilla; s.f., frontal spine. mammalian skull and_ the S.ml., Spill: masseterica: foramen rotundum, and provides for the passage outward of the third, fourth and sixth cranial nerves, together with the first and second divisions of the fifth. The lateral lamina of the pterygoid LES OKULE, Ass Al EOL: 79 process presents three foramina, of which the largest, anterior, and medial one, the anterior sphenoidal foramen, serves for the transmission of the internal maxillary artery, while the remaining two, the middle and posterior sphenoidal foramina, transmit muscular branches (masseterico- temporal and ptery egobuccinnator) of the mandibular nerve. On the medial side of the base of the medial lamina of the pterygoid process there is a shallow longitudinal groove, representing the pterygoid canal (canalis pterygoideus) of the human skull. Immediately in front of the ‘tympanic bulla, on the ventral surface of the skull, an irregular aperture, the foramen lacerum, leads directly into the cranial cavity. It is incompletely divided into two parts by a slender bony splints salt contains, in addition to the foramen lacerum, the foramen ovale of the typical mammalian skull, and serves to transmit the mandibular portion of the fifth nerve and the internal carotid artery. Looking into the aper- ture from the front, it is seen to communicate not only with the cranial cavity, but also with two apertures in the anterior portion of the auditory complex. One of these—that toward the middle line—is the internal carotid foramen (foramen caroticum internum). It is the an- terior end of a canal transmitting the internal carotid artery; the pos- terior end of this canal, the point at which the internal carotid artery enters the tympanic bone, or the external carotid foramen (foramen caroticum externum), being visible as a rounded aperture lying on the posteromedial surface of the tympanic bulla. The second, lateral aperture communicating with the foramen lacerum is that of the auditory (Eustachian) tube (tuba auditiva). It leads into the tympanic cavity, ‘and in the natural condition the tube places this cavity in com- munication with the nasal portion of the pharynx. Associated with the mastoid process 1s a small aperture, the stylomastoid foramen (foramen stylomastoideum), the external opening of the facial canal, which serves for the passage of the facial nerve. A slit-like aperture, the jugular foramen (foramen jugulare), lies in the jugular fossa, between the posterior ventral margin of the tympanic bulla and the occipital condyle. It transmits the first portion of the internal jugular vein from the trans- verse sinus of the dura mater, and also the ninth, tenth, and eleventh cranial nerves. Finally, immediately in front of the dorsal portion of the condyle, the occipital segment is perforated by several small apertures together representing the ‘hypoglossal canal (canalis hypoglossi), and serving for the transmission of the hypoglossal nerve. The roof of the cranial portion is largely formed by two pairs of thin membrane elements, the frontal and parietal bones. The former occupy a general position between the orbits, while the latter are interposed between the frontal bones and the occipital segment. A small portion of the roof is formed posteriorly, however, by a small lozenge-shaped element, the interparietal bone, and by the shield-shaped projection described above, which is part of the occipital bone. The space enclosed by the cranial portion of the skull is the cranial cavity (cavum cranii). Its form depends on the external configura- tion of the brain. It is divisible into three portions, known as the cranial fossae. The anterior cranial fossa (fossa cranii anterior) is a 80 ANATOMY OF THE RABBIT. small division lodging in the natural condition the olfactory bulbs of the brain. The middle cranial fossa, the largest division of the cavity, lodges the enlarged cerebral hemispheres. The posterior cranial fossa is a small division extending backward to the foramen magnum, and lodging in the natural condition the cerebellum and related posterior portions of the brain. It is partly set off from the middle cranial fossa by a fold of the dura mater, the tentorium cerebelli, which projects inward from the dorsal and lateral walls of the skull. This fold is usually found adhering to the internal surface of the skull, unless the latter has been very thoroughly cleared, and in all cases its posi- tion is indicated by a low ridge of bone. The marked difference in diameter between the middle and posterior cra- nial fossae is accounted for by the great thickness of the auditory portion of the skull. The anterior surface of the periotic bone will be observed to form an extensive posterior wall for the middle cranial fossa. The floor of the middle and posterior cranial fossae is not smooth, like the external base of the skull, but presents in its anterior portion a pro- minent elevation, the sella turcica, which is borne on the body of the posterior sphe- noid. It contains a large central depression, the hypo- physiai fossa (fossa hypophy- Seos), which in the natural Fic. 32. Ventral surface of the skull: AS, alisphenoid condition, ledges the hypo- Gee) ec (le physis or pituitary body. The EXO, exoccipital; M, maxilla; PL, palatine; PMX, = Siete _ premaxilla; PR, presphenoid (body of anterior sphe- aperture of the fossa 1s partly noid); rae supraoccipital (Squamous eee of occi- A A r y ; pital); SQ, squamosal; T, tympanic; ZY, zygomatic. enclosed laterally by a Ppalt ch, choana; c.hy., hypoglossal canal; c.o., occipital of pointed posterior clinoid condyle; f.c.e., external carotid foramen; f.in., incisive : : - foramen; f.j, jugular foramen; f.1., foramen lacerum,; processes (processus clinoidei f.m., mandibular fossa; f.m.o., foramen magnum; posteriores) the tips of which f.p.m., greater palatine foramen; f.s.a., anterior sphe- 2 noidal foramen; m.a.e., osseous portion of external are directed forward; and @ acoustic meatus; p.j., jugular process; DOs esnual we : : . occipital protuberance; p.pl., palatine process of maxilla; corre sponding pair of anterior p.pm., palatine process of premaxilla; p.pt., medial and clinoid processes lie at the lateral laminae of pterygoid process of posterior sphenoid; : 3 ¢ s.m., Spina masseterica. anterior end of the fossa, with the tips directed backward. The posterior, and also dorsal wall of the fossa, described as the dorsum sellae, leads by an abrupt curve Sine eagle «ai ee al ee ‘Tae SKULL AS A WHOLE, 81 backward on to the floor of the posterior cranial fossa, the sloping portion of the floor, or clivus, supporting in the natural condition the pons and medulla oblongata. Toward the anterior end of the middle cranial fossa, the lateral walls of the skull are greatly compressed, so that the anterior portion of the basicranium, especially the body of the anterior sphenoid, is largely excluded from the cranial cavity. The usually paired optic foramina are here confluent, there being a single aperture for the transmission of the optic nerves. The posterior ventral boundary of this aperture contains a broad groove, the sulcus chiasmatis, which lodges in the natural condition the optic chiasma. In the anterior cranial fossa the floor is largely formed by a perforated area, borne on the cribriform plate (lamina cribrosa) of the ethmoid bone, and serving for the transmission of the divided olfactory nerves. Its median portion projects slightly into the cranial fossa as a low ridge, the crista galli, which is interposed between the tips of the olfactory bulbs. In the ventrolateral portion of the cranial cavity may be found the internal openings of the foramina described above, namely, the superior orbital fissure, the foramen lacerum, the jugular foramen, and the hypo- glossal canal. The superior orbital fissure is almost ventral in position to the foramen opticum, and is connected backward with the foramen lacerum by a broad groove, the sulcus sphenoidalis, which lodges in the natural condition the roots of the fifth nerve. This groove continues to the medial surface of the periotic bone, where it is bridged over by the tentorium cerebelli. On the lateral wall of the posterior cranial fossa, and enclosed by the compact, white, petrous portion of the periotic bone, is a series of three apertures leading into its substance. One of these, much larger than the remaining two, is the subarcuate or floccular fossa (fossa subarcuata s. floccularis). It lodges in the natural condition the flocculus, a small stalked appendage of the cerebellum. Ventral to this fossa, and also somewhat in front of it, a thin ledge of bone extends over an oval open- ing, the internal aperture of the facial canal (canalis facialis), which serves for the transmission of the seventh cranial (facial) nerve. Imme- diately behind and below this aperture is the opening of the internal acoustic meatus (meatus acusticus internus) for the transmission of the eighth cranial (acoustic) nerve. The two apertures tend to be enclosed by a shallow bony ridge, largely formed by the projecting ledge described above, and resembling superficially the complete common tube repre- sented by the internal acoustic meatus of the human skull. The facial portion of the skull is largely formed by the investing bones of the upper jaw, palate, and mandible, but it encloses also the entire olfactory region of the primary skull, including the nasal fossae and associated turbinal bones. The upper jaw—the maxilla of the human skull—is formed of two primary, and, in the rabbit, separate, elements, the maxilla and premaxilla. They together form the greater portion of the facial region—in the adult condition also a large portion of the lateral walls of the nasal fossae—and bear in a ventrolateral position low alveolar processes (processus alveolares) for the sockets or alveoli of the incisor and cheek-teeth. The maxilla bears the anterior 82 ANATOMY OF THE Rappir. root of the zygomatic arch, the latter being formed partly by a short zygomatic process arising from its lateral surface, by the zygomatic bone, which is fused with it, and by the corresponding zygomatic process of the squamosal bone, constituting the posterior root. The anterior root of the zygomatic arch is perforated by a deep narrow infraorbital canal (canalis infraorbitalis), which opens on the facial surface by a vertical slit-like aperture, the infraorbital foramen. It serves for the trans- mission of the infraorbital vessels and nerves from the orbit to the face. The ventral portion of the maxilla is associated with the palatine bone to form the hard palate (palatum durum). This structure is repre- sented chiefly by a bony palatine bridge connecting the two sides of the skull between the more anterior cheek-teeth. It forms a portion of the roof of the oral cavity and a portion of the floor of the nasal cavity. Immediately in front of it, the palatal surface is perforated by a pair of large incisive foramina (foramina incisiva), which are broadly open to the nasal fossae. A considerable portion of the anterior and dorsal wall of the orbit is formed by the facial complex. Dorsally, the roofing element of this region, the frontal bone, bears.a curved lateral projection, the supra- orbital process (processus supraorbitalis), which overhangs the orbit. Its narrower base expands into anterior and posterior tips, which lie parallel to the adjacent portion of the skull, and enclose with the latter corresponding anterior and posterior supraorbital incisures. The latter are converted by ligament into foramina. The anterior wall of the orbit is formed in part by a loosely articulated element, the lacrimal bone, the lateral margin of which projects from the orbital rim as a blunt sub- cutaneous process (processus subcutaneus). On the ventral side of its base is the orbital opening of the nasolacrimal canal (canalis nasola- crimalis), the bony enclosure of the nasolacrimal duct, which in the natural condition leads from the corneal surface of the eye to the anterior portion of the nasal fossa. A smaller projection forming the ventral boundary of the nasolacrimal aperture is the hamulus lacri- malis. Finally, in the ventral anterior angle of the orbit, the bases of the three posterior cheek-teeth encroach to a considerable extent on the orbital space. They are separated from the orbital wall by a deep infraorbital groove (sulcus infraorbitalis), which leads forward into the canal of the same name. They partly conceal two important apertures of this region, the orbital opening of the pterygopalatine canal (canalis pterygopalatinus), leading to the palatal surface, and the sphenopalatine foramen (foramen sphenopalatinum), leading to the nasal fossa. The pterygopalatine canal opens ventrally in the palato-maxillary suture of the hard palate by a rounded aperture, the greater palatine foramen (foramen palatinum majus). The nasal cavity (cavum nasi) is enclosed by the maxilla and pre- maxilla, with the assistance of paired roofing elements, the nasal bones. Apart from the incisive foramina, which are closed in the natural con- dition, the cavity is open at two points. Posteriorly it communicates with the ventral surface of the skull by the choanae, which, in the rabbit, are incompletely divided. Anteriorly it opens to the outside by the THE SKULL AS A WHOLE: 83 piriform aperture (apertura piriformis). The cavity is divided into right and left portions, the nasal fossae. In the divided skull it is seen that the division is effected chiefly through a median vertical, cartilaginous plate, the nasal septum (septum nasi), or cartilaginous portion of the mesethmoid. This is continuous posteriorly with a small crescentic vertical plate of bone, the perpendicular plate (lamina perpendicularis) of the ethmoid bone—the bony portion of the mesethmoid—and the latter is also the terminal element of the series of median bones constituting the basicranium. Posteriorly, the ventral portion of the cartilaginous nasal septum is. supported by a vertical bony plate, the vomer, the dorsal margin of which is grooved to receive it. Anteriorly, the nasal septum bears on its ventral margin the paired enclosures of the vomero- nasal organ, which are also supported by the grooved surface formed in the middle line by the adjacent dorsal surfaces of the palatine processes of the premaxilla. The relations of these structures, as well as of the cartilage supporting the eso] ge eae duct, are best seen in very young animals (cf. Plate III). The delicate, foided, or scroll-like turbinated bones, characteristic of the nasal cavity, are borne on its posterior and lateral walls. Occupying Fic. 33. The skull in vertical section: BO, basioccipital (basilar portion of occipital): BS, basisphenoid (body of posterior sphenoid); ET, ethmoturbinal; F, frontal; I, inter- parietal; M, maxilla; MT, maxilloturbinal; N, nasal; NT, nasoturbinal; P, parietal; PE: palatine: PMX, premaxilla; PR, presphenoid (body of anterior sphenoid); PT, petrous portion of petromastoid; SO, supraoccipital (squamous portion of occipital); T, tympanic; V, vomer. a.p., piriform aperture of se c.f., internal aperture of facial canal; c.o., occipital condyle; f.c.a., f.c.m. and f.c »., anterior, middle, and posterior cranial fossae; f.f., floccular fossa; f.h., hy pophy sial fossa f.in., incisive foramen; f.s., sphenopalatine foramen; 1., per- pendicular plate of the ethmoid; m.a.i., internal acoustic meatus; 0., optic foramen; p.a., alveolar process of maxilla; p.d., hard palate; p.o.e., external occipital protuberance; p.pt., pterygoid process of posterior sphenoid; s.n., nasal septum; t.c., tentorium cerebelli. the anterior portion of the lateral wall of the nasal fossa is a finely-ridged mass of bone, the concha inferior, or maxilloturbinal. It is easily dis- tinguishable from a more dorsal and posterior series of broader folds, which together constitute the ethmoturbinal. In the rabbit,as in mammals generally, the latter is divisible into a more dorsal elongated portion attached to the nasal bone, the nasoturbinal, and a more ventral portion, also posterior portion, the ethmoturbinal proper, composed of several 8&4 ANATOMY OF THE RABBIT. shorter folds decreasing in length from above downward. In the natural condition the turbinated bones bear a considerable portion of the nasal epithelium, the surface of which is greatly increased by the folding of the underlying bone. That covering the ethmoturbinal contains the olfactory sense organs, while that covering the maxilloturbinal is non- sensory and possesses the mechanical function of freeing the air of the respiratory tracts from foreign materials, as well as of warming it slightly in its passage. On this account the respective structures are conveniently distinguished as sensory (olfactory) turbinals and respiratory turbinals. The mandible (mandibula) is composed of two portions, united anteriorly by the symphysis mandibulae. Each half comprises a hori- zontal portion, forming in conjunction with that of the opposite side the body of the mandible (corpus mandibulae), and a posterior, vertical portion, the ramus mandibulae, the latter serving for the insertion of the muscles of mastication and for artticulation with the skull. The Fic. 34. Lateral surface of the left ramus of the mandible: a.m., angle; c.m., body of mandible; cp.m., articular portion (head) of mandible; d.i., d.m., and d.pm., incisor, molar and premolar teeth; f.m., mental foramen; i.m.a.and i.m.p., anterior and posterior mandibular incisures; p.c., coronoid process; p.cd., condyloid process; t.m. and t.pt., masseteric and pterygoid tuberosities. body of the mandible bears on its dorsal margin the alveoli of the lower teeth. The mandibular ramus forms a broad plate, the lateral surface of which is occupied in the natural condition by the masseter muscle, while the medial surface forms an area of insertion for the external and internal pterygoids. The surface of the ramus is greatly increased in its posteroventral portion through the expansion of the bone to form the angle (angulus mandibulae), or angular apophysis. The elon- gated articular surface is borne at the end of a vertical, or slightly oblique condyloid process (processus condyloideus). The nerve and vessels of the mandible enter at the mandibular foramen (foramen mandibulare), the latter being situated on the medial surface of the bone immediately behind the last cheek-tooth. THE BONES OF THE SKULL. 85 Pa EE BONES OF THE SKULE: iH OCCIPITAL, BONE, The occipital bone (os occipitale) is the first of the basicranial seg- ments as numbered from the occipital articulation forward. It forms the posterior boundary of the skull, and establishes the connection of the latter with the vertebral column. Its external surface is identifiable for the most part with the nuchal surface, but a portion of it falls in the horizontal plane of the basis cranii. The internal surface is partly exposed to the cranial cavity, and forms in this relation the posterior, dorsal, and ventral boundaries of the posterior cranial fossa. The re- maining portion is excluded from the cranial cavity, being applied instead to the broad posterior surfaces of the petrotympanic bones. The occipital bone is divisible into four portions, namely, the basilar portion (pars basilaris), or basioccipital, the paired lateral portions (partes laterales), or exoccipitals, and the squamous portion (squama occipitalis), or supraoccipital. All four portions take part in the forma- tion of the foramen magnum. In the young animal (Fig. 11) they are represented by separate elements, formed in a continuous mass of cartilage, and united for a time by synchondroses, but in the course of growth they become fused to form a single occipital bone. The basioccipital is that portion lying below and in front of the foramen magnum. Its main surfaces are respectively dorsal and ven- tral. Its anterior margin is united with the posterior margin of the basisphenoid by a thin, transverse cartilage union, the sphenooccipital synchondrosis (synchondrosis sphenooccipitalis). _Posteriorly its dorsal and ventral surfaces come together in a thin concave edge which forms the ventral boundary of the foramen magnum. Laterally it is bounded by the petrotympanic bone and by the lateral portion of the occipital. The dorsal surface bears a median groove, deeper in its middle portion, where the lateral margins of the bone are raised to form a pair of rounded bosses for articulation with the petrotympanic. The groove represents the sloping portion or clivus of the occipital, and lodges in the natural condition, as described above, the ventral portion of the medulla oblongata. The ventral surface presents a similar groove, in the pos- terior portion of which there is a small ridge-like elevation, the pharyngeal tubercle (tuberculum pharyngeum). The exoccipital is directed dorsad from the basioccipital in such a way that it falls in the plane of the nuchal surface. It is applied to the posterior surface of the petrotympanic bone, and also extends down- ward beyond the latter as the jugular process. The occipital condyle is borne on the exoccipital, with the exception, however, of its ventral tip, which belongs to the basioccipital. The portion of the occipital bone con- necting the basioccipital and exoccipital contains the jugular fossa and the apertures representing the hypoglossal canal. Its anterior margin bears a jugular incisure (incisura jugularis), forming the occipital 8 86 ANATOMY OF THE RABBIT. boundary of the jugular foramen, the remaining portion of the latter being formed by the petrotympanice. The supraoccipital is the dorsal portion of the bone. Its dorsal margin is bent sharply forward, so that it tends to fall, like the basi- occipital, in a horizontal plane. Its external surface bears the superior nuchal lines and the external occipital protuberance. A pair of lateral wing-like expansions rest upon, and partly overlap, the dorsal margins of the petrotympanic bones. The anterior boundary is formed by the interparietal, parietal, and squamosal bones, but in young skulls the squamosal connection is represented by a vacuity. The internal sur- face bears a longitudinal groove, lodging in the natural condition the median vermis of the cerebellum. It is crossed at its anterior end by a shallow transverse groove (sulcus transversus), which marks the position of the transverse sinus of the dura mater. 2. DHE POSTERIOR: SPHENOLD: The sphenoid bone, as identified from the human condition, is a com- plex of elements belonging to two segments, namely, the posterior sphenoid (os sphenoidale posterius) and the anterior sphenoid (os sphenoidale anterius). In the rabbit, as in mammals generally, these - segments are separate throughout life. The posterior sphenoid comprises: (1) a median portion, the body, or basisphenoid; (2) paired dorsolateral expansions, the greater wings (alae magnae), or alisphenoids; and (3) paired ventral projections, the pterygoid processes. The basisphenoid continues the basis crani forward from the basi- occipital to the body of the anterior sphenoid. It is united with the latter by the intersphenoidai synchondrosis. Its surfaces correspond for the most part to those of the basioccipital. The ventral surface forms the chief part of the bony roof of the nasopharynx. It is perforated in its middle by a round aperture, the foramen cavernosum, which leads into the interior of the bone. The dorsal surface is occupied, as described above, by the hypophysial fossa and related structures, namely, the dorsum sellae and the posterior clinoid processes. On the lateral surface of the base of the posterior clinoid process a faint groove, the sulcus caroticus, marks the course of the internal carotid artery. The interior of the bone contains a cavity of considerable size, the sphenoidal sinus (sinus sphenoidalis), which communicates both with the foramen cavernosum and the hypophysial fossa. The alisphenoid extends at first laterad, but soon changes its direction so that its axis becomes dorsoventral. At the same time the bone is rotated in such a way that its surfaces tend to fall in a transverse plane. It is bounded anteriorly by the orbitosphenoid, dorsally by the squamosal, and posteriorly by the petrotympanic. The anterior margin of its root encloses with the basisphenoid, and to a certain extent with the orbito- sphenoid, the superior orbital fissure. The foramen lacerum is formed by the posterior margin of its root in association with the petrotympanice. The external surface of the alisphenoid is convex, both toward the THE BONES OF THE SKULL. 87 orbit and toward the ventral surface of the skull. In the posterior por- tion of the orbit this surface bears a jagged elevation, the crista alae magnae. The internal surface forms a portion of the floor and antero- ventral wall of the middle cranial fossa. At its base a broad groove, the sulcus sphenoidalis, indicates the position of the root of the fifth nerve and the related semilunar (Gasserian) ganglion. The pterygoid process comprises the two plates described above as the medial and lateral laminae. The former is vertical, and its medial surface is directed toward the nasopharynx. The latter is almost hori- zontal. The medial lamina ends ventrally in a hooked projection, the hamular process (hamulus pterygoideus). In the young animal this por- tion is formed of an elevation of cartilage tipped by a separate mem- brane element, the pterygoid bone. The pterygoid fossa is formed in part by the medial and lateral laminae and in part by the divided pos- terior end of the palatine bone. The posterior base of the lateral lamina is extensively excavated, like the adjacent portions of the alisphenoid. It bears a shallow groove, representing a pterygoid canal (canalis pterygoideus), and is perforated by the three apertures described above as the anterior, middle, and posterior sphenoidal foramina. Set ANTERIOR oPiBeNOL: The anterior sphenoid (os sphenoidale anterius) consists of two por- tions, namely, a median portion, the body, or presphenoid, and a pair of lateral expansions, the lesser wings (alae parvae), or orbitosphenoids. The presphenoid is a constricted bony splint which continues the basis cranii forward from the basisphenoid. It is joined anteriorly with the perpendicular plate of the ethmoid and with the cartilaginous nasal septum. In the divided skull, or better in one from which the roof has been removed, the actual dorsal surface of the bone is seen to be exposed to the cranial cavity only in its posterior portion, where it is occupied by the sulcus chiasmatis and the optic foramina. That part of the floor immediately in front of the optic foramina is formed by the coalesced roots of the orbitosphenoids, the dorsal surface of the pre- sphenoid being thus excluded. The orbitosphenoid forms a long, low plate, lying in the ventral por- tion of the orbit, and divided by a shallow notch at the level of the optic foramen into a posterior portion, the orbitosphenoid proper, and an anterior portion, the ethmoidal process (processus ethmoidalis). The orbitosphenoid proper lies behind the optic foramen. It is in contact dorsally with the orbital portion of the frontal, and ventrally with the alisphenoid; it assists the latter in the formation of the superior orbital fissure. Its posterior tip is in contact with the squamosal. Its internal surface forms a considerable portion of the anteroventral wall of the middle cranial fossa. The ethmoidal process extends forward from the optic foramen. Its dorsal margin is articulated with the orbital portion of the frontal, and its ventral margin with the orbital portion of the palatine. Anteriorly it projects toward the lacrimal bone, thus occupying, in part, a space 88 ANATOMY OF THE RABBIT. which, in the typical mammalian skull, is filled by the lamina papyracea of the ethmoid. Its internal surface is associated with the ethmoid bone and with the nasal cavity. It falls for the most part below the level of the cranial cavity. 4. THE SQUAMOSAL BONE. The temporal bone, or temporal complex, as recognized from the human condition, 1s an association of three elements—squamosal, tympanic, and periotic—which in the human skull are coalesced to form a single bone. It is usually described as consisting of four portions, of which the squamosal and tympanic portions are two, while the periotic bone is considered to consist of two others, one of which, the petrous portion, is a solid white portion lodging the internal ear, while the second, or mastoid portion, is a mass of less compact character appear- ing externally in the wall of the skull. In the rabbit the original elements are not coalesced, but the periotic and tympanic bones are so closely associated that it is proper to describe them as forming a petrotympanic bone. The squamosal bone (os squamosum) is a rectangular plate, forming part of the lateral wall of the cranium, and bearing the posterior root of the zygomatic arch. It is articulated anteriorly with the orbitosphenoid and with the orbital portion of the frontal, dorsally with’ the frontal and parietal posteriorly with the supraoccipital and petrotympanic, and ventrally with the alisphenoid. Its posterior margin bears a prominent, slightly decurved squamosal process (processus squamosus). It hes on the lateral surface of the petrotympanic immediately above the opening of the bony external acoustic meatus. The posterior root of the zygo- matic arch is formed by a lateral and afterwards ventral projection, the zygomatic process of the squamosal. Its base bears ventrally the mandibular fossa, and dorsally, in association with the body of the squamosal, the temporal fossa. The internal surface of the squamosal forms a considerable portion of the wall of the cranial cavity, the middle cranial cavity being, in fact, broadest in this region. 5. DHE PRLROT YMPANIC. BONNIE: The petrotympanic bone (os petrotympanicum) is a somewhat oblong bone lying in the lateral wall of the cranium between the posterior sphenoid and occipital bones. It is chiefly indicated externally by the tympanic bulla and the bony external acoustic meatus. It is articulated anteriorly with the alisphenoid and squamosal, dorsally with the supra- occipital, and posteriorly with the exoccipital. Except for the presence of the squamosal process of the squamosal bone, the lateral and ventral surfaces are exposed to the outside of the skull. The internal surface is exposed to the posterior cranial fossa, with the exception, however, of a small ventral portion which is articulated with the basioccipital bone. Only a small portion of the anterior surface is in contact with the squamosal bone, the larger part being applied to the tentorium cerebelli and forming with the latter a posterior wall for the middle THE BoNnES OF THE SKULL. 89 cranial fossa. The dorsal portion of the bone corresponds in thickness with the wing of the supraoccipital with which it is articulated. The posterior surface is applied to the anterior surface of the exoccipital, and is thus excluded both from the cranial cavity and from the external surface of the skull. Viewing the skull from behind, however, it is seen that a small dorsal portion protrudes in a triangular space formed by the dorsolateral margin of the exoccipital and the ventrolateral margin of the supraoccipital wing. This portion is distinguishable by its pitted character. It forms the mastoid portion (pars mastoidea) as distin- guished from the solid white petrous portion (pars petrosa), which is exposed to the cranial cavity, and which contains the structures of the internal ear. The mastoid portion lies for the most part above the tympanic cavity, but it is also continued ventrad between the external acoustic meatus and the exoccipital as the mastoid process. The stylo- mastoid foramen lies between the latter and the external acoustic meatus. The petrous portion, as viewed from its medial surface, is roughly oblong; it is placed obliquely with reference to the basioccipital and basisphenoid. The floccular fossa occupies its posterodorsal portion, and extends into the substance of the bone, forming a much larger depression than is indicated by the diameter of its rim. The related dorsal margin of the bone is occupied by a groove which leads into a canal at its posterior margin. It indicates the position of the lateral portion of the transverse sinus of the dura mater. The ventral, thicker portion of the bone, enclosing the apertures of the internal acoustic ' meatus and the facial canal, is also that lodging the vestibulum and cochlea of the internal ear. A small aperture at its anteroventral angle, only visible when the petrotympanic is freed from its connections, repre- sents the hiatus canalis facialis of the human skull. It transmits the great superficial petrosal nerve, a branch of the facial nerve passing to the sphenopalatine ganglion. The tympanic surface of the petrous portion is described below in connection with the structures of the tympanic cavity. The tympanic portion forms the spherical, expanded, shell-like, tym- panic bulla, which contains in its interior the tympanic cavity, and is continuous dorsally with the bony enclosure of the external acoustic meatus. The boundary between the two is indicated externally by a shallow oblique groove, the position of which indicates roughly that of the tympanic membrane within. The medioventral margin of the bone -is articulated with the basioccipital, but the swollen portion is separated from the latter by a broad groove terminating posteriorly in the jugular fossa and the jugular foramen. Immediately in front of the jugular fossa, the rounded aperture of the external carotid foramen, transmitting in the natural condition the internal carotid artery, leads into the carotid canal of the interior of tympanic portion. At the anterior end of the groove, communicating with the foramen lacerum, is the anterior opening of the carotid canal, the internal carotid foramen, and on its lateral side the much larger aperture of the auditory (Eustachian) tube. The rela- tions of these apertures are seen to best advantage when the petro- 90 ANATOMY OF THE RABBIT. tympanic is disarticulated from the associated posterior sphenoid bone. The auditory tube is then seen to lead directly into the tympanic cavity. A fine bristle may be passed through the carotid canal from one foramen to the other. 6. THE STRUCTURES ORME DY MiPANIC (CAVAR Ve The relations of the tympanic cavity and associated structures may be studied with advantage in a skull from which the lateral wall of the tympanic bulla and external acoustic meatus has been removed, the sur- face displayed being as indicated in Fig. 35. The tympanum or middle ear is enclosed by the tympanic and petromastoid portions of the temporal complex. The attached margin of the tympanic bulla encloses a roughly triangular area, into the ventral part of which the petrous portion of the petromastoid projects as a smooth, white, convex ridge, the prom- ontory (promontorium). Above and behind the promontory the tympanic cavity is extended toward the mastoid portion of the bone Fic. 35. Petrotympanic portion of the auditory complex of the left side x3. The lateral portions of the tympanic bulla and external acoustic meatus have been removed, exposing the structures of the tympanic cavity. MS, mastoid portion; P, petrous portion; T, tympanic portion (bulla tympani). c.m., mastoid cells; c.t., tympanic cavity; f.c., cochlear fenestra; in., incus; m.a.e., external acoustic meatus; m.m., manubrium of the malleus; mz.so., supraoccipital margin of petromastoid; p.m., mastoid process; st., stapes; t.a., aperture of auditory tube. as the tympanic or mastoid antrum (antrum tympanicum), and the interior of the mastoid portion is partly occupied by small extensions of the tympanic antrum, termed the mastoid cells (cellulae mastoideae). At the anteroventral angle of the area already described, a deep notch indicates the point of entrance of the auditory tube. The exposed surface of the petromastoid presents two apertures, one of which, situated posteroventrally, is open in the dried skull, and is the cochlear fenestra (fenestra cochleae). In the natural condition it is closed by a thin membrane which separates the tympanic cavity from the perilym- phatic space containing the membranous labyrinth. The second aperture, the vestibular fenestra (fenestra vestibuli), lies above and in front of that just described. It is closed bv the base of the stapes. The auditory ossicles (ossicula auditus) comprise three elements, namely, the malleus, incus, and stapes, which bridge the space inter- THE BoNES OF THE SKULL, 9] vening between the tympanic membrane and the opening to the internal ear as represented by the vestibular fenestra. They occupy the dorsal angle of the triangular area already described, and lie immediately above the promontory. The malleus is the lateral element. The main por- tion. termed the head, is concealed by the projecting edge of the external acoustic meatus. It bears a stout vertical process, the manubrium mallei, which in the natural condition lies in contact with the tympanic membrane. The incus is the intermediate element; it is directly articulated with the malleus, and bears a downwardly-directed long limb (crus longum), for articulation with the minute head of the stapes. The latter element is a small, stirrup-shaped bone, occupying an almost transverse position, and articulated at its base with the margin of the vestibular fenestra. EE NOE Re PAR VE A BONE: The interparietal (os interparietale) is a small, lozenge-shaped element, surrounded by the two parietal bones and the supraoccipital. It is the first of the membrane roofing elements of the cranium proceeding for- ward from the supraoccipital, and in the rabbit’s skull is not fused with the occipital segment. 8. THE PARIETAL BONE. The parietal bone (os parietale) is a characteristic roofing bone covering a large portion of the middle cranial fossa. It is somewhat rectangular in shape, and is connected by serrate sutures with the sur- rounding elements and with its fellow of the opposite side, the sutures producing a characteristic pattern on the external surface of the skull. The sutures are medial, anterior, lateral, and posterior in position, and are designated respectively as sagittal, coronal, squamosal, and lamb- doidal. The posterolateral angle of the bone is produced ventrally into a long, curved squamous process (processus squamosus), which lies in the angle formed by the tentorium cerebelli and the lateral wall of the middle cranial fossa. It is not exposed to the external surface of the skull. Uo tHE TRONTAL BONE: The frontal bone (os frontale) is a paired element, lying directly in front of the parietal, and forming with its fellow of the opposite side the anterior portion of the roof of the cranial cavity and also a considerable portion of its lateral, orbital wall. Unlike the condition in the human skull, the two bones are separate throughout life, so that there is a per- manent frontal suture. Each consists of a frontal portion (pars frontalis), the external or dorsal surface of which continues that of the parietal, and of an orbital portion (pars orbitalis), enclosing the dorsal portion of the orbit. The two parts are connected at the supraorbital border, with which is also connected the base of the divided supraorbital process. The anterior end of the frontal portion is deeply notched where it comes in contact with the nasal and premaxillary bones. Two processes are 92 ANATOMY OF THE RABBIT. thus formed, one medial, the other lateral to the nasal. The medial process is associated with that of the opposite side to form a triangular frontal spine, while the lateral or maxillary process (processus maxillaris) projects forward between the nasal and premaxillary bones, on the one hand, and the subcutaneous process of the lacrimal, the orbital process of the maxilla, and the body of the latter, on the other. The orbital portion of the frontal forms a considerable portion of the orbital wall. Its anterior margin is in contact with the lacrimal bone, its ventral margin with the slender sphenoorbital process of the maxilla, the ethmoid process of the orbitosphenoid, and the orbitosphenoid proper. Its internal surface is divided by a vertical ridge into anterior and posterior portions, in relation respectively to the anterior and middle cranial fossae. The anterior cranial fossa is enclosed by the frontal bones, with the exception, however, of a small portion of the floor which is formed by the cribriform plate of the ethmoid. 10.. THESBLAMOlMD.BONE: The ethmoid bone (os ethmoidale), the chief representative of the embryonic cartilaginous nasal capsule, is a delicate, greatly sculptured structure, almost completely enclosed by the membrane bones of the face. Its features may be studied either in the divided skull, or in one from which the roof of the nasal and cranial cavities has been re- moved. It consists of three main portions, namely, the cribriform plate, the perpendicular plate, and the paired lateral masses, or ethmoidal labyrinths. The cribriform plate (lamina cribrosa) is exposed to the anterior cranial fossa. It is somewhat heart-shaped, with its apex in contact with the ethmoidal processes of the orbitosphenoids. Its lateral por- tions are perforated by numerous foramina, giving passage in the natural condition to the branches of the olfactory nerves. Its median portion forms a low vertical ridge, the crista galli, continuous in front with the perpendicular plate. The perpendicular plate (lamina perpendicularis) is the bony, pos- terior portion of the nasal septum, and as such is exposed to the nasal cavity. It is united with the cartilaginous nasal septum and also with the presphenoid. It forms the terminal member of the chain of bones lying in the basicranial axis. The ethmoidal labyrinth (labyrinthus ethmoidalis) occupies for the most part the posterior portion of the nasal fossa, but the nasoturbinal extends forward to its anterior end, and is attached for the greater part of its length to the internal surface of the nasal bone. It is broadest in its middle portion, where it projects into the space left between the ethmoturbinal proper and the maxilloturbinal, and contains at this point a pouch-like cavity, termed the marsupium nasale. The whole struc- ture is comparable to one of the folds of the ethmoturbinal proper; but it is frequently seen to be divided into anterior and posterior parts by a thin vertical line of cartilage, the anterior division being probably allied to the maxilloturbinal. Its middle, ventral, portion bears a —— > ane iat THE BONES OF THE SKULL. 93 stout, backwardly-directed uncinate process (processus uncinatus), which is applied to the medial surface of the maxilla. The ethmoturbinal proper consists, as described above, of several shorter scrolls, decreasing in length from above downward. Like the posterior part of the nasoturbinal, they are attached. directly to the cribriform plate, the perforations of which may be seen in the divided skull opening into the ethmoidal cells (cellulae ethmoidales) or spaces contained by them. They are roughly comparable to the superior and middle turbinated bones of the human skull, but in the rabbit, as in most mammals, the ethmoturbinal surfaces are relatively much more extensive than in man. In the typical mammalian skull the ethmoid bone is exposed to the orbit, where it forms a thin plate of bone, the lamina papyracea. In the rabbit, however, the space usually occupied by the lamina papyracea is partly filled by the lacrimal bone, the ethmoidal process of the orbito- sphenoid and the sphenoorbital process of the maxilla. 11. THE INFERIOR TURBINATED BONE. The inferior turbinated bone (concha nasalis inferior), or maxillo- turbinal, is a finely ridged structure, situated anteriorly in the nasal fossa, and supported by the maxilla and premaxilla. It represents the simi- larly-named structure of the human skull, the lowermost of three scroll- like bones, of which the remaining two, the superior and middle turbinated bones, belong to the ethmoturbinal. In the natural con- dition it is covered by a non-olfactory epithelium, and is thus distinguish- able in function as well as in position from the latter. 1235 DEL evPAeIE IL A. The maxilla, the largest element of the facial region, is associated with its fellow of the opposite side to form the main portion of the upper jaw. It consists of a central portion, the body (corpus maxillae), and of five processes, namely, alveolar, palatine, orbital, zygomatic, and spheno- orbital. In the adult condition the zygomatic bone is fused with the maxilla, so that the extent of the zygomatic process appears to be greatly increased. The body of the maxilla is greatly fenestrated on its external sur- face, the perforated area extending backward to the anterior rim of the orbit, and thus including the maxillary fossa and the infraorbital foramen. The dorsal boundary of the bone is formed by the frontal process of the premaxilla and by the maxillary process of the frontal. . Anteriorly, it is united with the premaxilla, the ventral part of the suture appearing in the diastema separating the incisors from the cheek-teeth. The ventral portion of the bone forms part of the lateral boundary of the incisive foramen. Behind the palatine bridge it is applied to the lateral surface of the palatine bone, and is projected into the orbit as a broad ridge enclosing the alveoli of the four posterior cheek-teeth. In the divided skull the medial surface of the body of the maxilla is found to be concealed by the ethmoturbinal. It contains a deep longitu- 04 ANATOMY OF THE RABBIT. dinal excavation, the maxillary sinus (sinus maxillaris), widely open to the nasal fossa, but only seen to advantage when the ethmoturbinal is removed. The lateral wall of the sinus corresponds in position with the fenestrated area of the external surface. It bears the chief part of the nasolacrimal canal. The alveolar process (processus alveolaris) is that portion of the maxilla lodging the sockets of the cheek-teeth. In the rabbit it is separated by the diastema, in which no teeth occur, from a corresponding but imperfectly differentiated process of the premaxilla. The palatine process (processus palatinus) extends toward the median plane. It forms with its fellow of the opposite side about two-thirds of the palatine bridge. The orbital process (processus orbitalis) is directed obliquely toward the dorsal surface of the skull. In conjunction with the lacrimal bone and the maxillary process of the frontal, it forms the anterior orbital rim. It is continuous with the fenestrated portion of the body, and its appearance as a process is largely due to its solid character as com- pared with the perforated surface lying in front of it. The zygomatic process (processus zygomaticus) forms the anterior root of the zygomatic arch, and in the adult condition has fused with it the anterior end of the zygomatic bone. Its ventral angle bears a prominent masseteric spine for the attachment of the ligament of the masseter muscle. The sphenoorbital process (processus sphenoorbitalis) lies on the medial wall of the orbit, in a position opposite to the middle portion of the ridge lodging the posterior cheek-teeth. It forms a stout buttress, the tip of which is applied to the anteroventral angle of the frontal bone. In this position it is visible from the orbit, lying between the lacrimal bone and the ethmoidal process of the orbitosphenoid. 13), SEE PRE MAXIE AS The premaxilla, or incisive bone (os incisivum), forms the anterior portion of the upper jaw. It comprises a central portion, the hody— including with the latter the scarcely differentiated alveolar portion con- taining the large and small incisors—a frontal process, and a palatine process. The body forms a portion of the palatal surface of the skull and of the lateral boundary of the incisive foramen. Its dorsal surface forms part of the boundary of the piriform aperture, the remaining por- tion of this being formed by the nasal bone. The palatine process extends backward on the medial side of the bone, closely applied on the palatal surface to its fellow of the opposite side, and forms in this way a medial boundary for the incisive foramen. Its dorsal surface, in conjunction with that of the corresponding process of the other side, bears a broad palatine groove (sulcus palatinus), lodging a portion of the cartilage of the vomeronasal organ and nasopalatine duct. The frontal process (pro- cessus frontalis) is a thin bony splint, extending backward between the nasal and maxillary bones, and terminating between the former and the maxillary process of the frontal. THE BonEs OF THE SKULL. 95 14. THE ZYGOMATIC BONE. The zygomatic bone (os zygomaticum) is a separate element only in very young animals. In the adult it is fused anteriorly with the zygo- matic process of the maxilla, the position of the original suture being roughly identifiable as the point where the free horizontal portion of the zygomatic arch arises from the transverse zygomatic process. It forms an almost sagittal plate of bone bridging the orbit and serving for the attachment of the masseter muscle of the mandible. Its dorsal margin forms posteriorly a smooth, horizontal articulation with the zygomatic process of the squamosal, the end of the bone projecting con- siderably behind the articulation. 15.) THE NASAL; BONE: The nasal bone (os nasale) is a thin, elongated bone forming the roof of the nasal fossa, and, in conjunction with its fellow of the opposite side, the dorsal boundary of the piriform aperture. It is loosely articulated with the maxilla and with the bone of the opposite side by smooth (harmonic) sutures. The medial margin is supported by the dorsal edge of the nasal septum. The internal surface bears the nasoturbinal scroll. 16. THE VOMER. The vomer is the median, somewhat sickle-shaped, vertical plate of bone separating the ventral portions of the nasal fossae. It is visible from the palatal surface through the incisive foramina, but its extent is best shown in the divided skull. It forms a support for the ventral border of the nasal septum, and its posterior portion bears a shelf-like projection, the ala vomeris, which assists in the support of the ethmo- turbinal. i, THe TACRIMAL, BONE. The lacrimal bone (os lacrimale) is a small element lying in the anterior wall of the orbit. It is loosely articulated with the surrounding bones, and in the dried skull is frequently missing unless care has been taken to keep it in place. It consists of a basal portion, somewhat rectangular on its orbital surface, and of two processes, namely, the subcutaneous process and the hamulus lacrimalis. The subcutaneous process is the prominent hook-like projecting laterad beyond the orbital rim. The hamulus lacrimalis is a small process, directed toward the nasal cavity. It bears a groove which, in association with a corresponding groove of the maxillary bone, forms the first portion of the nasolacrimal canal. S20 Ee Ae AEN EB ONES The palatine bone (os palatinum) forms the posterior portion of the palatine bridge and the major portion of the lateral wall of the naso- pharynx. It consists of two portions—horizontal, and perpendicu- lar. The horizontal portion (pars horizontalis) is that lying in the plane 96 ANATOMY OF THE RABBIT. of the palatal surface. It is articulated in front with the palatine process of the maxilla, the suture between the two bones enclosing the greater palatine foramen, the ventral termination of the pterygopalatine canal. The perpendicular portion (pars perpendicularis) is the vertical plate extending backward from the palatine bridge. Its medial surface is divided by a low ridge into a dorsal portion, in particular relation to the nasopharynx, and a ventral portion, in relation to the oral cavity, the ridge indicating the position of the soft palate. Its lateral surface is partly applied to the maxilla and partly exposed to the orbit. Its dorsal margin is articulated with the presphenoid and with the ethmoidal process of orbitosphenoid, but a small posterior portion is free, so that the anterior portion of the basisphenoid is visible from the orbit. The free ventral margin forms posteriorly a thick projecting angle, the pyramidal process (processus pyramidalis), the base of which is cleft where it articulates with the medial and lateral laminae of the pterygoid process. Between the pyramidal process and the alveolus of the last cheek-tooth there is a conspicuous palatine notch (incisura palatina), connecting the orbit with the palatal surface. In the entire skull only the posterior portion of the lateral surface is visible from the orbit, the anterior portion being concealed by the projecting bases of the posterior cheek-teeth. The ridge of bone on which the alveoli of these teeth are borne is separated from the palatine bone by the infraorbital groove. The medial wall of the latter, formed by the palatine bone, contains the orbital opening of the pterygopalatine canal and the sphenopalatine foramen. 19. THE MANDIBLE. The mandible (mandibula), or lower jaw, comprises the two dentary bones (ossa dentalia), which, in the rabbit, as in mammals generally, are united by a fibrous or fibrocartilaginous connection (symphysis mandibulae); not coalesced, as in the human skull, to form a continuous structure. As indicated above, each of the dentary bones comprises: (1) a horizontal, tooth-bearing portion which, in conjunction with that of the opposite side, forms the body of the mandible (corpus mandibulae) ; and (2) a posterior, vertical plate, the mandibular ramus (ramus mandi- bulae), for muscle attachment and articulation. The horizontal portion is deep posteriorly, where it lodges the alveoli of the cheek-teeth. Anteriorly, in the diastema separating the latter from the incisors, its dorsal surface is rounded and depressed, the space thus formed corres- ponding to a similar space in the upper jaw and serving chiefly for the accommodation of the lips, which in this region encroach medially on the oral cavity. The medial surface of the horizontal portion forms an acute angle with that of the bone of the opposite side, except anteriorly, where it bears a roughened area for articulation with the latter. Run- ning backward from the symphysis there is a broad horizontal ridge, representing the mylohyoid line (linea mylohyoidea), the line of attach- ment of the mylohyoid muscle. The mandibular foramen, through which, in the natural condition, the inferior alveolar nerve and artery gain access to the interior of the bone, lies on this surface at the junction pas Sal ee THe Hyor. 97 of the horizontal portion with the ramus. The corresponding mental foramen (foramen mentale), through which branches of these structures leave the mandible, is situated on the lateral surface in front of the first premolar. The mandibular foramen is closely connected with a second aperture lying at the ventral end of the sulcus ascendens, directly behind the last molar, and serving for the transmission of a vein connecting the inferior alveolar and inferior orbital veins. The mandibular ramus forms in general an obtuse angle with the horizontal portion. As in other herbivores, the ventral part, distin- guished as the angle, is greatly increased in size at the expense of the condyloid process and to a still greater extent of the coronoid process, the latter being vestigial. In addition to a low pterygoid tuberosity (tuberositas pterygoidea), situated at the posterior projecting point of the angle, the posterior and ventral margins of the angle are excavated on the medial side of the bone, so that they form the boundary of a pro- nounced, though shallow, pterygoid fossa for the insertion of the ptery- goideus internus muscle. A somewhat similar, but less developed, masseteric fossa occupies the corresponding lateral surface of the angle, its raised ventral margin terminating posteriorly in the masseteric tuberosity (tuberositas masseterica). The articular portion, or head of the mandible is greatly elongated in the anteroposterior direction in accordance with the anteroposterior action of the lower jaw, this feature being one which is of general occurrence in the rodent order, and more fully expressed in the great extension forward and backward of the attachment areas of the muscles of mastication. The connection of the articulating portion with the condyloid process, the so-called neck of the mandible (collum mandibulae), is a thin plate of bone, the anterior and posterior margins of which are barely notched by the anterior and posterior mandibular incisures. Connecting the anterior incisure with the rim of the alveolus of the last cheek-tooth there is a deep groove, the sulcus ascendens, the lateral margin of which is formed by the re- duced coronoid process (processus coronoideus). Its low medial margin is formed by a bony stay which extends to the medial surface of the horizontal portion opposite the last cheek-tooth, and is continued for- ward into the mylohyoid line. The sulcus ascendens lodges in the natural condition the insertion portion of the greatly reduced temporalis muscle. Ceti VOM Ab PARATLUS: The hyoid bone (os hyoideum) (Fig. 36) is a stout, somewhat wedge- shaped bone lying in front of the larynx and between the angles of the mandible. Its ventral portion is connected with the thyreoid cartilage of the larynx by the median hyothyreoid ligament. With its lateral portion are articulated two independent elements, termed the lesser and greater cornua. The lesser cornu (cornu minus) is a small, partly car- tilaginous structure, attached to the anterodorsal angle of the hyoid, and connected through the stylohyoideus minor muscle with the jugular process of the skull. The greater cornu (cornu majus) is a larger element extending obliquely dorsad, and similarly suspended from the jugular 98 ANATOMY OF THE RABBIT. process by the stylohyoideus major muscle. The connection of the lesser cornu with the styloid process through the stylohyoideus minor replaces the stylohyoid ligament of the human skull and the chain of elements commonly occurring in mammals and other vertebrates in this region. The muscle tendon contains near the jugular process a small ossification representing a detached styloid process. This connection, together with the hyoid bone itself, indicates the relation of the em- bryonic hyoid arch, from which the skeletal structures in question are derived. The greater cornu belongs to the succeeding visceral arch, and is connected with the superior cornu of the thyreoid cartilage of the larynx by the lateral hyothyreoid ligament. THE SKELELFON OF THE ANTERIOR LIMB. The skeleton of the anterior limb is divisible into two portions, namely, a proximal portion, comprising the scapula and the clavicle, and a distal portion, comprising the supports of the free extremity. The scapulae and clavi- cles of the two sides together form the pectoral girdle. The pectoral girdle is 3 Pic: oe ie surface of us . . y mx: tas ryt i lightly constructed, and, apart from its gid and Jaryns. ca, arvteno muscular connections, which constitute inferior commu’ of thyreoid cartilage: : 5 5 é ce.m., lesser cornu of hyoid; c.mj., its main support, is directly attached to greater cornu of hyoid; ¢.s , superior as zs * cornu of thyreoid cartilage; c.t., left the axial skeleton only through the Ite OF Gen RRR EO id cart ieee amen sternoclavicular ligament Epis tie) ea lake | a ae 7 ; . e oramen; JO ew coe ka, yot yreoi The skeleton of the free extremity 1s ligament; 1.h.m., median Ce oe Si sehive . e . a lig: ont; set... cothyre . divisible into proximal, middle, and SHG ih Is bance a. distal segments. The proximal segment stylohyoideus minor muscle; s.mj., : . 5 stylohyoideus major muscle; irs contains a single bone, the humerus; cartilaginous tracheal rings. the middle segment two elements, the radius and ulna; while the distal segment comprises, in addition to the accessory sesamoid bones, twenty-eight elements of the regular series, of which nine form the carpus, five the metacarpus, and fourteen the phalanges of the digits. DEE SCAPRWIEA: The scapula (Fig. 37) is a somewhat triangular plate of bone lying in the natural position on the lateral surface of the anterior part of the thorax, with its apex directed downward and forward. ‘In the rabbit, as in quadrupedal mammals generally, the main surfaces are respectively medial and lateral, and differ in this respect from the human condition, in which, from the transverse widening of the thorax, the corresponding surfaces are more nearly ventral and dorsal. Of its Tue ANTERIOR LIMB SKELETON. 99 three borders, one, the superior border (margo superior), is directed toward the occiput; another, the vertebral border (margo vertebralis), toward the vertebral column; and the third or axillary border (margo axillaris), toward the armpit. The corresponding angles are medial, inferior, and lateral. The lateral surface bears a stout bony plate, the scapular spine (spina scapulae), which arises from the body of the bone through about two-thirds of its extent, and ends ventrally in a free pro- jection, the acromion. The posterior margin of the acromion bears ¢ backwardly-directed process, the metacromion (processus hamatus). Through the presence of the scapular spine, the lateral surface of the bone is divided into two areas for muscular attachment. One of these, the supraspinous fossa (fossa supraspinata), lies in front of the spine, the other, the infraspinous fossa (fossa infraspinata), behind it. The infraspinous fossa is the more extensive one. The medial surface, on the other hand, presents a single large shallow depression, the sub- scapular fossa (fossa subscapularis), which is triangular in shape and Leathe yy Fic. 37. Lateral surface of the left scapula: a., acromion; a.i., a.l., and a.m., inferior, lateral, and medial angles; c.g., glenoid cavity; c.s., neck of the scapula; f.s. and f.i., supraspinous and infraspinous fossae: m., metacromion; m.a., m.s., and m.v., axillary, superior, and vertebral borders; p.c., coracoid process; s.s., scapular spine. occupies practically the entire surface. The apex or lateral angle of the scapula, sometimes termed the head of the bone, is expanded to a considerable extent in comparison with the slender portion—the so- called neck of the scapula (collum scapulae)—connecting it with the body of the bone. It bears a concave depression, the glenoid cavity (cavitas glenoidalis), for articulation with the humerus. The articu- lating surface is borne chiefly on that part of the bone corresponding to the axillary border, but it also extends in an anterior direction to the base of an overhanging projection, the coracoid process (processus coracoideus). The free portion of the latter forms a blunt, hook-like projection lying toward the medial surface of the bone. 100 ANATOMY OF THE RABBIT. THE CLravicTrs. The clavicle (clavicula) is imperfectly developed in the rabbit, con- sisting of a slender, arcuate rod of bone, tipped by cartilage, which lies in the interspace between the manubrium sterni and the head of the humerus. It occupies only a portion of this interspace, being attached medially by the sternoclavicular ligament and laterally by the cleido- humeral ligament. THE HUMERUS. The humerus (Fig. 38) is typical of the long bones of the proximal and middle segments of the fore and hind limbs in consisting of a central portion, the body or shaft of the bone, and of proximal and distal extremities for muscle attachment and articulation. The proxi- mal extremity bears on its medial side a smooth, convex projection, the head of the humerus (caput humeri), for articulation with the scapula. The articulation is nominally a ball-and-socket joint, or enar- throsis, but the articulating surfaces are somewhat restricted, and the muscular arrangements of the limb are such that the range of lateral motion (abduction and adduction) is small. Immediately in front of the head of the bone there is a small elevation, the lesser tubercle (tuberculum minus). It is separated by a longitudinal furrow of the anterior surface, the inter- tubercular groove (sulcus intertubercularis), from a much larger lateral elevation, the greater tubercle (tuberculum majus). Ex- tending distad from the latter is a tri- angular area, the humeral spine (spina humeri), the tip of which reaches almost to the middle of the bone and forms a pronounced angle on its anterior surface. The distal extremity of the humerus bears a grooved articular surface, the trochlea humeri, for articulation with the th. radius and ulna. On its lateral side IS eas pee eos ete ee eee melee smaller surface, the capitulum humeri, for humerus; 11. and e.m., lateral and articulation with the radius alone. Im- Pee ee pein ce mediately above the trochlea the medial and — cular Brot eee searaeee a lateral portions of the bone are thickened to _ tubercles. form two areas for muscular attachment. One of these, the lateral epicondyle (epicondylus lateralis), is a general point of origin for the extensor muscles of the dorsal surface of the hand, while the other, the medial epicondyle (epicondylus medialis), is — pg a = ACN II LT i NS AEE Ae ee ee ee ee ee THe ANTERIOR LIMB SKELETON. a similar point of origin for the flexor muscles of the ventral or volar surface. Between the epicondyles the extremity of the bone is greatly excavated, so that the projecting portions of the radius in front and of the ulna behind are received into depressions of the surface when the forearm is greatly flexed or extended. On the anterior side is the radial fossa (fossa radialis); on the posterior side the olecranon fossa (fossa olecrani), so- called because it accommodates the olecranon process of the ulna. THEARADEWUS SAND: ULNA: The radius (Fig. 39) is the shorter of the two bones of the forearm, since its proximal extremity does not extend backward beyond the front of the elbow joint. It is antero- dorsal in its general position, but is crossed on the ulna in such a way that its proximal extremity tends to be lateral, while its distal extremity is medial. The proximal extremity, termed the head of the radius (capitulum radil) is immovably articulated with the ulna. It bears an extensive articular surface, meet- ing both the trochlea and capitulum of the humerus, and thus forming a considerable portion of the elbow-joint. The body of the bone is solidly united with the ulna by the interosseous ligament of the forearm. distal extremity is largely formed by an epiphysis, which is well- marked even in older animals. surface (facies articularis carpea), for articula- tion with the navicular and lunate bones. The ulna (Fig. 39) is a somewhat S-shaped bone, the shaft of which is vertically flattened, so that it possesses two main surfaces, respec- tively anterodorsal and posteroventral. The former, in conjunction with the related sur- face of the radius, continues the area of origin of the extensor muscles of the hand from the lateral epicondyle of the humerus distad on to the forearm, while the latter has a similar function with respect to the flexor muscles. The proximal portion of the bone is laterally compressed. It bears a crescentic depression, the semilunar notch (incisura semilunaris), 9 athe: It bears a grooved, carpal articular p tr n. mi. C. ate Cp. M Fic. 39. Skeleton of the fore- arm and hand from the dorsal surface: R, radius; U, ulna; C, carpus; M, metacarpus; 12). phalanges; I-V, metacarpal bones; c., central bone; cp., capitate; . c.r., head of radius; f:a.c., carpal articular surface of radius; h., hamate bone; iss., semilunar notch of the ulna: 1., lunate bone; mi., lesser mul- tangular; mj., greater mul- tangular; n., navicular; ol., olecranon; p.s., styloid process of the ulna; tr., triquetral bone; u., ungual phalanges. 102 ANATOMY OF THE RABBIT. the articulating surface of which continues that of the medial portion of the head of the radius, and is received into the trochlea humeri. Behind the elbow-joint the bone forms the large projecting portion of the elbow, the olecranon, which is a strong process for the insertion of the extensor muscles (anconaei) acting on the forearm. The distal extremity of the bone is formed by an epiphysis, similar to but much longer than that of the radius. It is immovably articulated with the radius, and its tip is formed by a blunt styloid process (processus styloideus), which is articulated with the triquetral bone of the carpus. The elbow-joint is formed by the trochlea and capitulum of the humerus in conjunction with the semilunar notch of the ulna and the corresponding articular surface of the head of the radius. It is a hinge- joint, or ginglymus, permitting motion in one plane, i.e., extension and flexion of the forearm. The trochlear surface of the humerus, however, has a slight spiral trend, the anterior portion being medial in comparison with the posterior portion. Through the immovable articulation provided by the respective proximal and distal ends of the bones, and also through the interosseous ligament, the radius and ulna are unable to change their positions with respect to one another; in other words, the radius is unable to rotate on an axis formed by the ulna, the fore foot being fixed in a position comparable to that of pronation in the human hand (cf. p. 33). Es eC ARSEIGS: The carpus (Fig. 39) comprises nine small elements, the wrist or carpal bones (ossa carpi), which are interposed between the forearm and the digits. They are arranged in two main rows, namely, a proximal row, the elements of which are articulated with the radius and ulna; and a distal row, the elements of which are articulated with the five bones of the metacarpus. Enumerated from the medial side of the wrist laterad, the proximal row contains four elements, namely, the navicular, lunate, triquetral and pisiform bones. The navicular and lunate are articulated with the distal extremity of the radius; the triquetral with the styloid process of the ulna. The pisiform bone lies on the ventral surface of the extremity of the ulna, and is therefore not exposed to the dorsal surface of the wrist. The distal row contains five elements, namely, the greater multangular, lesser multangular, central, capitate, and hamate bones. The first, second, and fourth are in association respectively with the first, second, and third metacarpals. The central bone lies to the lateral side of the articulation at the base of the second metacarpal. As its name implies, it is originally an element interposed between the proximal and distal rows. The hamate is a comparatively large element associated with the fourth and fifth metacarpals, but extending also to the articulation of the third, where it tends to replace the greatly reduced capitate. Tur PostERIOR LIMB SKELETON. 103 THE METACARPUS AND PHALANGES. The metacarpus (Fig. 39) comprises five stout elements, the meta- carpal bones (ossa metacarpalia), which form the basal supports of the digits. Each consists, in addition to a main portion or body, of a flattened proximal end, or base, and a rounded distal extremity, or head. The four lateral bones are normally developed, while the first, which belongs to a reduced digit, is of very small size. The phalanges or bones of the digits are distributed according to the formula 2, 3, 3, 3, 3. They are similar in form to the metacarpals, with the exception, however, of the terminal, ungual phalanges, which are laterally compressed, pointed, and cleft at their tips for the attachment of the claws. SESAMOID BONES. Accessory elements, sesamoid bones (ossa sesamoidea), developed in connection with the ligaments of muscles, are found on the volar surface of the foot in association with certain of the joints. They occur in trans- verse pairs at the metacarpophalangeal articulations and in linear pairs at the articulations of the second with the third phalanges. The pisi- form bone of the carpus is also a sesamoid, being formed in the insertion tendon of the flexor carpi ulnaris muscle. ai SKE LON TOK THE: POSTERIOR LIMB. In the posterior limb the proximal or girdle portion comprises the paired coxal bones, which are united ventrally at the pelvic symphysis, thus forming the pelvic girdle. The distal portion—comprising, as in the anterior limb, the supports of the free extremity—is divisible into proximal, middle and distal segments. The proximal segment contains a single element, the femur; the middle segment two elements, the tibia and fibula, which, however, are extensively coalesced; and the distal segment twenty-three elements, of which six form the tarsus, five the metatarsus, and twelve the phalanges. LEE COXAL BONE: The coxal bone (os coxae) (Fig. 40) is a somewhat triradiate struc- ture, the posterior limbs of which are united, so that they enclose a large aperture, the obturator foramen (foramen obturatum). It is firmly articulated with the sacrum, and is united with its fellow of the opposite side by a thin strip of cartilage containing a small amount of fibrous material. The latter connection is the pubic symphysis (symphysis pubis), better termed in the rabbit the pelvic symphysis, since it is somewhat more extensive than the corresponding articulation of the human pelvis. In the young animal each half of the pelvis consists of three elements, namely, the ilium, ischium, and pubis. They form the three rays of the coxal bone, and are united with one another in the region of the acetabulum, which is the basin-like depression for the articulation of the pelvis with the femur. Only two of the original elements, however, actually take part in the formation of the acetabulum, the pubis being 104 ANATOMY OF THE RABBIT, excluded through the development in the acetabular depression of a small triangular element, the os acetabuli. Although completely coalesced in the adult condition, and showing but few traces of their original separation, the three chief elements are nevertheless describedas if distinct. The ilium (os ilium) is the anterior, also somewhat dorsal, portion of the bone; that part extending forward from the acetabulum. It comprises a basal portion, the body (corpus oss. ilium), which includes the anterior portion of the acetabulum and the cylindrical part of the bone in front of it, and an ex- panded portion, the iliac wing (ala oss. ilium), for muscle attachment and articulation with the sacrum. The body is somewhat triangular in section, its surface being divided into three areas, which are respectively medial, or sacral, ventrolateral, or iliac, and dorso-lateral, or gluteal. The corresponding borders are respectively ventral, or pubic, lateral, or acetabular, and dorsal, or ischial. The acetabular border terminates a short distance in front of the acetabulum in an abruptly truncated projection, the inferior anterior spine (spina anterior inferior). The ischial border forms the anterior half of a long depression of the dorsal surface of the coxal bone, the greater sciatic notch (incisura ischiadica major). The pubic border presents on its medial side a faint, ridge-like eleva- tion, the iliopectineal line (linea iliopect- inea), which connects the sharp anterior border of the pubis with the articular surface for the sacrum. The wing of the ilium forms a shovel- like expansion, the natural position of which is almost sagittal. Its lateral surface provides a fairly extensive area for the origin of the gluteal muscles. Its medial surface is a muscle surface only in its anterior portion, the posterior portion being occupied by the roughened auricular surface (facies auricularis), Fic. 40. Lateral surface of the left coxal bone: IL, ilium; IS, ischium; P, pubis; a., acetabulum; a.i., iliac wing; c.i., body of ilium; c.is., body of ischium; cp., body of pubis; cr., iliac crest; e.i., iliopectineal eminence; f.a., acetabular fossa; f.o., obturator foramen; rlefeh acetabular notch; i.mi., lesser sciatic notch; i.mj., greater sciatic notch; 1.i., iliopectineal line; p-l., lateral process of ischial tuberosity; r.i.i., inferior ramus of ischium; r.i.p., inferior ramus of pubis; r.s.i., Superior ramus of ischium; T1.s.p., superior ramus of pubis; s.a.i., inferior anterior spine of the ilium; s.a.s., superior anterior spine; s.i., ischial spine; s.p., symphysis pubis; s.p.i., inferior posterior spine; t.i., ischial tuberosity; t.p., pubic tubercle. for connection with the sacrum. The dorsal margin is thin and straight. Posteriorly, where it is associated with the greater sciatic notch, there is a small projection, the inferior posterior spine (spina posterior inferior) Anteriorly it passes by a broad angle into the anterodorsal margin of nA a ia i eal Tue Posterior LIMB SKELETON. 105 the bone, the latter forming the projecting end of the wing, which is dis- tinguished as the iliac crest (crista iliaca). This portion is considerably thicker than the related dorsal and ventral margins, and also bears on its medial side a somewhat hook-shaped process. Its anteroventral angle is the superior anterior spine (spina anterior superior). The ven- tral margin is slightly longer than the dorsal margin, and is also concave. It is associated with the pubic border of the body of the ilium, and is not connected with the inferior anterior spine. The ischium (os ischii) extends backward from the acetabulum, its axis continuing thatof the illum. It consists of a basal portion, or body (corpus oss. ischii), a superior ramus, and aninferiorramus. The body of the ischium is for the most part cylindrical. It forms the posterior part of the acetabulum, and presents in connection with the latter a deep acetabular notch (incisura acetabuli), which tends to interrupt the articular surface. The acetabular notch leads forward into a depression of the centre of the articular basin, the acetabular fossa (fossa acetabuli). In the natural condition the combined depressions serve for the attach- ment of the round ligament of the head of the femur. The dorsal margin of the bone, belonging in part to the body and in part to the superior ramus, bears a short hook-like projection, the ischial spine (spina ischiadica). The spine divides this margin into two parts, one of which forms the posterior half of the greater sciatic notch, already described, while the other forms a similar, and, in the rabbit, scarcely less extensive, posterior depression, the lesser sciatic notch (incisura ischiadica minor). The superior ramus of the ischium is the continuation backward of the body of the bone. It is a somewhat flattened plate of bone, the thicker dorsal portion of which terminates in two blunt projections. One of these, the ischial tuberosity (tuber ischiadicum), forms the pos- terior end of the bone, while the other extends in a lateral direction and is described as the lateral process (processus lateralis). The inferior ramus is that part of the ischium which extends from the superior ramus downward and forward between the obturator foramen and the symphysis to meet the corresponding ramus of the pubis. The pubis (os pubis) consists of a basal portion, or body lying imme- diately below the acetabulum, a superior ramus extending from the body to the symphysis, and an inferior ramus extending backward along the symphysis to its junction with the ischium. The anterior margin of the bone, described as the pecten oss. pubis, is thin and sharp. Near the symphysis it bears a minute elevation, the pubic tubercle (tuberculum pubicum), and laterally a more extensive elevation, the iliopectineal eminence (eminentia iliopectinea). The latter is more conspicuous in older specimens, where it is easily recognizable by its jagged outline. Its lateral margin is continuous with the iliopectineal line. THE FEMUR. The femur (Fig. 41) is a somewhat S-shaped bone, the body being very slightly arcuate, while of the two extremities, the distal one is bent downward, forming the articulation of the knee, the proximal one, 106 ANATOMY OF THE RABBIT. with its various processes, slightly upward in association with the pelvis. In considering the general form, it will be remembered that in the natural sitting posture of the rabbit, the position of the femur is approxi- mately horizontal, the convex surface of the shaft, which is equivalent to the anterior surface in man, being uppermost. The proximal extremity of the femur bears an extensive rounded portion or head (caput femoris), for articulation with the pelvic girdle. This portion is separated from the main part of the extremity by a con- stricted area or neck (collum femoris), so that, unlike the case of the anterior limb, the points of muscle attachment fall a considerable distance from the point of articulation. The actual extremity of the bone is formed by a large process for muscular attachment, the great trochanter (trochanter major). It is divided into two portions, one of which, the first trochanter (trochanter primus), forms the large termi- nal, hook-like projection, while the other, the third trochanter (trochanter tertius), is the smaller lateral crest. On the medial side of the bone, immediately distal to the head, there is a triangular elevation, the lesser, or second trochanter (trochanter minor s. secundus). Posteriorly, these pro- jections form a smooth surface for muscle attachment, except, however, at the base of the trochanter major, where the surface of the bone presents a deep, though narrow, depression, the trochanteric fossa (fossa trochanterica). The distal extremity bears an extensive surface for articulation with the tibia. It is divided into two portions, known as the medial and lateral condyles, through the presence of a deep excavation, the inter- condyloid fossa (fossa intercondyloidea). Immediately above the condyles, on thé anterior surface of the bone, the inter- condyloid fossa is replaced by a broad groove, the patellar surface (facies patel- laris), which, in the natural condition, accommodates the convex internal surface of the patella. The medial and lateral Fic. 41. Anterior surface of the left portions of the bone, intervening between femur: ciand cm. lateral and the distal portion of the patellar surface and the tips of the condyles, provide slightly elevated, roughened surfaces, the medial and lateral epicondyles, for muscular attachment. medial condyles; cl.f., neck of femur; cp.f., articular portion (head); e.l. and e.m., lateral and medial epicon- dyles; f.p., patellar surface; t.mi., trochanter minor; t.mj., trochanter major. including t.p. and t.t., the first and third trochanters. THE PosteRIonk Limp SKELETON. 107 THE TIBIA AND FIBULA. The tibia (Fig. 42) is the larger of the two bones of the leg, lying on the medial side of the fibula, and fused with the latter for more than one- half of its length. Its proximal extremity is triangular in section, the main surfaces being respectively anterolateral, anteromedial, and pos- terior. The anterior border is formed by a stout ridge-like elevation, the tuberosity of the tibia (tuberositas tibiae), which in the natural con- dition serves for the insertion of the quadriceps femoris, the extensor tendon being carried over the knee by the patella and the patellar hgament. The arti- biG: nee cular portion is slightly differentiated into y Ge medial and lateral condyles corresponding to those of the distal end of the femur. On the articular surface the concave areas for the reception of the condyles of the femur are separated from one another by a small intervening, partly divided hillock, the inter- condyloid eminence (eminentia intercondy- loidea), and also posteriorly by a depression of the articular border, the posterior inter- condyloid fossa. A corresponding anterior intercondyloid fossa lies in front of the inter- condyloid eminence, but is poorly differ- entiated. The fibula (Fig. 42) is the smaller, lateral bone of the leg, and in the rabbit is so extensively fused with the tibia that scarcely more than a third of it is distinguishable. The free portion forms a flattened bony splint, the medial margin of which is firmly united with the tibia by the interosseous ligament of the leg. Its proximal extremity is connected with the lateral condyle of the tibia by an elongated epiphysis, the latter, like those of the distal ends of the radius and ulna, being distinguishable even in older animals. The combined distal extremities of the tibia and fibula bear a roughly rectangular articular 7” ™— surface for the tarsus. The tibial portion of this surface presents two grooves, separated |, P1s. 2. Anterior surtace of the by a ridge, for articulation with the trochlea cl. and c.m., lateral and medial Pile Onits medial sideis a small) projection, Cue Pas) proximal ane the medial malleolus (malleolus medialis). ™-™., lateral and medial malleoli; ° t.t., tuberosity of tibia. The fibular portion presents a_ trans- verse groove for the convex articular surface of the calcaneus. Imme- diately above it, on the lateral side of the bone, is a prominent projec- tion, the lateral malleolus (malleolus lateralis). It forms the anterior boundary of a groove which in the natural condition lodges the inser- tion tendons of the peroneal muscles. 108 ANATOMY OF THE RABBIT. THE EARS US, The tarsus (Fig. 43) comprises six elements, the tarsal, or ankle-bones (ossa tarsi), which, like the corresponding bones of the carpus, are arranged in proximal and distal rows. An exception is to be made, how- ever, for one element, the navicular, which occupies an intermediate position. The proximal row contains two elements, the talus and cal- caneus. The talus is medial and also slightly dorsal in position. Its proximal end, described as the body (corpus tali), bears an extensive pulley-like surface, the trochlea tali, for articulation with the tibia, these two surfaces together forming the chief portion of the ankle-joint. Its distal end, termed the head of the talus (caput tali), pro- vides a convex articular surface for the navi- cular bone, and is separated from the larger trochlear portion by a slightly constricted intermediate portion or neck (collum tali). Its ventrolateral border is extensively artt- culated with the calcaneus. The latter is a cylindrical element, fully twice as long as the talus, since it is extended backward behind the ankle-joint, as the tuber calcanei, or bone of the heel. Its dorsal surface bears a prom- inent elevation for articulation with the fibular side of the tibiofibula. Its medial surface bears a flat, shelf-like process, the sustentaculum tali, which forms a ventral support for the talus. The distal extremity of the bone articulates with the cuboid and also with the navicular. The intermediate element, the navicular bone, is a somewhat cubical bone, lying on the medial side of the tarsus between the talus, on the one hand, and the proximal end of the second metatarsal bone and the second and third cuneiform bones, on the other. Fic. 43. The bones of the left foot, Its position is more nearly that of a central element than is the case with the bone called by this name in the rabbit’s carpus. In this connection it will be remembered that the carpus and tarsus, like other parts of the limbs, are primarily constructed on the same plan. The distal row of the tarsus contains three elements, namely, the second and third cuneiform bones and the cuboid bone. viewed from the dorsal surface: aye tarsus) Me smetatansus ties phalanges. II-V, the four me- tatarsal bones: cb., cuboid; cl., calcaneus; c.s., second cuneiform; c.t., third cuneiform: fans articular surface for fibular side of the tibofibula: mn mnavicular; t, talus; t:c., tuber calcanei; ‘t:t., trochlea tali. The two former and especially the first, are smaller bones, articulated respectively with the second (first developed) and third metatarsals. In the rabbit the first cuneiform bone—the first element of the distal row in the usual con- = X Tuer Posterior LIMB SKELETON. 109 dition—is fused with the proximal end of the second metatarsal. The cuboid is a larger element articulating, like the hamate bone of the carpus, with two distal elements, the fourth and fifth metatarsals. Its ventral surface bears a transverse elevation, the tuberosity of the cuboid (tuberositas oss. cuboidei), in front of which is a groove for the accom- modation of the peculiar insertion tendon of the peronaeus primus muscle. THE METATARSUS AND PHALANGES. The metatarsus (Fig. 48) comprises five elements, of which four are fully developed and greatly exceed in size the corresponding bones of metacarpus, while one, the first metatarsal, is vestigial. The vestigial element lies on the plantar surface of the foot, for the most part ventral to the navicular and at the base of the second metatarsal. In each developed metatarsal there may be distinguished a main portion or body, a proximal extremity or base, and a distal extremity or head, the last- named portion articulating with the proximal phalanx of the digit. The base of the fifth metatarsal bears a tuberosity for the insertion of the peronaeus secundus muscle. The phalanges are distributed according to the formula 0, 3, 38, 3, 3 the terminal, ungual phalanges being modified like those of the anterior limb. SESAMOID BONES. The sesamoid bones of the posterior limb occur at the knee-joint and on the plantar surface of the foot. On the anterior surface of the knee is the knee-pan or patella, through which, as indicated above, the tendon of the quadriceps femoris muscle is carried over the knee and continued as the patellar ligament to the tuberosity of the tibia. On the posterior surface there are three sesamoid bones, of which one lies in association with the medial condyle of the femur, while the remaining two are asso- ciated respectively with the lateral condyle of the femur and that of the tibia. The sesamoids of the foot are situated at the metatarsophalangeal joints and at those connecting the second and third phalanges. PART III. DISSECTION OF THE RABBIT. The plan of dissection as outlined in the following pages presupposes in the first place that the entire dissection is to be made on a single specimen, and secondly that the latter has been prepared for gross dissection according to the method given in the appendix. These points may be mentioned as explaining many details of procedure and also to a certain extent the selection in preference to others of those structures which are more readily made out by the method employed. Because of the convenience of dissecting in circumscribed regions, the plan has been divided, although of necessity very unequally, into several parts. The order of these is such that the visceral dissection is introduced at an early stage. The somewhat more logical plan of completing first the dissection of the anterior and posterior limbs may be followed, but on account of the fact that it involves a lengthy muscular dissection to begin with, it is perhaps not to be recommended. The account itself aims at a statement of the various structures as met with in order of dissection and the features by which they may be identified, rather than at a full description. The student should make his own observations and prove them by personal drawings and de- scriptions of selected parts. In this connection he will do well to bear in mind that while dissection is nominally a means of obtaining anatom- ical information, its chief value as a laboratory exercise consists in the training to be acquired from critical observation and analysis. It is therefore of quite as much practical importance that he should make his observations extensive and accurate as that he should employ only good instruments, or maintain the proper sequence in dissection. I. EXTERNAL FEATURES. The external structures, subdivisions of the body, and superficial skeletal points may be made out as follows: 1. The division of the body into the head (caput), neck (collum), trunk (truncus), tail (cauda), and the anterior and posterior limbs or extremities (extremitates). 2. Imethe head: (a) The division into a posterior, cranial portion (cranium), and an anterior, facial portion (facies). (b) The mouth (os), bounded by the cleft upper lip (abium superius) and the undivided lower lip (labium inferius). The large sensory hairs or vibrissae. (c) (d) (e) (f) EXTERNAL FEATURES. ight The nose (nasus), and its external apertures (nares an- teriores). The eye (oculus), and its coverings, the eyelids, including the upper eyelid (palpebra superior), the lower eyelid (palpebra inferior), and the third eyelid (palpebra tertia). The third eyelid occupies the anterior angle of the eve, and is compar- able to the conjunctival fold of the human eye. The external ear (auricula), and its canal, the external acoustic meatus (meatus acusticus externus), leading to the tympanic membrane. Points on the head skeleton, to be identified by feeling through the skin; zygomatic arch, supraorbital process, external occipital protuberance, angle of the mandible, symphysis of the mandible, and the hyoid bone. 3. In the trunk : (a) (b) (c) (d) (e) (f) The division into thorax, abdomen, and back, or dorsum. The inclusion with the trunk of the proximal portions of the limbs. The angle formed by the anterior limb with the trunk represents the axillary fossa (fossa axillaris). The depression is much less evident than in man on account of the different positions of its enclosing folds formed by the pectorales and latissimus dorsi muscles. A corresponding inguinal furrow separates the posterior limb from the abdomen and pelvis. The anal aperture (anus), and on either side of it the inguinal spaces, hairless depressions, on which the ducts of the inguinal glands open. In the male: the urinogenital aperture at the extremity of the penis; the latter enclosed by a fold of integument, the prepuce (praeputium); the scrotal sacs (scrotum), lateral sacs of the integument lodging the testes. In the female: the urinogenital aperture, enclosed by folds of the integument, forming the vulva. The clitoris, the homologue of the penis, is contained in its ventral wall. The mammary nipples (papillae mammarum), eight (to ten) in number on the ventral surface of the breast and abdomen. The following skeletal points: on the axial skeleton, the manubrium sterni, xiphoid process, costal arch, spinous processes of thoracic and lumbar vertebrae; on the pectoral girdle, the acromion, clavicle, and respective borders and angles of the scapula; on the pelvic girdle, the iliac crest, pubic symphysis, and ischial tuberosity. 4. In the anterior limb: (a) (b) The division of the free portion into three segments, the arm (brachium), forearm (antibrachium), and hand (manus). The position of the elbow (cubitus) in comparison with the knee. 112 ANATOMY OF THE RABBIT. (c) The five digits, designated from the medial side as: first (d. primus), or pollex, second (d. secundus), or index, third or middle (d. tertius s. medius), fourth (d. quartus), and fifth - (d. quintus s. minimus). 5. In the posterior limb: (a) The division into three segments, the thigh (femur), leg (crus), and foot (pes). (b) The knee (genu), and the popliteal fossa of its posterior surface, the latter not well defined. The projection of the heel (calx), and the angle formed by the foot with the leg. (c) The four digits (dd. secundus-quintus). The vestigial first digit, or hallux, does not appear above the integument. THE ABDOMINAL WALL. HS Il. THE ABDOMINAL WALL. 1. Place the animal in the supine position. Make a median in- cision of the skin of the ventral surface extending from the pubic symphysis to the tip of the mandible, being careful not to cut through more than the skin itself. Make three transverse incisions on the left side, the first on the medial surface of the arm and extending to the elbow, the second midway between the anterior and posterior limbs, the third on the medial surface of the thigh and extending to the knee. Work the flaps loose from the surface, using the handle of the scalpel, until the side of the trunk is well exposed. On the right side of the body it is sufficient to clear the middle line. Identify the structures as follows: On the skin: (a) The thick compact connective tissue forming the corium. (b) The imbedded hair-follicles. (c) The loose subcutaneous tissue (tela subcutanea) by which the skin is attached. (d) In the female: the mammary glands (mammae), forming a layer on the inner surface, and more or less closely aggregated about the mammary nipples. On the exposed surface: (e) The linea alba, a white aponeurotic line extending from the pubic symphysis to the xiphoid process of the sternum. ({) The cutaneus maximus muscle, a thin sheet of muscle fibres covering the entire lateral surface of the thorax and abdomen. Origin: The linea alba, the ventral surface of the sternum in its posterior portion, and the humeral spine. The last- named portion appears on the medial surface of the humerus. Insertion: The skin of the dorsolateral surface of the trunk. The fibres are directed upward and backward. The muscle is continuous across the back with its fellow of the opposite side, and is extended backward to the dorsum of the tail. It is used in shaking the skin. The artery passing forward for a short distance in the inguinal region, and lying in the subcutaneous tissue, is the superficial epigastric, a branch of the femoral (p. 149). The abdominal vein, a conspicuous vessel in the female, traverses the lateral portion of the abdominal wall from the inguinal furrow to the axillary fossa, lying on the external surface of the cutaneus maximus. It is a tributary of the inferior epigastric (p. 134), and anastomoses forwards with the external mammary vein of the axilla. The corresponding arteries are the external mammary branch of the long thoracic (p. 137), and the abdominal branch of the inferior epigastric, the latter arising directly from an external spermatic trunk. The inguinal lymph nodes (lymphoglandulae inguinales) are small, oval, brownish bodies lying in the inguinal furrow. 114 ANATOMY OF THE RABBIT. 2. Remove the cutaneus maximus from the surface. Identify the ms following points of attachment of the abdominal muscles proper: (a) The linea alba. (b) The linea semilunaris, a slightly curved line situated laterally a short distance from the linea alba. (c) The ribs and the costal arch. (d) The lumbodorsal fascia (fascia lumbodorsalis), a broad, white sheet of connective tissue extending over the posterior thoracic and lumbar regions. (e) The inguinal ligament (ligamentum inguinale), a stout white cord, stretched between the symphysis pubis and the iliac Crest 3. Identify on the surface the external oblique muscle (m. obliquus externus abdominis). Origin: The posterior ten ribs by separate slips, the xiphoid process, and the lumbodorsal fascia. Insertion: The linea alba and the inguinalligament. The fibres are directed from an anterior dorsal position downward and backward, the more dorsal ones almost directly backward. Some of the anterior slips of origin interdigitate with those of the thoracic portion of the serratus anterior muscle. Some are concealed by the pectoral muscles. The muscle crossing the breast from the sternum to the arm is the pectoralis major. That passing forward from the lumbodorsal fascia to the medial surface of the humerus is the latissimus dorsi. The margins of these muscles may be raised where they conceal the external oblique. 4. Taking a line between the iliac crest and the xiphoid process, divide the muscle, and then separate it fully from the next, which may be distinguished by the cross direction of its fibres. Note the separate slips of origin and the difference in appearance between the fleshy and aponeurotic portions of the muscle; then remove it from the surface. Examine the following muscles, proceeding in a similar manner: (a) The internal oblique muscle (m. obliquus internus abdominis). Origin: The inguinal ligament, a second sheet of the lumbo- dorsal fascia, and the posterior four ribs. Insertion: The linea alba. The fibres pass downward and forward. The ventral aponeurosis is much broader than that of the external oblique. It contains the rectus abdominis. (b) The transverse muscle (m. transversus abdominis). Origin: Seven posterior ribs, a third sheet of the lumbodorsal fascia, and the inguinal ligament. Insertion: The linea alba. The fibres are directed downward and slightly backward. (c) The rectus abdominis muscle. Origin: Lateral border of the sternum, including the xiphoid process; also the ventral surfaces of the second to seventh costal cartilages. Insert- ion: At the anterior end of the pubic symphysis. It is a thin, strap-like muscle, enclosed by the aponeurosis of the internal ee ga Tur ABDOMINAL WALL. $5 oblique, and separated from its fellow of the opposite side by the linea alba. The artery passing forward, for the most part in this muscle, is the inferior epigastric, a branch of the external iliac (p. 184). It anastomoses with the internal mammary (p. 174). It gives off the external spermatic artery, a small vessel which perforates the abdominal wall and extends backward, supplying the sac of the testis in the male and ending in the female in the wall of the vulva. An abdominal branch of this vessel passes forward on the abdominal wall as described above. 5. Divide the remaining portion of the wall on the left side, and its whole thickness on the right, by a transverse incision, so that the viscera are fully exposed. Note on the internal surface of the wall the smooth serous investment here forming the parietal peritoneum (peritonaeum parietale). 116 ANATOMY OF THE’ RABBIT. Ill. THE STOMACH AND SPLEEN. The cavity disclosed by the division of the abdominal wall is the peritoneal cavity (cavum peritonaei), the largest of the four great serous sacs representing the primary body-cavity, or coelom (p. 49). The major portion of the cavity is abdominal, but it extends into the pelvis, and, in the male, also into the scrotal sacs. Its lining membrane is that appearing on the body-wall as the parietal peritoneum, noted above, and on the visceral structures as the visceral peritoneum (peritonaeum viscerale). The visceral structures here include the major portions of the digestive and urinogenital systems. The general relations of the visceral peritonaeum should first be examined by raising a portion of the small intestine from the left side of the visceral mass. Note its enclosure by a complete serous coat, similar in appearance to the membrane covering the body wall, and the ex- tension of the latter into a mesentery for the support of the structure from the dorsal body-wall. Note the parallel arrangement of the arteries and veins, and also their frequent anastomoses. | Lymphatic vessels (lacteal vessels) accompany the bloodvessels in the mesentery, but being transparent are not readily recognizable. Lymph nodes also occur, but in this portion of the mesentery they are aggregated at its dorsal attachment, or root (radix mesenteril). 1. Displace the posteroventral portion of the liver forward, exposing in this way the ventral surface of the stomach. Without injuring the enclosing peritoneum, examine the contour of the organ and its divisions, as follows: (a) The greater curvature (curvatura ventriculi major), its convex posterior surface. (b) The lesser curvature (curvatura ventriculi minor), the contracted, concave anterior surface. (c) The main portion, or body of the stomach (corpus ventricul1). It lies for the most part to the left of the median plane. (d) The cardia, or area of junction with the oesophagus. (e) The fundus, a small sac-like portion lying to the left of the cardia. (f) The pyloric limb (pars pylorica), lying to the right of the body of the organ. (g) The pylorus, the point of communication of the stomach with the intestine (duodenum). It is marked by an annular constriction, preceding which is a greatly thickened muscular portion of the pyloric limb, known as the pyloric antrum (antrum pyloricum). 2. Raise the posterior portion of the stomach and turn it forward. Note on the dorsal surface of the greater curvature a dark red, elongated THE STOMACH AND SPLEEN. 17 body, the spleen (lien). Trace the course of the peritoneum from the dorsal abdominal wall to the liver, as follows: (a) (b) A broad fold of peritoneum, the mesogastrium, connects the dorsal abdominal wall and the diaphragm with the left side and dorsal surface of the greater curvature. Its posterior portion is divided into two parts by the spleen. The dorsal part, the phrenicosplenic ligament (lig. phrenicolienale) connects the spleen with the dorsal body-wall. The ventral part, the gastrosplenic ligament (lig. gastrolienale) connects the spleen with the greater curvature. The peritoneum is projected backward from the greater curvature as a free fold, the greater omentum (omentum majus), which covers the intestines to a certain extent. It usually contains fat. It is composed of four leaves, of which two come from the surface of the stomach and two others pass forward in a dorsal position from the free posterior border to unite with the transverse mesocolon. The lesser omentum (omentum minus) passes from the lesser curvature and the duodenum to the posterior surface of the liver. Its thickened margin on the right side forms the hepatoduodenal ligament (lig. hepatoduodenale) which carries three important vessels of this relation, namely, the common bile duct, the hepatic artery and the portal vein. Its left portion forms a thin membrane, the hepatogastric omentum, connecting the caudate lobe with the lesser curvature. 3. Working on the left side between the dorsal surface of the stomach and the body-wall, remove sufficient of the peritoneum to expose the first portion of the abdominal aorta as it emerges from the diaphragm. Passing in the direction of the stomach is a median ventral branch, the coeliac artery, the distribution of which may be traced. The following structures, however, should first be identified. 10 (a) (b) (c) (d) (e) The superior mesenteric artery (a. mesenterica superior), a second, also much larger, median branch of the aorta, given off a little distance behind the coeliac artery and passing in the direction of the intestine. The suprarenal gland (gl. suprarenalis) of the left side, a yellowish disc-shaped body lying some distance from the anteromedial margin of the left kidney. The inferior caval vein (v. cava inferior), a large thin-walled vessel lying to the right ef the aorta. It is not conspicuous if empty. The coeliac ganglion (g. coeliacum) of the sympathetic nervous system lies in front of the superior mesenteric artery. The superior mesenteric ganglion (g. mesentericum superius) of the sympathetic system lies immediately behind the su- perior mesenteric artery, and also on its lateral walls. 118 ANATOMY OF THE RABBIT. (f) The nerves proceeding from the coeliac and superior me- senteric ganglia accompany the corresponding arteries, forming the coeliac and superior mesenteric plexuses. (g) The (greater) splanchnic nerve (n. splanchnicus major) of the left side passes backward from its origin in the thorax (see p. 178), around the reduced left crus of the diaphragm and crossing the aorta obliquely enters the superior mesen- teric ganglion. (h) A portion of the pancreas (cf. p. 122) is seenin the peritoneum after the branches of the splenic artery have been severed (4, a). Trace the plan of branching of the coeliac artery, beginning at the point of origin, and exposing the vessels in order. The coeliac artery (a. coeliaca) is a short trunk, its first main branch, the splenic artery, being given off near its origin from the aorta. The remaining portion of the vessel passes to the right in the direction of the lesser curvature, and divides into two parts, the left gastric and hepatic arteries. Small vessels the inferior phrenic arteries (aa. phrenicae inferiores) are given off from the anterior wall of the coeliac and are distributed to the diaphragm. The distribution of the main branches is as follows: (a) The splenic artery (a. lienalis) passes in the direction of the spleen, giving off small branches (rr. pancreatici) to the pan- creas and one or more large vessels, the short gastric arteries (aa. gastricae breves), to the left portion of the greater curvature. Passing along the concave surface, or hilus, of the spleen it gives off several splenic branches (rr. lienales) to that organ, and also several more branches, comparable to the short gastric arteries, to the greater curvature. Toward the end of the spleen the artery passes into the greater omentum, and at this point there is given off a large vessel, the left gastroepiploic artery (a. gastroepiploica sinistra), which passes to the right on the greater curvature and anastomoses with the right gastroepiploic artery. The gastrosplenic ligament, together with its vessels, may be divided, the spleen being allowed to fall backward toward the intestine. (b) The left gastric artery (a. gastrica sinistra) forms a short trunk, or more commonly a group of vessels, the branches of which pass in a somewhat radiate manner toward the lesser curva- ture of the stomach, reaching in this way both dorsal and ventral surfaces. Two larger vessels appear on the ventral surface respectively to the right and left of the cardia. That on the left distributes small branches (rr. oesophagei) to the cesphagus, while that on the right bears a small pyloric branch which anastomoses across the lesser curvature with the right gastric artery. The chief nerves of the coeliac plexus accompany the branches of the artery to the stomach where they connect with the terminal ramifications of the vagus (p. 119). THE STOMACH AND SPLEEN. 119 (c) The hepatic artery (a. hepatica), the continuation of the coeliac, passes forward and to the right, giving off small branches to the pancreas. Its first main branch is the gastroduodenal artery (a. gastroduodenalis). The latter is distributed chiefly to the first portion of the intestine as the superior pancreaticoduodenal artery (a. pancreaticoduoden- alis superior), but a small recurrent branch, the right gas- troepiploic artery (a. gastroepiploica dextra), traverses the greater omentum to the greater curvature where it ana- astomoses with the left gastroepiploic artery. After giving off the gastroduodenal artery, the hepatic enters the lesser omentum on its way to the liver. A small branch, the right gastric artery (a. gastrica dextra) passes to the pylorus and anastomoses across the lesser curvature with a branch of the left gastric artery. The veins of the stomach and spleen are tributaries of the portal vein. Accompanying the branches of the splenic artery are the tributaries of the splenic vein (v. lienalis), including the left gastroepiploic vein. Accompanying the branches of the left gastric artery are the tributaries of the coronary vein (v. coronaria ventriculi). The splenic and coronary veins enter the left wall of the portal vein through a short common trunk. On the right side of the stomach the superior pancreaticoduo- denal vein is united with the right gastroepiploic vein to form a short trunk, the gastroduodenal vein (v. gastroduodenalis), which enters the right wall of the portal vein. The left gastroepiploic vein receives tributaries from the dorsal surface of the pyloric antrum. The abdominal portion of the tenth cranial, or vagus nerve (n. vagus) may be traced from the oesophagus to the surface of the stomach. The left cord appears on the left wall of the oesophagus; crossing the ventral surface of the latter obliquely to the right, it ramifies on the ventral portion of the lesser curvature. The right cord passes backward in a similar manner on the dorsal surface of the oesophagus. 4. Cut across the stomach at the pyloric antrum. Divide the oesophagus, and remove the stomach from the body. Open the organ by means of an incision extending around the greater curvature to the oesophagus. . On the cut end of the pyloric antrum the mucous and muscular tunics (cf. Fig. 15) may be distinguished and separated from one another by dividing the loose tissue of the tela submucosa. On the surface of the mucous tunic may be seen the gastric areas (areae gastricae), formed by the longitudinal folds and imperfect transverse ridges which tend to connect them. They are well marked only in the contracted condition of the stomach. The mucous tunic of the stomach is sharply differen- tiated from that of the oesophagus. 120 ANATOMY OF THE RABBIT. LV. THE ViVi The liver (hepar) is noteworthy, first, as being the largest of the glandular structures of the body, and, secondly, as containing, in ad- dition to the primary circulation formed by the hepatic artery and veins, the ramifications of the portal system. It is an appendage of the digestive tube, its connection with the latter being through the common bile duct. 1. Examine the general contour and plan of division as follows: (a) (b) (c) (d) (e) The convex anterior surface, applied to the diaphragm; the concave posterior surface fitting the convexity of the stomach; the thickened dorsal portion of the organ, and its thin posteroventral margin. The division of the organ into right and left lobes, and of each lobe into anterior and posterior lobules. The gall bladder (vesica fellea), situated on the posterior surface of the right anterior lobule. The quadrate lobe (lobus quadratus), an imperfectly de- veloped lobe lying to the medial side of the gall bladder. The caudate lobe (lobus caudatus), a small independent lobe projecting backward from the base of the left posterior lobule, and accommodated in the natural condition in the space enclosed by the lesser curvature of the stomach. 2. Trace the peritoneal connections as follows: (a) (b) (c) (d) The lesser omentum, represented by the hepatoduodenal ligament. The falciform ligament (lig. falciforme hepatis), a broad median sheet connecting the anterior surface of the liver with the diaphragm and extending backward to the ventral abdominal wall. The position of this ligament indicates the line of division of the liver into right and left lobes. The free arcuate border of the ligament contains a thin cord, the round ligament (lig. teres hepatis), which marks the position of the umbilical vein in the foetus. The coronary ligament (lig. coronarium hepatis), a short circular fold, continuous with the falciform, and connecting the anterior surface of the liver with the middle of the diaphragm. The left triangular ligament (lig. triangulare sinistrum), a lateral continuation of the coronary connecting the left lobe with the diaphragm. 3. Trace the branches of the common bile duct, the hepatic artery and the portal vein. These structures traverse the lesser omentum side by side and their branches are similarly arranged. (a) (b) (c) THE LIVER. 121 The common bile duct (d. choledochus) is formed on the posterior surface of the liver by the union of a left hepatic duct (d. hepaticus) with a similar duct from the right anterior lobule. The latter receives the cystic duct (d. cysticus) from the gall bladder. Special ducts from the right posterior lobule and from the caudate lobe enter the common bile duct through a short common trunk. The common bile duct passes backward on the right side of the portal vein, and enters the digestive tube on the dorsal surface of the first (superior) portion of the duodenum immediately beyond the pylorus. The hepatic artery (a. hepatica) approaches the liver by passing forward on the right side of the portal vein ventral to the bile duct. ‘It distributes branches to the right posterior lobule and the caudate lobe, and at the base of the liver proper divides into right and left rami. The right ramus sends a branch, the cystic artery (a. cystica), to the gall bladder. The portal vein (v. portae), a vessel of large calibre, but usually found in a collapsed condition, enters the lesser omentum from the dorsal surface of the pyloric antrum. It distributes branches to the right posterior lobule and the caudate lobe; then passing directly forward to the base of the left lobe is distributed to the latter, a right branch being given off to the right anterior lobule. 4. Divide the lesser omentum with the structures described above. Divide the coronary and triangular ligaments, beimg careful not to injure the central tendon of the diaphragm, which resembles the coronary ligament. following: (a) (b) (c) Remove the liver and examine its dorsal surface for the The inferior vena cava; it is accommodated in a depression of the thickened dorsal portion of the organ. The vessel should be opened lengthwise. The hepatic veins (vv. hepaticae) open almost directly from the substance of the liver into the inferior cava. They are typically four in number, there being separate vessels for the anterior and posterior parts of the right lobe and for the caudate lobe, in addition to a large vessel for the left lobe. The renal impression (impressio renalis), an extensive excavation of the right posterior lobule for the accommoda- tion of the right kidney. 122 ANATOMY OF THE RABBIT. V. THE INTESTINES. The posterior portion of the digestive tube, or that portion extending from the pyloric aperture of the stomach to the anal aperture, is divisible into two main parts, not wholly distinguishable in calibre, namely, the small intestine (intestinum tenue), and the large intestine (intestinum crassum). Both are greatly elongated and convoluted. In examining them care must be taken to avoid injury to the bloodvessels and me- senteries, especially the dorsal attachments of the mesenteries, in which the chief plexuses and related ganglia of the sympathetic system will afterwards be traced. For the general relations of the intestines and mesenteries see pp. 41, 50. 1. Beginning at the pylorus, trace the course of the small intestine, as follows: Its first portion, the duodenum, forms a U-shaped loop lying on the dorsal wall of the abdominal cavity to the right of the vertebral column. The end of this loop, when traced from the right side, disappears in the peritoneum and may then be picked up in a forward position on the left side of the mass. This point marks the beginning of the second portion, the mesenterial small intestine (intestinum tenue mesenteriale), which may be traced to its termination on the greatly enlarged caecum. The connection with the caecum is through a rounded semi-expanded sac, the sacculus rotundus. The terminal portion is somewhat more difficult to follow on account of the adhesions of its peritoneum with that of the large intestine. 2. Examine the divisions of the duodenal loop and related structures, as follows: (a) The superior, descending, transverse (horizontal), and as- cending portions of the duodenal loop. (b) The common bile duct, opening on the dorsal wall of the superior portion. (c) The mesoduodenum, a fold of peritoneum joining the various parts of the loop. (d) The pancreas (Fig. ,3B, p. 11). Its principal portion is here seen as a diffuse brownish mass lying in the mesoduodenum. Its duct (d. pancreatis) opens into the posterior portion of the ascending hmb. (e) The superior pancreaticoducdenal artery, a branch of the gastroduodenal (see p. 119), passes backward on the first portion of the descending limb. ({) The inferior pancreaticoduodenal artery (a. pancreaticoduo- denalis inferior), a branch of the superior mesenteric (p.125), enters the mesoduodenum from the left side and supplies the major portion of the loop. An anterior branch an- astomoses with (e). weet THE INTESTINES. 123 3. In the mesenterial small intestine the following features may be identified : (a) The lighter coloration, due to the thicker wall and greater vascularity, of the first or duodenal portion, thus distin- guished as the jejunum (intestinum jejunum). (b) The darker coloration, due to the thinner walls, which allow the contents to show through, and diminished vascularity of the terminal or caecal portion, thus distinguished as the ileum (intestinum ileum). The two portions are not distinctly separable. Also the circular folds (plicae circulares), or valvulae conniventes, of the mucous tunic, which in many mammals contribute to the thickness of the wall in the duodenum and jejunum, are in the rabbit not definitely expressed. (c) The mesentery, the peritoneal support of the mesenterial small intestine, is distinguished in its major portion by its broad frill-like character, which allows great freedom of movement to this portion of the digestive tube. Its terminal portion, however, beginning at a point where the intestine turns sharply forward on its way to the caecum, is adherent to the mesocolon. (d) The mesenteric lymph glands (lymphoglandulae mesentericae) are aggregated at the dorsal root of the mesentery, where they form a compact mass surrounding the base of the superior mesenteric artery. (e) The wall of the sacculus rotundus shows externally a hexagonal pattern on account of the presence init of a large number of lymph follicles. Similar structures, forming oval areas about 3mm. in diameter and 5mm. in length, may be found along the intestinal wall (aggregated lymph nodules of Peyer). ({) The finger-like processes, or villi, of the mucous tunic of the small intestine may be seen by making an incision of the wall and examining its internal surface. A small portion of the wall may be excised and examined under water. 4. Trace the course of the large intestine, beginning at the sacculus rotundus, as follows: Its first portion, the blind intestine or caecum (intestinum caecum), distinguished by its great size, is connected with the large intestine proper only in the region of the sacculus rotundus. Its course is comparable to two turns of a spiral. Its closed end, formed by the vermiform process (processus vermiformis), les in a dorsal position, and is directed backward. The second portion, the colon, comprising the major portion of the large intestine proper, leaves the caecum in the region of the sacculus rotundus, in which position it is distinguished by its greatly sacculated walls. The third portion, the straight intestine, or rectum, (intestinum rectum) is a small terminal division situated in the middle line and enclosed for the most part by the pelvis. It is scarcely distinguishable from the related portion of the colon. 124 ANATOMY OF THE RABBIT. 5. In the caecum the following features may be distinguished: (a) (b) The wall, which is otherwise smooth, is divided by a spirally arranged constriction, the latter denoting the position, internally, of a fold of the mucous tunic, the spiral valve. The vermiform process is a narrow, light-colored tube of about five inches in length, the wall patterned externally by lymph follicles, in the same way as that of the sacculus rotundus, and greatly thickened in comparison with that of the caecum proper. Fic. 44. Plan of the connections of the large and small intestines: c., ascending colon; cae., caecum; h., haustra; i., ileum; s.r., sacculus rotundus; t.c., band of the colon; v.S., Spiral valve. 6. The colon is divisible into ascending, transverse, and descending portions, the relations of which may be traced as follows: (a) The ascending colon (colon ascendens) passes from its origin on the caecum to a point forwards on the right side of the dorsal body-wall. This portion is greatly elongated in the rabbit, and instead of passing directly forward follows a tortuous course. It is composed of five principal limbs, united by flexures. Three of the limbs are directed for the most part forward, the remaining two backward. The first limb of the colon bears three rows of small saccu- lations, the haustra, separated by three longitudinal muscle stripes, distinguished as the bands of the colon (taeniae coli). Two of these bands are free, while the third is enclosed by the supporting peritoneum, the mesocolon. The two free bands run together toward the anterior end of the first limb. (b) The transverse colon (colon transversum) is a short segment, beginning forwards on the right and crossing the middle line transversely to the left, where it bends sharply backward, and is replaced by the descending colon. ih ica ——E——— (c) THE INTESTINES. 125 The descending colon (colon descendens) passes backward to a point in front of the pelvis, where it is replaced by the rectum. The descending mesocolon, which connects this portion with the dorsal body-wall, should be noted on account of its relation to the inferior mesenteric artery and sympathetic plexuses. It is connected for a considerable distance with the mesentery of the ascending limb of the duodenum. 7. Displace the caecum, turning it over to the right. Lay out the mesenterial small intestine, so that the mesentery and its bloodvessels are exposed. Remove the lymph glands from about the base of the mesenteric artery, and trace its branches as follows: (a) (b) (c) The middle colic artery (a. colica media), a small vessel arising from the left wall and passing to the transverse colon. The inferior pancreaticoduodenal artery (p. 122) arises at the same level, but from the right wall. The ileocaecocolic artery (a. ileocaecocolica), a large branch, equalling in size the superior mesenteric trunk, is distributed to the terminal portion of the ileum, the caecum (including the vermiform process), and the ascending colon. Its branches are arranged in two series, a proximal group being given off near the point of origin of the main vessel, and a distal group, including the terminal portion of the vessel, at about two inches from the point of origin. The proximal branches include: (1) Small branches to the third, fourth, and fifth limbs of the ascending colon. (2) The appendicular artery (a. appendicularis) to the vermiform process. A branch of this vessel, given off near the point of origin, passes to the ileum, anastomosing with ‘a branch of the superior mesen- teric trunk. (3) An anterior ileocaecal artery to the terminal fourth (third limb) of the caecum proper and related por- tion of the ileum. (4) An anterior right colic artery to the flexure uniting the first and second limbs of the ascending colon. (5) A posterior right colic artery to the second limb of the ascending colon. This vessel anastomoses with (4) and with the special branch to the third limb (1). The distal branches include: (6) A posterior ileocaecal artery to the middle portion of the third limb of the caecum and the adjacent portion of the ileum; anastomosing with (38). (7) A caecal artery to the posterior portion of the third limb of the caecum. 126 ANATOMY OF THE RABBIT. (8) Terminal branches to the parts about the sacculus rotundus; anastomosing with (4). (d) The intestinal arteries (aa. intestinales), about twenty in number, are given off from the superior mesenteric artery, and are distributed to the free portion of the mesenterial small intestine. The successive vessels are connected by anastomoses. The end of the superior mesenteric artery has the relation of one of the intestinal arteries. It anastomoses forwards witha single branch given off from the side of the vessel opposite to the intestinal arteries, and the latter in turn anastomoses forwards with a branch of the appendicular artery. 8. Locate in the descending mesocolon the inferior mesenteric artery (a. mesenterica inferior), a small median vessel arising from the abdominal aorta. It has two main branches—the left colic artery (a. colica sinistra) to the anterior portion of the descending colon (anastomosing with the middle colic), and the superior haemorrhoidal artery (a. haemorrhoidalis superior) to the posterior portion of the colon and the rectum. 9. The superior mesenteric vein (v. mesenterica superior), the chief tributary of the portal, collects the blood distributed by the superior mesenteric artery, its tributaries being similar in arrangement to the branches of the artery. The inferior mesenteric vein (v. mesenterica inferior) collects blood from the descending colon and rectum; it may be traced forward in the descending mesocolon. 10. Sympathetic plexuses. In the descending mesocolon will be found the inferior mesenteric ganglion (g. mesentericum inferius), a narrow curved body situated in front of the inferior mesenteric artery. Surrounding the abdominal aorta and appearing in the mesocolon is the abdominal aortic plexus (plexus aorticus abdominalis). It is connected anteriorly with the coeliac and superior mesenteric plexuses (p. 118) accompanying the corresponding vessels, and with the renal plexuses accompanying the renal vessels to the kidneys; posteriorly with the inferior mesenteric and spermatic plexuses about the inferior mesenteric and internal spermatic arteries, and with the hypogastric plexus about the pelvic vessels. 11. By dividing the rectum close in front of the pelvis, and severing the peritoneal attachments, the intestines may be separated and laid out in an extended condition. The relations to one another of the ileum, caecum, and colon are studied to much better advantage than in the natural position. so eee ted THE URINOGENITAL SYSTEM. 127 VI. THE URINOGENITAL SYSTEM. A. The Urinary Organs. The central organs of excretion, the kidneys (renes), occupy an anterior position on the dorsal wall of the abdomen. The right kidney is placed a little farther forward than the left, and is largely covered by the right posterior lobule of the liver. In addition to a fibrous coat immediately surrounding the kidney substance, each organ is imbedded, in a mass of fatty material, the adipose capsule (capsula adiposa), and is also held in position by the peritoneum, which is stretched across its ventral surface. 1. By removing the peritoneum and adipose capsule from the left kidney, the external features and vascular connections may be made out as follows: (a) The general convexity of contour. (b) The renal hilus (hilus renalis), a concavity of the medial sur- face of the organ. (c) The ureter, or duct of the kidney, a white tube passing back- ward from the hilus. (d) The renal artery (a. renalis), arising from the abdominal aorta and entering the kidney at the hilus. A branch of this vessel, the twelfth intercostal artery (a. intercostalis x1), passes to the body-wall in front of the kidney, giving off a small suprarenal artery to the suprarenal gland. (e) The renal vein (v. renalis), leaving the kidney at the hilus, and joining the inferior cava. 2. Divide the kidney, beginning the incision at the hilus and re- moving the ventral half. Examine the cut surface of the dorsal half for the following: (a) The renal pelvis (pelvis renalis), formed by the expanded funnel-like end of the ureter, which is fitted into the renal hilus. The enclosed space is largely occupied by a conical projection of the kidney substance, the renal papilla (papilla renalis). (b) The cortical substance (substantia corticalis); distinguish- able as a narrow peripheral zone of the kidney substance. (c) The medullary substance (substantia medullaris), forming the central and medial portion of the kidney, including the renal papilla. It is distinguished by its radial striations. (d) The fibrous coat (tunica fibrosa) of the kidney may be stripped from the surface. In the rabbit the kidney is not lobulated. Hence there is a single renal papilla, and the division of the kidney substance into renal pyramids is imperfectly expressed. The medullary substance, however, possesses a slightly divided margin. 128 ANATOMY OF THE RABBIT. The distinction of medullary and cortical portions depends on differ- ences in the arrangement of the minute tubules which compose the kidney. The medullary substance is largely composed of straight collecting tubules converging on the renal papilla, while the cortical substance is occupied by convoluted portions of the tubules and their vascular connections, the glomeruli. In the natural condition the two parts are also distinguishable in coloration, the cortical substance being darker. In embalmed animals, however, the color features are usually reversed. 3. The urinary bladder (vesica urinaria) lies in the ventral posterior portion of the abdominal cavity. It is a muscular sac, capable of a considerable amount of distension, but usually found in preserved animals in a greatly contracted condition. Its rounded anterior end, the vertex, projects forward into the abdominal cavity, while its posterior portion or fundus, narrows to a canal, the urethra, which receives on its dorsal wall the apertures of the genital ducts and those of the related glands. The connections may be made out as follows: (a) The peritoneum is reflected from the ventral surface of the rectum in the male and from the uterus in the female, to the bladder, and after investing the latter passes to the ventral abdominal wall. The dorsal peritoneum forms in the male a double rectovesical fold (plica rectovesicalis), and in the female a similar vesicouterine fold, a recess of considerable extent (rectovesical or vesicouterine pouch) being left between the adjacent structures. The ventral peritoneum forms a broad median vertical sheet, the middle umbilical fold (plica umbilicalis media). The free edge of this fold, extending from the vertex of the bladder to the umbilicus, contains a slender cord, the middle umbilical ligament (lig. umbilicale medium). The latter marks the position of the peripheral portions of the umbilical arteries in the foetus. “| (b) The umbilical artery (a. umbilicalis), a branch of the hypo- gastric, passes along the side of the bladder to the vertex. From the base of the artery branches are given off to the ureter (a. ureterica) and related portions of the genital ducts. B. The Male Genital Organs. 1. Continue the median ventral incision of the skin backward along the symphysis to the penis. Reflect the skin on both sides to clear the attachments of the penis to the ischium, and on the left to a point -beyond the scrotum. Note the cremaster muscle (m. cremaster), a thin layer of muscle fibres forming the outer layer of the sac of the testis. It is continuous with the internal oblique muscle of the abdominal wall, and also contains fibres from the transverse muscle. Make a longi- tudinal incision through this muscle, cutting forward into the abdominal cavity. On spreading apart the two flaps the following features may be made out: (a) The parietal layer (lamina parietalis) of the tunica vaginalis propria, a layer of peritoneum, continuous with that of the THE URINOCENITAL SYSTEM. 129 abdominal wall, forms the internal lining of the sac of the testis (cf. p. 51). The sac is widely open to the abdominal cavity, so that the testis passes freely from one cavity to the other. (b) The male reproductive gland, the testis, with its associated vessels and duct, occupy the cavity of the sac, the testis being suspended from its dorsal wall. (c) The gubernaculum, a short thick cord containing smooth muscle fibres, joins the posterior end of the testis with the end of the sac. (d) The visceral layer (lamina visceralis) of the tunica vaginalis propria forms the peritoneal coat of the testis and is con- tinuous with the mesorchium, a broad vertical fold of peri- toneum connecting the testis forwards with the dorsal body- wall. (e) The first portion of the duct of the testis, the epididymis, forms a thickened mass at the anterior end of the testis, and extends backward as a thinner cord along its side. The thickened anterior portion is the caput epididymidis and the contracted posterior portion the cauda epididymidis. The second portion, the ductus deferens, leaves the testis at its posterior end, where it is firmly attached to the guberna- culum. The connection with the epididymis may be shown by carefully separating the duct from the gubernaculum and the side of the testis. (f) The internal spermatic artery (a. spermatica interna) arises from the abdominal aorta, that of the left side immediately behind the origin of the inferior mesenteric. It enters the anterior end of the testis. (g) The spermatic vein (v. spermatica) is formed by a network of vessels, the plexus pampiniformis, which surrounds the internal spermatic artery in its posterior portion. It opens forwards into the inferior caval at the angle formed by the latter with the renal artery. (That of the right side enters the inferior caval at about the level of the spermatic arteries. ) Owing to the open communication of the testis sac with the abdominal the association of the ductus deferens with the spermatic vessels to form a spermatic cord (funiculus spermaticus) 1s very imperfectly expressed. ) The connections of the deferent ducts with the common urino- genital tube and related parts may be displayed by dividing the sym- physis and dissecting in the interior of the pelvis. The structure and attachments of the penis should first be examined. Apart from its terminal portion, the glans penis, and the urethra, the body of the penis is chiefly formed of a pair of hollow fibrous structures, the cavernous bodies (corpora cavernosa penis). Each of these is firmly attached to the posterior margin of the ischium by a fibrous cord, the crus penis. The latter is largely concealed by a short thick ischiocaver- 130 ANATOMY OF THE RABBIT. nosus muscle, the origin of which occupies a similar position on the ischium. The penis is also attached to the symphysis by a short but stout suspensory ligament (lig. suspensorium) and by a small pubo- cavernosus muscle lying between the latter and the ischiocavernosus. The attachments of the penis should be severed and the symphysis divided. By pressing apart the two sides of the pelvis the urinogenital tubes, with the penis and the pelvic portion of the rectum, may be dissected out and removed from the body. The middle haemorrhoidal artery (a. haemorrhoidalis media), a branch of the hypogastric, passes to the side of the rectum and to the seminal vesicle. The internal pudendal artery (a. pudenda inter- na), accompanied by the corres- ponding nerve and vein, passes to the side of the penis, giving off the inferior haemorrhoidal artery to the terminal portion of the rectum and to the associated rectal or anal gland. The rectum is connected with the root of the tail by the rectocaudalis muscle, a somewhat spindle - shaped aggregation of smooth muscle fibres, arising from the body of the second caudal ver- tebra, and inserted a short distance forwards on the dorsal surface of the rectum. The sphincter ani externus and sphincter ani internus are two closely related muscles enclosing the rectum and urethra, the former arising from the dorsum of the tail. The following parts of the urinogenital system may be made out: (a) The connection of the bladder with the outside of the body through the irethra; It. come prises a short pros- tatic portion in rela- tion to the genital ducts, al much longer membranous portion traversing UV.—-——+ Fic. 45. The male urinogenital ducts and related structures, viewed from the lateral surface. After Rauther: a., anal aperture; bu., bulbourethral gland; c.c., corpus cavernosum; d:d., ductus de- ferens; g.a., anal (rectal) gland; g,i., inguinal gland; g.p., glans penis; par., paraprostatic glands; pr., prostate; r., rectum; ur., ureter; u.v., urethra (membranous portion); v.s., seminal vesicle; v.u., urinary bladder. iad the pelvis, and a terminal cavernous portion in the penis. (b) The seminal vesicle (vesicula seminalis) lies on the dorsal surface of the base of the bladder. It is a flattened median pouch, the anterior, slightly divided tip of which is directed forward. THE URINOGENITAL SYSTEM. 131 (c) The deferent ducts lie between the seminal vesicle and the dorsal wall of the bladder. They terminate in the ventral wall of the seminal vesicle. (d) The prostate (prostata), a white or yellowish mass of glandular tissue, lies in the posterior portion of the dorsal wall of the seminal vesicle. Its ducts, three or four in number on either side, open into the urethra. Accessory paraprostatic glands (gl. paraprostaticae), minute finger-like projections of the urethal wall, lie on either side of the base of the seminal vesicle. (e) The bulbourethral gland (gl. bulbourethralis); situated on the dorsal wall of the urethra behind the prostate. 3. The internal surface of the dorsal wall of the urethra may be exposed by a longitudinal incision extending into the bladder. The crescentic aperture of the seminal vesicle lies immediately in front of an oval elevation, the colliculus seminalis, on either side of which some of the minute apertures of the prostate may be made out. C. The Female Genital Organs. 1. The organs may be traced from the abdominal cavity backward, as follows: (a) The ovary (ovarium) is a small—in young animals minute— elongated structure of greyish or yellowish coloration lying on the dorsal body-wall some distance behind the kidney. It is readily distinguished by the circular translucent dots representing the larger vesicular ovarian follicles. In some cases the darker radiate impressions (corpora lutea) left by extruded eggs are discernible. (b) The mesovarium, a short fold of peritoneum suspending the ovary from the body-wall. (c) The internal spermatic artery (a. spermatica interna) arises from the abdominal aorta, immediately behind the origin of the inferior mesenteric artery, and crosses the body-wall transversely to the ovary. (d) The spermatic vein (v. spermatica) leaves the medial side of the ovary, and crossing the body-wall enters the inferior caval vein. (e) The uterine tube (tuba uterina), the first portion of the ovi- duct, distinguishable by its narrow calibre, opens into the abdominal cavity through a broad funnel-like expansion, the ostium abdominale tubae uterinae. The margin bears a large number of short folds and processes, the fimbriae tubae, which tend to enclose the margin of the ovary. ({) The mesosalpinx is the peritoneum supporting the uterine tube. It is continuous with the mesovarium. (g) The uterus, the second portion of the oviduct; distinguished 132 ANATOMY OF THE RABBIT. by its greater diameter and muscular walls. The size of this portion is enormously increased in animals which contain or have borne young. (h) The mesometrium is the supporting peritoneum of the uterus, and is a continuation of the mesosalpinx. The mesometrium, mesosalpinx and mesovarium together constitute the broad ligament (lig. latum uteri). (1) The round ligament of the uterus (lig. teres uteri) crosses the broad ligament, and may be traced from the anterior end of the uterus to the body-wall below the posterior portion of the inguinal ligament where it is inserted (cf. p. 49). (j) The vagina is a flattened median tube with muscular walls; it receives anteriorly the apertures of the right and left uteri. 2. Preparatory to dissecting the urinogenital structures of the pelvis, the median incision of skin of the ventral surface should be continued backward to the base of the clitoris, the structure and attachments of the latter being then examined as follows: (a) The cavernous bodies (corpora cavernosa clitoridis) form the body of the structure, as in the male, and are attached to the posterior border of the ischium through short fibrous cords, the crura clitoridis. (b) The suspensory ligament (lig. suspensorium clitoridis) is a short median cord joining the base of the clitoris with the posterior end of the symphysis. (c) The pubocavernosus and ischiocavernosus muscles pass from the posterior border of the ischium to the base of the clitoris on either side, the former being medial in position. The attachments of the clitoris should be severed and the symphysis divided. By pressing apart the two sides of the pelvis the urinogenital tube may be dissected out and removed, together with the terminal portion of the rectum. The related structures, the middle haemorrhoidal artery, etc., appearing in this dissection are as in the male (see note p. 130). In the urinogenital ducts examine the extent of the vagina back- wards and its connection with the canal of the bladder to form the common vestibulum. The latter is comparable to the male urethra (cf. p. 47, Fig. 25). The bulbourethral gland (gl. bulbourethralis) (cf. p. 131) lies on the dorsal wall. By slitting open the vestibulum and extending the incision into the bladder and also forward into the left uterus the apertures of these structures may be examined from the interior. There is an external uterine aperture (orificium externum uteri) for each division of the uterus. THE ABDOMINAL AORTA, ETC. 135 VII. THE ABDOMINAL AORTA, INFERIOR CAVAL VEIN, AND SNe Ash DiC EReUINICS: The dissection and removal of the intestines and urinogenital organs clears the dorsal body-wall for an examination of the abdominal portion of the aorta, the inferior caval vein, and the sympathetic trunks. If the inferior cava does not contain blood its tributaries should be cleared first, in order to keep them from being damaged; otherwise the branches of the aorta should first be traced. The anterior portion of the inferior cava has been removed with the liver. 1. The abdominal portion of the aorta, described as the abdominal aorta (aorta abdominalis), extends from the hiatus aorticus of the diaphragm to the seventh lumbar vertebra, where it is replaced by the paired common iliac arteries. It passes backward in a median position along the ventral surfaces of the bodies of the vertebrae. Its natural continuation backwards on the sacrum and the caudal vertebrae is represented by the greatly reduced median sacral artery. The branches of the vessel are distributed in two series: (1) visceral branches (rami viscerales) to the parts of the digestive tube and the urinogenital organs; and (2) parietal branches (rami parietales) to the body-wall. The visceral branches comprise the paired renal and spermatic arteries, and the unpaired coeliac, superior mesenteric and inferior mesenteric arteries, which have already been traced. The parietal branches comprise: (a) The superior phrenic arteries (aa. phrenicae superiores), small vessels arising in the hiatus aorticus and passing to the diaphragm. (b) The twelfth intercostal artery (a. intercostalis x1), arising on either side from the renal artery, and passing laterad to the body-wall. (c) The lumbar arteries (aa. lumbales), seven pairs of vessels distributed metamerically to the lumbar portion of the body-wall. Six pairs arise from the dorsal wall of the aorta, the seventh from the median sacral artery (d), each through a common trunk. (d) The median sacral artery (a. sacralis media) arises from the dorsal wall of the aorta near its posterior end, and passes backward on the ventral. surface of the sacrum. Its first portion is concealed from the ventral surface by the common hypogastric vein. 2. The common iliac artery (a. iliaca communis) is a short trunk, the branches of which pass to the posterior limb, the wall of the pelvis, and the pelvic viscera. It gives off the iliolumbar artery (a. iliolumbalis), which passes laterad to the body-wall, and then divides into two branches the connections of which may be traced as follows: 11 134 ANATOMY OF THE RABBIT. (a) The external iliac artery (a. iliaca externa) is the larger lateral branch, directed toward the inguinal hgament, over which it passes to the medial surface of the limb as the femoral artery. It gives off the inferior epigastric artery (a. epigastrica inferior), which passes forward on the medial portion of the abdominal wall (see note p. 115). (b) The hypogastric artery (a. hypogastrica) is the smaller medial branch, directed backward on the dorsal wall of the pelvis. At its junction with the external iliac the vessel gives off the umbilical artery (a. umbilicalis) to the bladder, or in the female first to the vagina and uterus (a. uterina). The obturator artery (a. obturatoria) passes laterad to the pelvic wall, and the middle haemorrhoidal (see note p. 130) to the side of the rectum. The main vessel leaves the pelvic cavity, passing to the lateral side of the abductor caudae anterior, and reappears posteriorly as the internal pudendal (note p. 180). The intermediate branches pass to the posterior limb and the side of the tail (p. 149). 3. The inferior caval vein (v. cava inferior) is formed on the dorsal surface of the posterior end of the aorta by the union of the paired external iliac veins with the common hypogastric. From this position it passes to the right side of the aorta (rarely to the left) almost to its ventral surface, and then runs forward on the right side to the diaphragm. Its visceral roots or tributaries (radices viscerales) comprise the paired renal and spermatic veins, and the hepatic veins from the liver (p. 121). Its parietal tributaries (radices parietales) include the inferior phrenic veins (vv. phrenicae inferiores), which enter the inferior cava from either side of the diaphragm, the lumbar veins (vv. lumbales), a_series of vessels corresponding to the lumbar arteries, and the paired iliolumbar vein (v. iliolumbalis). 4. The external iliac vein (v. iliaca externa), the continuation of the femoral vein of the thigh, approaches the inferior cava from the dorsal side of the inguinal lhgament. It receives the inferior epigastric vein from the abdominal wall and the vesical vein from the bladder, the latter including in the female also the veins of the uterus. 5. The sympathetic trunk (truncus sympathicus). Its lumbar, sacral, and caudal portions may be traced on either side by working between the abdominal aorta (or its continuation, the median sacral artery) and the body-wall. Except on the ventral surface of the sacrum, the ganglia of opposite sides lie close together. The lumbar portion of each trunk comprises seven ganglia with their connections. The ganglia lie on the lateral surfaces of the lumbar arteries near the points where the latter disappear dorsally in the body-wall. The rami communicantes may be found passing from the ganglia toward the spinal nerve-roots. The sacral portion comprises four ganglia of which the first two are much larger than the others. The caudal portion comprises two minute ganglia and an unpaired terminal ganglion uniting the two trunks. THE ANTERIOR Limes. 135 Vite TEE ANTERIOR IMB: For this dissection the skin must first be reflected from the lateral surface of the limb and the side of the neck to the dorsal median line. Covering the side and ventral surface of the neck is a broad thin sheet of muscle, the platysma, replacing the cutaneus maximus of the trunk. It forms a continuotis layer over the dorsal surface of the neck, at which place it is also continuous with the cutaneus maximus. Passing forward from the manubrium sterni is a narrow band of fibres, in- separable from the platysma but lying beneath it, the depressor conchae (parotideoauricularis) posterior, which is inserted into the external base of the ear. The entire sheet of muscle should be raised from the surface, separated posteriorly from its attachment, and turned forward on the head. The dissection is mainly muscular, but the arteries and nerves should be kept intact for later examination. For the general muscular relations of anterior and posterior limbs see p. 33. *1. Muscles arising from the axial skeleton and inserted on the scapula and clavicle. (a) The cleidomastoideus. Origin: Mastoid portion of the skull. Insertion: Middle portion of the clavicle. The muscle lying on its medial side and arising from the manubrium sterni 1s the sternomastoideus, one of the muscles of the head. (b) The basioclavicularis (basiohumeralis). Origin: Basiocci- pital bone. Insertion: Lateral third of the clavicle and the cleidohumeral ligament. (c) The levator scapulae major. Origin: Sphenooccipital syn- chondrosis. Insertion: Metacromion. The superficial cervical artery (p. 173) passes obliquely forward and outward under cover of these muscles, ramifying beneath the superior portion of the trapezius in the fat-mass of the side of the neck. Its ascending cervical branch passes forward on the lateral surface of the external jugular vein. (d) The trapezius. Origin in two portions: Superior (cervical) portion: External occipital protuberance and the ligamentum nuchae. Insertion: Metacromion and supraspinous fascia. Inferior (thoracic) portion. Origin: Spinous processes of the thoracic vertebrae and the lumbodorsal fascia. Insertion: Dorsal two-thirds of the scapular spine. The muscle forms a broad triangular sheet on the lateral surface of the shoulder. The levator scapulae major, basioclavicularis, and trapezius should be divided. On the ventral surface of the origin of the superior portion of the trapezius and levator scapulae major may be found the ventral rami of the third and fourth cervical spinal nerves. The great auricular nerve (n. auricularis magnus) passes from the third to the ear. — *The structures of Group 2 may be dissected first if preferred, the serratus anterior muscle being exposed from the lateral surface and divided together with the latissimus dorsi. 136 (e) (f) (g) (h) ANATOMY OF THE RABBIT. The rhomboideus minor. Origin: Ligamentum nuchae. Insertion: Anterior two-thirds of the vertebral border of the scapula. The rhomboideus major. Origin: Spinous processes of the first seven thoracic vertebrae. Insertion: Posterior third of vertebral border. By dividing the rhomboidei the scapula may be displaced laterad. The operation is facilitated by dividing the latissimus dorsi, the re- lations of which should, however, first be noted (2, a.). The levator scapulae minor. Origin: Mastoid and supra- occipital portions of the skull. Insertion: Medial surface of the inferior angle of the scapula. The serratus anterior. Origin in two portions: Cervical portion: Transverse processes of the posterior five cervical vertebrae and the anterior two ribs. Thoracic portion: Third to ninth ribs by separate slips alter- nating with those of the external oblique. Common insertion: Medial surface of the vertebral border of the scapula. The transverse artery of the neck (a. transversa colli) lies on the medial side of the cervical portion. 2. Muscles arising from the axial skeleton and the pectoral girdle and inserted on the humerus, for the most part at its proximal extremity. (a) (b) (c) (d) The latissimus dorsi. Origin: Lumbodorsal fascia and four posterior ribs. Insertion: Humeral spine. A long flat triangular muscle, covering a considerable portion of the lateral surface of the thorax; having its dorsal angle covered by the thoracic portion of the trapezius. Its insertion end passes to the medial side of the humerus. The pectoralis primus (p. tenuis). Origin: Manubrium sterni. Insertion: Humeral spine. A branch of the thoracoacromial artery appears between this muscle and the deltoidens. ' The pectoralis secundus (p. major). Origin: Entire lateral portion of the sternum. Insertion: Anteromedial surface of the humerus, beginning below the head, and extending to the boundary between the middle and distal thirds. The anterior fibres are covered by those of (b). Some of the posterior fibres are inserted highest on the humerus. The two muscles should be separated and cut across. The pectoralis tertius (p. minor). Origin consisting of two portions: First portion: Middle line of the sternum between the attach- ments of the second to fourth ribs. Second portion: Manubrium sterni, extending to the level of the first rib. Insertion: The superficial fibres of the first portion are attached to the clavicle. The remaining fibres, combined with those (e) THE ANTERIOR LIMp. 137 of the second portion and those of the pectoscapularis, pass to the dorsal side of the clavicle and over the shoulder to be inserted on the scapular spine. The pectoralis quartus. Origin: The sternum, from the attachment of the fourth to seventh costal cartilages. In- sertion: Anterior surface of the head of the humerus, passing thence to its medial side. The muscle overlaps the more posterior fibres of the first portion of (d). ({) The pectoscapularis. Origin: The sternum at the point of attachment of the first costal cartilage. Insertion as indi- cated above. m.a.i., internal acoustic meatus; Sa branous labyrinth, consisting of a sacculus; s.e., endolymphatic sac; st., second series of spaces contained stapes; t.a., auditory tube; u., utriculus: within the first, and comprising v, vestibulum; VIII, acoustic nerve. the duct of the cochlea, the sacculus, the utriculus, and the semicircular ducts, together with their connections and the endolymphatic duct and sac. The membran- ous labyrinth contains the sensory portion of the ear and its cavity is occupied by a fluid material, the endolymph. The wall is separated from that of the bony labyrinth by an extensive perilym- phatic space also occupied by a fluid material termed the perilymph. 172 ANATOMY OF THE RABBIT. Xx, THE THORS. 1. Examination of the thoracic wall. For this purpose the lateral surface of the thorax may be conveniently cleared, on the side from which the anterior limb has removed, by dissecting away the attachments of the muscles already examined in the previous dissections. These include the origins of the pectorales, pectoscapularis, serratus anterior (thoracic portion), obliquus externus, and rectus abdominis. The dorsal portion of the exposed surface is occupied by the spinal musculature; to be examined at a later stage. On the ventral portion appear the ribs, and between them, filling the intercostal spaces, the intercostal muscles. The external intercostals (mm. intercostales externi) arise from the posterior margins of the bone ribs, the fibres passing obliquely downward and backward to be inserted on the anterior margins of the next succeeding ribs. The internal intercostals (mm. intercostales interni), the fibres of which are disposed in the opposite direction, are concealed for the most part from this surface, but appear ventrally between the costal cartilages, where they are not covered by the external intercostals. They are best examined at a later stage from the interior of the thorax. In preparation for the removal of a section of the thoracic wall, the pectorales should be divided on the opposite side of the thorax, close to the sternum, so that the limb may be displaced. The nerves and vessels of the neck must be kept intact until the following dissection accounts for their thoracic connections. The sealeni muscles (p. 182) should be examined, since it is necessary to destroy their costal insertions. A triangular section of the wall, including the sternum and the costal cartilages, may be removed by making three incisions, one on either side of the sternum, extending from the middle of the first rib backward to the end of the ninth bone rib, and the third across the ventral surface close in front of the diaphragm. The transversus thoracis muscle appears on the inner surface of the section removed. It is a thin sheet of fibres arising from the body and xiphoid process of the sternum and inserted on the ribs, from the second to the sixth, at the junctions of the bone ribs with the costal cartilages. The artery passing along the ventral wall between the foregoing muscle and the internal intercostals is the internal mammary (p. 174). 2. Dissection of structures in the superior thoracic aperture. ° The nerves and bloodvessels of this region are concealed by the thymus gland, a large triangular flattened structure of fatty consis- tence, extending forward from the base of the heart to the anterior end of the thorax. The thymus should be carefully scraped away, all vessels except those of the structure itself being kept intact. The following structures, including the aortic arch and the arteries arising from it, the superior caval veins, and the vagus, phrenic, and Tur THORAX. 173 sympathetic nerves, cannot be dissected in the order given below, but must be separated from one another and identified as they appear. (a) The arch of the aorta (arcus aortae). Beginning at the base of the heart, the aorta passes at first forward, and then describing a curve, in the course of which it lies slightly to left of the median plane, turns backward along the ventral surfaces of the bodies of the thoracic vertebrae. With the exception of the coronary arteries (p. 176), the first branches are the large paired vessels arising from the anterior wall. They comprise the common carotid and subclavian arteries. On the right side the carotid and subclavian arise from a short common trunk, the innominate artery (a. anonyma). The left common carotid arises immediately to the left of this vessel or from its base. The branches of the common carotid artery have already been traced, with the exception of the intra-cranial portion of the internal carotid (p. 190). The subclavian artery (a. subclavia) is the first portion of the artery of the anterior limb. It passes from its point of origin laterad to the anterior margin of the first rib, where it is replaced by the axillary artery. Near its point of origin it gives off several branches, the relations of which are subject to considerable variation. They include: (1) The vertebral artery (a. vertebralis). This vessel passes into the costotransverse foramen of the seventh cervical vertebra, * and traversing the canal formed by this and the correspond- ing foramina of the remaining cervical vertebrae, reaches the interior of the cranial cavity. Its union on the ventral surface of the medulla oblongata with its fellow of the opposite side to form the basilar artery will be seen at a later stage (p. 190). (2) The superficial cervical artery (a. cervicalis superficialis)— divided in a previous dissection (p. 135)—is a small vessel which passes forward and outward beneath the insertions of the cleidomastoideus, basioclavicularis and levator scapulae major muscles, ramifying extensively in the fat mass of the side of the neck under cover of the superior portion of the trapezius. Its ascending cervical branch lies on the lateral side of the external jugular vein. (3) The transverse artery of the neck (a. transversa colli), also divided in a previous dissection (p. 186), passes laterad around the neck of the first rib to the wall of the thorax. It passes through the loop formed by the eighth cervical and first thoracic spinal nerves. It runs dorsad, first on the medial side of the scalenus anterior, then on the medial side of the cervical portion of the serratus anterior. A strong branch passes to the inferior angle of the scapula. The artery sup- plies the serratus anterior and the rhomboidei. 1 4. ANATOMY OF THE RABBIT. (4) The a. intercostalis suprema passes backward to the internal surface of the thoracic wall, giving off the first three inter- costal arteries in the intercostal spaces, and also small branches to the oesophagus and trachea. (5) The internal mammary artery (a. mammaria interna), the (b) (d) first portion of which has been removed with the ventral wall of the thorax, passes backward to the ventral abdominal wall as the superior epigastric artery (a. epigastrica superior) anastomosing with the inferior epigastric (p. 115). The superior caval vein (v. cava superior) is formed at the base of the neck by the union of the internal and external jugular veins, the latter vessel receiving at this point the subclavian vein (v. subclavia). The right superior caval passes almost directly backward, crossing the = ventral surface of the je right subclavian artery, 2 and enters the anterior portion of the right atrium. The left vessel crosses both the left subclavian artery and the arch of the aorta, reaching the right at- rium from the dorsal surface of the heart. The vagus nerve. On the right side the nerve crosses the ventral sur- face of the subclavian artery, passing dorsad to the wall of the oeso- phagus. It gives off the recurrent nerve (n. recurrens), the latter curving around the sub- Fic. 52. Plan of the venous and lymphatic Sits ; trunks of the anterior portion of the body. clavian artery and After McClure and Silvester. ee eee : a., azygos vein; ao., aorta; c.s., left superior passing forward along caval vein; d.th., thoracic duct; j.e., j.i., and the side of the trachea j-tr., external, internal, and transverse jugular ! veins; s., left subclavian vein; tr.s., transverse to the larynx. On the © scapular vein. left side the vagus passes between the archof the aorta and the base of the heart to the ventrolateral wall of the oesophagus. The recur- rent nerve passes forward on the dorsal side of the arch. The ramus cardiacus of the vagus. Infront of the subclavian artery the nerve is at first closely associated with the vagus trunk, lying on its medial side. On the right side it passes to the dorsal surface of the subclavian, and on the left to the THE THORAX. WAS) dorsal surface of the aortic arch. It ends in the cardiac plexus (plexus cardiacus), a network of sympathetic nerves lying between the aortic arch and the pulmonary artery. (e) The phrenic nerve (n. phrenicus) is a stout cord arising chiefly from the fourth cervical spinal nerve. That of the left side crosses the ventral surface of the subclavian artery and the aortic arch, passing then along the pericardium to the diaphragm. That of the right side accompanies the thoracic portion of the inferior caval vein. ({) The sympathetic trunk. At the base of the neck the cervical portion of the svmpathetic trunk enters the inferior cervical ganglion (g. cervicale inferius). The latter lies in front of and somewhat dorsal to the subclavian artery. The first thoracic ganglion lies behind the artery and is connected with the inferior cervical by the ansa subclavia, a loop formed by two cords, one of which passes to the dorsal, the other to the ventral side of the subclavian artery. The nerves proceeding from the inferior cervical ganglion enter the cardiac plexus and the sympathetic plexuses of the sub- clavian and its branches. 3. Dissection of the heart- The character and relations of the enclosing serous sac, the peri- cardium, should first be noted. Its relation to the heart is similar to that of the peritoneum and pleura investing other visceral organs (p. 49). It comprises a parietal layer, that portion commonly known as the peri- cardium, and a visceral layer, the epicardium, which forms an immediate investment for the heart substance. The parietal layer forms a loose, capacious sac, the serous membrane being greatly strengthened by the presence of a thin layer of connective tissue which forms an external investment and is usually considered as part of the pericardium. The paired pleural cavities containing the lungs are broadly separated by a longitudinal] vertical partition, the mediastinum or mediastinal septum, the space enclosed by the latter being largely occupied by the heart and by the cavity of the pericardium. For a considerable area ventrally the pericardium is loosely applied to the thoracic wall, the intervening space, which is bounded laterally by the membrane lining the pleural cavities, being known as the anterior mediastinum. A corresponding dorsal space lying between the heart and the bodies of the thoracic vertebrae, and also bounded laterally by the pleura, is the posterior mediastinum. It is occupied by several structures, namely, the oesophagus, the thoracic aorta, the bronchi, and the pulmonary bloodvessels. The pericardium should be removed, and the external features of the heart and its great vessels examined as follows: (a) The posterior, somewhat conical, ventricular portion of the heart. The left ventricle (ventriculus sinister) may be dis- tinguished both by its position and by the more solid charac- ter of its wall. The right ventricle (ventriculus dexter) is less muscular, and the wall is readily pressed inward. The line of division is indicated on the ventral surface by a faint de- pression, the anterior longitudinal sulcus. 176 ANATOMY OF THE RABBIT. (b) The pulmonary artery (a. pulmonalis) leaves the base of the right ventricle, passing forward and to the left in a some- what spiral fashion around the aorta. On the dorsal sur- face of the latter it divides into the right and left pulmonary arteries, one for each lung. At the point of division the pulmonary artery is connected with the aorta by a short fibrous cord, the arterial ligament (lig. arteriosum), repre- senting the foetal connection of the two vessels through the ductus arteriosus (p. 44). (c) The left coronary artery (a. coronaria sinistra) passes back- ward on the ventral surface of the heart, lying in the anterior longitudinal sulcus. A corresponding right coronary artery (a. coronaria dextra) passes to the right side of the heart, lying between the right ventricle and the right atrium. The two vessels supply the substance of the heart. They are the first branches of the aorta, arising from the aortic sinuses at its base. (d) The left atrium (atrium sinistrum) is the thin-walled chamber lying to the left at the base of the heart. The pulmonary veins (venae pulmonales), several on either side, enter the left atrium, passing from the medial portions of the lungs. (e) The right atrium (atrium dextrum) resembles the left in the character of its wall. It receives the right and left superior caval and the unpaired inferior caval veins. The heart may be removed by dividing the great bloodvessels. The arch of the aorta should be removed with the heart by dividing the vessel at a point beyond the origin of the left subclavian, and then severing the carotids and subclavians at their bases. This exposes the surface for the subsequent examination of the posterior end of the trachea and its connections with the lungs. Open the right ventricle by a longitudinal incision of the ventral wall, extending the incision forward into the pulmonary artery; also both atria by transverse incisions. By washing out the cavities, the internal features of the wall, including the arrangement of the valvular structures, may be examined as follows: In the right ventricle: (a) The trabeculae carneae; muscular ridges of the internal sur- face of the wall. : (b) The tricuspid valve (valvula_ tricuspidalis). The thin membranous flaps composing the valve enclose the atrio- ventricular aperture, and project into the cavity of the ventricle. Their margins, which are otherwise free, are con- nected by slender fibrous cords, the chordae tendineae, with the papillary muscles (mm. papillares), the latter being thick muscular projections, of somewhat conical shape, arising from the opposite walls. (c) The semilunar valves (valvulae semilunares) of the pulmonary artery are three extremely thin folds guarding the entrance to the vessel from the right ventricle. Two of the valves THE THORAX. 77 are usually to be found, the third being destroyed on opening the vessel. In the atria: (a) The respective positions of the pulmonary and systemic veins at their points of entrance. (b) The complete separation of the two chambers. In the par- tition separating them may be seen a thin fibrous portion denoting the position of the embryonic foramen ovale. Open the left ventricle by a ventral longitudinal incision, cutting well through the tip of the ventricle and extending the incision across the pulmonary: artery and into the aorta.. On account of the great thick- ness of the wall the internal structure is not so easily examined as in the right ventricle. (a) The bicuspid valve (valvula bicuspidalis) is similar in general structure to the tricuspid valve of the right ventricle. (b) The semilunar valves of the aorta are three in number, and are similar to those of the pulmonary artery. 4. Examination of the lungs and their connections. The removal of the ventral wall of the thorax opens the pleural cavities by taking away a considerable portion of the costal pleura, which is adherent to the internal surfaces of the ribs. The chief features may be made out as follows: (a) Each pleural cavity (cavum pleurae) is a closed serous sac, the lining membrane, or pleura, being distributed over the costal surface as the costal pleura, partly over the anterior surface of the diaphragm as the diaphragmatic pleura, and over the surface of the lung as the pulmonary pleura. Pos- teriorly the pulmonary pleura passes from the medial margin of the left lung to the medial lobule of the right and thence backward to the diaphragm, forming a broad sheet of attach- ment, the pulmonary ligament (lig. pulmonale). (b) The lungs (pulmones) are paired expansible structures, the surfaces of which are free, except posteriorly, where they are attached to the diaphragm through the pulmonary liga- ment, and medially where they are connected with the pulmonary bloodvessels and the respiratory tubes. (c) Each lung is divided into three portions, the superior, middle, and inferior lobes. On account of the smaller size of the left lung, the superior lobe is imperfectly developed. On the right side the inferior lobe is divided into two portions, the medial and lateral lobules, the inferior caval vein passing between them. (d) The trachea divides at its posterior end into two portions, the right and left bronchi, one for each lung. Each bronchus is _ again divided into smaller portions, the bronchial rami, which penetrate the substance of the organ and redivide into smaller tubes. ANATOMY OF THE RABBIT. (e) The branches of the pulmonary artery and the pulmonary (f) The lun care being veins may be traced for a short distance on the medial por- tion, or hilus, of each organ. The vagus nerve passes to the dorsal side of the bronchus, giving off a number of branches, which accompany the bronchus to the lung. gs may be removed, together with a portion of the trachea, taken to leave the vagus nerves in place. The lungs may then be examined to better advantage, and the surface also prepared for the next dissection. 5. The following structures may now be traced on the dorsal wall of the thorax: (a) (b) (c) (d Se (e) (g) The oesophagus. It traverses the thorax in a median posi- tion, entering the diaphragm at the hiatus oesophageus. The vagus nerves. The right and left nerves pass backward along the lateral walls of the oesophagus, and are connected with one another through nerve plexuses lying on its dorsal and ventral surfaces. The left cord is that appearing at the posterior end of the oesophagus in a ventral position and passing to the ventral surface of the stomach. The right cord occupies a corresponding dorsal position and passes to the dorsal surface of the stomach (p. 119). The thoracic aorta (aorta thoracalis) passes backward on the ventral surface of the vertebral column, leaving the thorax through the hiatus aorticus, the latter being the aperture enclosed by the crura of the diaphragm. Its branches in the thorax are the intercostal arteries (aa. intercostales), beginning with the fourth, which are given off metamerically in the intercostal spaces, and pass laterad to the thoracic wall. The thoracic portions of the sympathetic trunks lie on the lateral surfaces of the bodies of the thoracic vertebrae. The posterior ganglia give origin to the splanchnic nerve, the latter passing backward into the abdominal cavity (p. 118). The levatores costarum; a series of small muscles arising from the transverse processes of the vertebrae and the heads of the ribs and inserted on the anterior margins of the next succeeding ribs. The intercostal nerves (nn. intercostales) accompany the intercostal arteries to the lateral wall of the thorax. The azygos vein (v. azygos) is a small, asymmetrical, venous trunk lying to the right of the dorsal surface of the aorta. It receives the majority of the intercostal veins, which accompany the corresponding arteries and nerves, the tributaries extending backward to the first lumbar veins. It opens forward into the right superior caval. The more anterior intercostal veins are tributaries of the right and left supreme intercostal veins which open into the corresponding superior cavals. THE THORAX. 179 6. The diaphragm (diaphragma) is a muscular and tendinous sheet forming the posterior wall of the thorax and separating the pleural cavities from the peritoneal cavity. It is somewhat dome-shaped in the relaxed condition, but in contraction it becomes flattened in such a way that the space occupied by the lungs is considerably increased, while the liver and related structures of the abdominal cavity are dis- placed backward. As a muscle the diaphragm arises in three portions. The first, or lumbar portion, consists of two muscular and fibrous cords, the crura, the right much larger and stronger than the left, arising from the anterior spinous processes of the first three lumbar vertebrae. ~The second, or costal portion, arises from the internal surfaces of the posterior ribs. The third, or sternal portion, arises from the xiphoid process of the sternum. Its insertion is represented by its own tendinous central portion, or centrum tendineum, although the latter is virtually attached forwards to the lungs and pericardium through the broad pulmonary ligament. The centrum tendineum is shaped somewhat lke a trefoil, the fibres of the costal and sternal portions radiating outward from its margin. The following may be made out on the posterior surface: (a) The cut margins of the falciform, coronary, and left triangular ligaments. (b) The hiatus aorticus, an aperture enclosed by the two crura and serving for the transmission of the aorta. (c) The hiatus oesophageus, ventral to the foregoing, and serving for the passage of the oesophagus. (d) The foramen venae cavae, situated slightly to the right and serving for the transmission of the vena cava inferior. (e) The superior phrenic arteries (aa. phrenicae superiores) arise from the aorta at about the point of origin of the eleventh intercostals or from one of the latter, and enter the crura. The inferior phrenic arteries are minute branches arising at the base of the coeliac. ({) The inferior phrenic veins (vv. phrenicae inferiores), one on either side of the foramen venae cavae, at which point they enter the inferior cava. The superior phrenic veins pass forward from the diaphragm, opening into the superior cavals. € 180 ANATOMY OF THE RABBIT. XII. THE VERTEBRAL AND OCCIPITAL MUSCULATURE. Dissection on the dorsal surface of the body from the occiput back- ward; also on the lateral and ventral surfaces of the neck. The serratus posterior muscle lies on the dorsolateral surface of the thorax. It arises from the ligamentum nuchae and from the lumbo- dorsal fascia back to the last rib, and is inserted on the lateral surfaces of the eight posterior ribs. The splenius muscle is a somewhat triangular sheet arising from the ligamentum nuchae and inserted on the supraoccipital and mastoid portions of the skull, extending also to the transverse process of the atlas. These two muscles should be divided, the serratus posterior being removed from the surface. 1. The long muscles of the vertebral column. Apart from the iliopsoas, psoas minor, and quadratus lumborum— muscles of appendicular insertion lying on the ventral surface of the Vertebral column—the vertebral musculature comprises chiefly modified segmental muscles lying on the dorsal surface, for the most part in the area enclosed by the spinous and transverse processes of the vertebrae. They include the sacrospinalis, semispinalis, and intertransversarii. Their insertions are extended in part laterad to the ribs. In the cervical region they are represented by short muscles, separated for the most part from the thoracic and lumbar portions, and arising by accessory bundles from the anterior ribs, the corresponding thoracic, and the posterior cervical vertebrae. In the cervical region the muscles are easily separated from one another, but in the posterior part of the body it is necessary to dissect away the tough investment of lumbodorsal fascia which covers them. (a) The sacrospinalis. Origin: Crest ‘of the ilium and medial surface of the iliac wing; mammillary processes of the six posterior lumbar vertebrae; investing lumbodorsal fascia. This muscle is the largest and strongest muscle of the body. It extends forward over the surfaces of the ribs. Its medial border is separated from the middle line by a space of con- siderable width, in which the semispinalis and multifidus muscles are accommodated. In the lumbar region it is inserted in a continuous mass on the long transverse processes of the vertebrae and in the interspaces. In the thoracic region the muscle divides into two portions, namely, a slender lateral portion, the iliocostalis, and a thick medial portion, the longissimus. The latter receives in the posterior portion of the thorax strong accessory bundles from the semt- spinalis muscle on its medial side, the two muscles being inseparable at this point. 14 THE VERTEBRAL MUSCULATURE. 181 The iliocostalis is inserted laterally on the ribs. It receives medially from the ribs a number of accessory bundles, which are inserted forwards to the seventh cervical vertebra. The longissimus is inserted by broad fleshy bands on the posterior margins of the ribs, medial to the accessory origins of the iliocostalis, this portion of the muscle forming the longissimus dorsi. Continuing to the neck it is inserted on the trans- verse processes of the four posterior cervical vertebrae, medial to the origin of the cervical portion of the serratus anterior, but a number of accessory slips carry the insertion forward to the transverse process of the atlas. This portion is the longissimus cervicis. A separate band of fibres arising chiefly from the transverse processes of the first three thoracic vertebrae joins the medial, ventral portion of the splenius, and forms the longissimus capitis. (b) The semispinalis and multifidus. The band of muscle lying between the longissimus and the middle line, is composed of partly fused slips, arising for the most part by very long tendons from the mammillary and transverse processes, and inserted forwards on the spinous processes. It is divisible into two portions, which are superficially separated by a constricted area lying at the level of the last thoracic vertebra, this being also the point where the muscle is fused with the longissimus. The anterior portion, the semispinalis dorsi, is inserted by a series of fleshy slips on the spinous process of more anterior thoracic vertebra, but extends to the spinous process of the third or fourth cervical vertebra. The posterior portion, the multifidus, increases in size backward to the sacrum, where it is continuous with the abductor caudae posterior. An almost separate muscle, the semispinalis capitis, arises from the articular processes of the five posterior cervical and the transverse processes of the first four thoracic vertebrae. It is inserted on the,lateral surface of the external occipital protuberance. The more posterior and medial portion of the muscle is composed of separate slips arising in common with the longissimus capitis, two closely applied slips, how- ever, at the free margin of the muscle, arising from the semispinalis dorsi and the longissimus. A second muscle, the semispinalis cervicis, is covered by the (c) foregoing one. It arises from the articular processes of the posterior cervical and first thoracic vertebrae, and is inserted on the spinous processes of the cervical vertebrae, chiefly on that of the epistropheus. The intertransversaril are short muscles connecting the lateral portions of the vertebrae. They are distinguishable in part by their darker coloration. They increase in size backwards, being most conspicuous in the lumbar region, 182 ANATOMY OF THE RABBIT. where they form thick muscular pads interposed between the mammiullary and accessory process. The last slip is attached to the crest of the ilium. 2. The following muscles constitute an occipital group, composed of short muscles arising from the atlas and axis and inserted on the atlas and the occipital portion of the skull. (a) (b) (c) (d) (e) (f) The rectus capitis posterior superficialis. Origin: Spinous process of the atlas. Insertion: External occipital pro- tuberance. The obliquus capitis superior. Origin: Transverse process of the atlas. Insertion: Lateral surface of the occipital pro- tuberance. The foregoing muscles should be divided. The rectus capitis posterior minor. Origin: Posterior tubercle of the atlas. Insertion: External occipital protuberance. The rectus capitis posterior major. Origin: Spinous process of the epistropheus. Insertion: Laterally on the supra- occipital bone. The obliquus capitis inferior. Origin: Spinous process of the epistropheus. Insertion: Dorsal surface of the transverse process of the atlas. The rectus capitis lateralis. Origin: In common with the obliquus capitis superior, which covers it. Insertion: Base of the jugular process of the occipital. 3. Muscles of the lateral and ventral surfaces of the neck. (a) (b) (c) (e) The scalenus anterior. Origin: Transverse processes of the four posterior cervical vertebrae. Insertion: First rib. The scalenus medius. Origin: Transverse process of the fifth cervical vertebra. Insertion: Lateral surfaces of the third to fifth ribs. The scalenus posterior. Origin: Transverse processes of the fourth to sixth cervical vertebrae. Insertion: First rib. The scalenus medialis. Origin: The sternum at the point of attachment of the first rib. Insertion: Middle of the ventral portion of the first rib. The foregoing muscles, comprising the scalenus group, are destroyed by the removal of the ventral thoracic wall (p. 172). The longus colli. Origin: Bodies of the first six thoracic vertebrae. Insertion: The muscle passes forward on the ventral surface of the bodies of the vertebrae, giving off insertion fibres, and also receiving strands of origin. It terminates on the anterior tubercle of the atlas. (f) The longus capitis is partly fused with the foregoing muscle, but its origin is in a more lateral position from the trans- verse processes of the first six cervical vertebrae. Insertion: Sphenooccipital synchondrosis. (g) (h) THE VERTEBRAL MUSCULATURE. 183 The longus atlantis. Origin: Lateral to the longus capitis, from the transverse processes of the third to sixth cervical vertebrae. Insertion: Transverse process of the atlas. The longus capitis should be divided near its insertion. The rectus capitis anterior. Medial portion of the ventral surface of the transverse process of the atlas. Insertion: Basioccipital bone. The caudal musculature in the rabbit comprises, in addition to the posterior extension of the cutaneus maximus, the following axial muscles: (a) The extensor caudae medialis. It lies inthe furrow between the spinous and articular processes of the sacral and caudal vertebrae, and is inserted on the transverse processes and dorsal surfaces of the caudal vertebrae. (b) The abductor caudae posterior arises as a continuation of the multifidus in the groove between the articular and transverse process and is inserted on succeeding vertebrae. (c) The abductor caudae anterior. Origin: Ischialspine. Insertion: Lateral surface of the sacrum and the transverse processes of the caudal vertebrae. (d) The flexor caudae, Origin: Ventral surface of the sacrum and anterior caudal vertebrae. Insertion: Ventral surfaces of succeeding vertebrae. 184 ANATOMY OF THE RABBIT. XII.. THE CENTRAL, NERVOUS] SMSTEM. 1. The spinal cord and nerve roots. To expose the whole cord or a portion of it from the dorsal surface, the muscles should be removed on both sides of the vertebral arches and the latter broken away with the bone forceps. The following features may be made out according to the extent of exposure: (a) The spinal cord (medulla spinalis) is a thick cylindrical white cord traversing the vertebral column in the vertebral canal. It is of almost uniform diameter, but exhibits two slight enlargements, one in the cervical, the other in the lumbar region. At about the middle of the sacrum it contracts to a slender filament, the filum terminale, which may be traced backward to the middle of the tail. (b) The enclosing membranes or meninges of the cord are three in number, of which two are readily identified. Lining the internal surface of the bone is a thick fibrous investment, the dura mater; on the surface of the nervous matter a thin vascular membrane, the pia mater. The dura and pia are connected by a loose web of connective (c) tissue, the arachnoidea. The regional distribution of the nerve roots—eight cervical, twelve dorsal, seven lumbar, four sacral, and six caudal. The cervical nerves are numbered from the vertebrae lying behind the intervertebral foramina from which they proceed, the remaining nerves from the vertebrae lying in front of the intervertebral foramina. The nerve transmitted by the intervertebral foramen between the seventh cervical and first thoracic vertebrae is described as the eighth cervical. (d)gThe origin and primary divisions of the nerve roots may be worked out by removing carefully the lateral portions of the arches of one or two vertebrae. The posterior, or sensory root (radix posterior), arises from the dorsolateral surface of cord, and expands immediately into a spinal ganglion (g. spinale). The more slender anterior, or motor root (radix anterior), arises by a number of filaments from the ventro- lateral surface. The combined roots give origin to the posterior, anterior, and communicating rami. The posterior ramus is an inconspicuous branch (except in the first two cervicals) passing to the dorsal musculature and skin. The anterior ramus is the chief portion of the spinal nerve, the successive rami appearing as the components of the cervical and lumbosacral plexuses or as individual spinal nerves. The ramus communicans is a slender filament which passes ventrad to join the sympathetic trunk. THE CENTRAL NERvouS SYSTEM. 185 2. A small portion of the spinal cord may be excised and examined (preferably under water) for the following: (see p. 22, fig. 18). (a) The division of the cord into lateral halves by two median depressions, the anterior median fissure (fissura mediana anterior) and the posterior median sulcus (sulcus medianus posterior). (b) The division of each half into three columns by shallow grooves, the anterior and posterior lateral sulci. The grooves are marked by the attachments of the anterior and posterior nerve roots. The three columns of each half of the cord are the anterior, lateral, and posterior funiculi. (c) On the cut surface the white substance (substantia alba) is seen to form a peripheral investment enclosing the grey sub- stance (substantia grisea) of the centre of the cord. The grey portion is somewhat H-shaped in section, each half being composed of anterior larger and posterior smaller masses, known in section as the horns of the grey matter, or as complete structures, the anterior and posterior grey columns. In the median plane, but nearer the dorsal than the ventral surface, is the minute central canal (canalis centralis), the cavity of the spinal cord. 3. The brain may be exposed by breaking away the supraorbital processes of the frontal bone and then removing the roof of the skull with bone forceps. In order to clear the brain and the roots of the cerebral nerves, it 1s necessary to remove the entire lateral wall of the skull on both sides. The chief part of the operation consists in removing the temporal portion of the skull by successive steps, exposing first the flocculus of the cerebellum, a small stalked body which is almost com- pletely enclosed by the dorsal portion of the petrosal. The entire petro- tympanic bone is easily detached, and if removed en masse the flocculus and probably also the roots of the facial and acoustic nerves will be destroyed. The arches of the first three or four cervical vertebrae should be removed, unless the anterior portion of the cord has been exposed in the previous dissection. The spinal cord may be divided at about the level of the third ver- tebra. The brain should then be raised very gently from the ventral wall of the skull and the nerve roots divided with a small sharp knife or with fine scissors. This operation requires considerable care, since the nerves are strongly attached at their points of exit from the cranial wall, but very lightly attached to the brain, so that they are in danger of being torn away. The anterior end of the brain may be freed by cutting across the small anterior expansions formed by the olfactory buibs. Portions of the dura mater removed with the brain may be cut away with scissors. It is adherent chiefly along two lines: .one representing the longitudinal fissure between the cerebral hemispheres, the other the 186 tentorium ANATOMY OF THE RABBIT. cerebelli. These parts contain the superior sagittal and transverse venous sinuses. On the ventral surface of the brain as removed appear the basilar and internal carotid arteries and their branches. These vessels should be kept intact for examination at a later stage. For the primary divisions of the brain see p. 34. 4. The superficial features may be identified as follows: IN THE PROSENCEPHALON: (a) (b) (c) (d) (e) The greatly enlarged cerebral hemisphere (hemisphaerium cerebri) forms with its fellow of the opposite side the largest portion of the brain. The two structures are separated by the longitudinal cerebral fissure, but are connected with one another by the commissures indicated below. The olfactory bulb (bulbus olfactorius) is a small expansion lying at the anterior end of each hemisphere. Its cut end represents the point of origin of the first cranial or olfactory nerve, the various divisions of which may be found on the skull, where they may be traced into the perforations of the cribriform plate. The olfactory bulb is the anterior portion of the olfactory lobe, or olfactory brain. When traced backward on the ventral surface of the brain it is seen to be replaced by a white band of fibres, the latter forming the olfactory: tract (tractus olfactorius). The tract ends posteriorly in an expanded portion of the brain, which, from its shape, is described as the lobus piriformis. The olfactory brain as thus defined is separated superficially from the remaining portions of the cerebral hemisphere by a longitudinal furrow, the limbic fissure (fissura limbica), which passes along its lateral margin. The anterior portion of the furrow, known as the anterior rhinal fissure, separates the olfactory tract laterally from the narrow anterior’ portion of the cerebral hemisphere. The corresponding posterior portion of the fur- row, the posterior rhinal fissure, separates the lobus piriformis from the posterior enlarged portion of the hemisphere. The slight angle formed at the junction of the anterior and posterior rhinal fissures is the point of origin of a faint depression extending upward on the cerebral hemisphere. It represents a rudimentary, lateral cerebral (Sylvian) fissure. The corpus callosum is a broad white commissural band passing transversely from one hemisphere to the other. Its median portion may be exposed by pressing apart the medial margins of the hemispheres at the longitudinal cerebral fissure. The pineal body (corpus pineale) is a small somewhat conical structure lying between the dorsal posterior tips of the cerebral hemispheres, and connected by a hollow stalk (f) (g) (h) (j) (k) THE CENTRAL NERVOUS SYSTEM. 187 with the unpaired portion of the brain (the thalamencephalon) lying below it. The connection is concealed by a mass of pigmented vascular tissue, the beginning of the chorioid plexus of the third ventricle, and usually, also by a small portion of the dura mater containing part of the sagittal venous sinus. The latter may be carefully detached. By pressing apart the tips of the hemispheres and removing the pineal body the dorsal surface of the thalamencephalon will be sufficiently exposed to make out the following features: The slit-like aperture appearing in the middle line after the removal of the pineal body represents the dorsal portion of the third ventricle (ventriculus tertius) (Fig. 53), the roof of which is formed anteriorly by the superficial portion of the chorioid plexus (plexus chorioideus), the latter entering the ventricle at this point. The lateral margins of the aperture are largely formed by two minute spindle- shaped masses,one on either side, the habenulae. Their posterior ends are united by a slender transverse band, the habenular commissure (com- missura habenularum). The fibres of the commissure are faintly trace- able forward, where they form a pair of thin whitish filaments (medullary striae). The posterior commissure (commis- sura posterior) crosses the posterior portion of the roof immediately behind and below the habenular commissure. Mie wallssot sthethirdsventrcle-are ..) 22) sees toe formed by the thalami. They are of the thalamencephalon as x viewed from the dorsal sur- broadly connected by the inter- face, after removal of the mediate mass (massa intermedia), or pineal body: a., anterior thalamic tubercle; che, middle commissure, which may be habenular commissure; c.p., e 7 49 posterior commissure; Cisse seen from the dorsal surface crossing | superior colliculus (of mesence- the cavity phalon); g.l. and g.m., lateral and medial geniculate bodies; : h., habenula; m.i.,massa inter- Onveither side ot te middie lime, O10 7 aa: o. celvinse oid of the habenulae, the dorsal portion — ventricle. of the thalamus forms a low, some- what oval projection, the pulvinar or posterior tubercle. The anterior tubercle of the thalamus is a faint elevation of very small dimensions lying in the angle enclosed between the pulvinar and the anterior portion of the aperture of the third ventricle. The parts of the metathalamus are distinguishable externally as two rounded projections of the lateral surfaces of the thalamencephalon. One of them, the lateral geniculate body (corpus geniculatum laterale), lies to the lateral side of the pulvinar, anal is only separated from it by a faint depres- 188 (1) (m1) (0) (p) ANATOMY OF THE RABBIv. sion of the surface. The medial geniculate body occupies a more posterior and medial position. The optic tract (tractus opticus) connects the geniculate bodies, especially the lateral one, with the ventral surface of the brain, ending in the optic chiasma.. (m) On the ventral surface, the optic chiasma (chiasma opticum), forms a conspicuous elevation, the posterior portion of which is traceable into the optic tracts, the anterior portion into the bases of the second cranial, or optic nerves. The hypophysis, or pituitary body, lies immediately behind the optic chiasma. On account of its enclosure by the walls of the hypophysial fossa, and also its slight attachment to the brain, the hypophysis is commonly detached in removing-the brain from the skull, in which case a slit- like aperture, representing the ventral portion of the third ventricle is exposed. The tuber cinereum is a small elevation of grey matter appearing on the ventral surface after the removal of the hypophysis. It is the base of attachment of the infundi- bulum, the latter being the slender extension of the brain downward toward the hypophysis. The mammillary body (corpus mammillare) forms a con- spicuous rounded elevation, lying at the posterior end of the tuber cinereum. The structure is single, but there is an indication of lateral lobes. IN THE MESENCEPHALON: (a) (b) (c) The dorsal surface is marked by four elevations, the corpora quadrigemina. The anterior pair, distinguished as the colliculi superiores, are much larger than the posterior pair, the colliculi inferiores. The ventral surface is occupied by a pair of divergent cords, the cerebral peduncles (pedunculi cerebri), separated by a faint median depression, the interpeduncular fossa (fossa interpeduncularis). The third cranial, or oculomotor nerve (n. oculomotorius), arises from the ventral surface of the cerebral peduncle. IN THE RHOMBENCEPHALON: (a) (b) The cerebellum forms an arch over the posterior portion of the brain, and is supported by stout pillars from its sides. In addition to the ridges of its surface the whole structure is divided into several lobes, including the median portion or vermis, the lateral hemispheres, and the stalked bodies, or flocculi, arising from the hemispheres. The anterior medullary velum (velum medullare anterius) is the thin membrane underlying the anterior portion of the cerebellum and connecting the latter with the inferior colliculi. It forms a small anterior portion of the roof of the fourth ventricle. en (c) (d) (e) On the ventral surface (Fig. 54), (f) (g) (h) (1) THE CENTRAL NERVOUS SYSTEM. 189 The fourth cranial, or trochlear nerve (n. trochlearis), arises from the anterior medullary velum. The first portion of the nerve is usually to be found on the lateral surface of the cerebral peduncle. The posterior medullary velum (velum medullare posterius) underlies the posterior margin of the cerebellum, and extends backward over the triangular space enclosed by the walls of the fourth ventricle. The structure forms a chorioid plexus, similar in character to that of the third ventricle but much less extensive. It is commonly torn away in the preparation of the brain, in which case the interior of the fourth ventricle is exposed. the pons forms a broad com- missural band extending trans- versely across the brain and upward into the supports of the cerebellum. its) *suiriaces 1s divided into two parts by a median depression, the sulcus hasilaris, occupied by the basilar artery. The anterior median fissure of the spinal cord ends at the posterior margin of the pons in a faint depression, the fora-. men caecum. The anterior funiculus of the cord is largely replaced forwards by the pyramid (pyramis), a narrow band extending forward on either side of the middle line to the posterior margin of Fic. 54. The rhombencephalon. the pons Ventral view (the cerebellum not ‘ figured). The trapezoid body (corpus trapezoideum) hes in the angle formed by the lateral margin of the pyramid with the pos- terior border of the pons. The fifth cranial, or trigeminal c.t., trapezoid body; f.c., foramen caecum; fl.c., cervical flexure; f.m.a., anterior median fissure; De pons: p.c., cerebral peduncle (mesencep- halon); py., pyramid. III, oculomotor nerve; IV, troch- lear; V!, portio major of the trige- minus; V2, portio minor; VI, abducens; VII, facial; VIII, acoustic: IX-XI, glossopharyngeal, vagus, and spinal accessory group; XII, hypo- oe 3 : z ENS (n. trigeminus), HEI ES by glossal; CI, first cervical spinal. two roots, a larger sensory root, the portio major, and a smaller motor root, the portio minor. The two parts appear at the lateral border of the pons. The portio major is the common trunk of the ophthalmic, maxillary and mandibular nerves, the portio minor joining the mandibular, so that the latter becomes a mixed nerve. The cut end of the portio major may be identified on the cranial wall and traced forward in the semilunar ganglion, the latter lying in a depression at the anterior ventral end of the petrosal bone. 190 ( ()) (k) (1) m1) ANATOMY OF THE RABBIT. The sixth cranial, or abducent nerve (n. abducens), is a slender cord arising at the anterior end of the pyramid. The seventh cranial, or facial nerve (n. facialis), and the eighth, or acoustic nerve (n. acusticus), arise from the lateral margin of the trapezoid body. The two nerves are closely associated, the former being slightly anterior in position. Its chief portion is the motor root. In addition the nerve receives a sensory filament, the portio intermedia or intermediate nerve. The glossopharyngeus, vagus, and accessorius arise by several roots arranged in a linear series along the lateral margin of the medulla. The trunk of the accessorius extends back- ward on the spinal cord, its roots, about ten in number, arising as far back as the fifth cervical spinal nerve. The twelfth cranial, or hypoglossal nerve (n. hypoglossus), arises by several roots from the ventral surface of the medulla and at the lateral margin of the pyramid, its point of origin corresponding to that of the ventral root of a spinal nerve. 4. The arteries of the brain may be traced on its ventral surface as follows: (a) (b) (c) (d) (e) The basilar artery (a. basilaris) is a median trunk formed on the ventral surface of the medulla oblongata by the union of the vertebral arteries, the latter here represented by their cut ends. The inferior cerebellar artery (a. cerebelli inferior) is the largest of several transverse vessels arising from the basilar and passing laterad to the cerebellum. The posterior cerebral artery (a. cerebri posterior) is formed on either side at the level of the anterior margin of the pons by the division of the basilar. It passes to the posterior portion of the cerebral hemisphere. The superior cerebellar artery (a. cerebelli superior) is a branch of the posterior cerebral, passing to the anterior portion of the cerebellum. The cut end of the internal carotid artery lies on either side of the tuber cenereum. It is connected backwards with the posterior cerebral. The middle cerebral artery (a. cerebri media) is given off from the internal carotid, passing to the middle portion of the hemispheres. The anterior cerebral artery (a. cerebri anterior), the con- tinuation of the carotid, passes to the anterior portion of the ventral surface and the olfactory bulb. The anterior cerebral is connected with that of the other side, a complete anastomotic loop being formed around the hypothalamus by combined branches of the vertebral and carotid arteries. This is the circle of Willis. — THE CENTRAL NERVOUS SYSTEM. 191 5. By dividing the supports of the cerebellum on either side, the entire structure may be removed and the surface exposed, as in Fig, 55, for an examination of the structures of the dorsal surface of the rhombencephalon. The posterior medullary velum is removed with the cerebellum, but a small portion of the anterior medullary velum should remain in place. (a) The fourth ventricle (ventriculus quartus) is the extensive space enclosed by the rhombencephalon. It is connected forwards with the cerebral aqueduct and backwards with the central canal of the spinal cord. Its roof is formed principally by the anterior and posterior medullary vela, the latter being attached to the cerebellum close to one another, so that they underlie it. The rhomboid fossa (fossa rhomboidea) is the shallow de- pression enclosed by the thick lateral and anterior walls and floor of the ventricle. The middle line shows a shallow depression, the posterior me- dian fissure (fissura mediana posterior), on either side of which the floor is raised into a low ridge, described as the medial eminence (eminentia medialis). The posterior end of the fossa forms with the enclosing wall the somewhat triangular figure described as the calamus scriptorius. The lateral supports of the cerebellum, now represented by their cut ends, are formed by (b) ce) fibre-bands passing into the cerebellum from adjacent ven- tral portions of the brain. In each a middle band, the brachium pontis, enters the pons ; an anterior band, the brachium conjunctivum, enters the mesencephalon, while a posterior connection is estab- BIGs455: The rhombencephalon. Dorsal view; after removal of the cerebellum: b.c., brachium conjunc- b.p., brachium pontis; cl., clava; cl.i., inferior colliculus (me- sencephalon); c.r., restiform body; e.m., medial eminence; f.c., tasciculus cuneatus; f.g., tasciculus gracilis; f.m.p., posterior median fissure of the rhomboid fossa; f.r., rhomboid fossa; s.m.p., posterior median sulcus of the medulla; s.l.p., posterior lateral sulcus; v.m.a., anterior medullary velum. tivum; lished with the medulla through the restiform body. The lateral wall of the fossa is formed by a thick ridge of nervous matter, convex on its medial side, the restiform body (corpus restiforme). Its anterior portion bears a large spherical elevation overlying the origin of the acoustic nerve. (d) 192 ANATOMY OF THE RABBIT. (e) The posterior funiculus, in passing forward from the cord, is divided into medial and lateral portions. The medial portion, the fasciculus gracilis, forms a narrow band ter- minating forwards in a club-shaped expansion, the clava. The lateral portion, the fasciculus cuneatus, passes into the restiform body. 6. The brain may be divided by a median vertical section, and one- half examined from the medial surface (Fig. 56). In addition to many of the features already made out on the surface the following may be noted: (a) The deep but extremely narrow cavity formed by the third ventricle is the first space appearing in the brain from the anterior end, the first two (paired) ventricles lying laterally in the hemispheres. They are connected with the third ventricle by a narrow transverse canal, the interventricular foramen (foramen interventriculare). Fic. 56. The brain in median section: a., anterior commissure; a.c., cerebral aque- duct; b.o., olfactory bulb; cb., cerebellum; c.cl., corpus callosum; c.f., body of the fornix; cl.i., inferior colliculus; cl.s., superior colliculus; c.m., mammillary body; c.0., optic chiasma; c.p., pineal body; fl.c., cervical fiexure; h., habenular commissure; h.c., cerebral hemisphere; hp. hippocampus; inf., infundibulum; 1.t., laminal terminalis; m.o., medulla oblongata; p., posterior commissure; p.c., chorioid plexus of the third ventricle; pd.c., cerebral peduncle; pn., pons; sp., splenium; s.pl., septum pellucidum; t.c., tuber cinereum; th., thalamus, massa intermedia; v.m.a., anterior medullary velum; v.m.p., posterior medullary velum;. v.q., fourth ventricle; v.t., third ventricle. I, olfactory nerve (origin); II, optic nerve. (b) The anterior boundary of the third ventricle is formed ventrally by the narrow partition separating the two hemi- spheres, in the dorsal portion of which is the small anterior commissure (commissura anterior). The ventral portion of the ventricle is projected toward the optic chiasma forming the recessus opticus, and into the infundibulum, forming the recessus infundibuli. (c) The mesencephalon contains no ventricular expansion, its substance being perforated only by a narrow tube, the cere- (d) (e) THE CENTRAL Nervous System. 193 bral aqueduct (aquaeductus cerebri), which connects the third with the fourth ventricle. The corpus callosum is shown in section. Anteriorly it appears to end in a somewhat club-shaped expansion, but in reality is extended as a thin sheet of fibres downward toward the lamina terminalis. Posteriorly it bends downward, forming the splenium, the latter being attached to the body of the fornix, which lies below it. The fornix consists of a pair of longitudinal fibre bands, fused for a short distance in the middle line to. form the unpaired body of the fornix (corpus fornicis). They begin in the mammiullary body, and passing upward as the columns of the fornix (columnae fornicis), meet in the body of the fornix, and afterwards diverge laterad as the pillars of the fornix (crura fornicis), ending in the hippocampus. Between the body of the fornix and the anterior portion of the corpus callosum is a thin area of the wall, the septum pelluci- dum, the lateral ventricles lying close together in this region. 7. The nervous matter covering the corpus callosum may be removed from one hemisphere by first marking out a triangular area on the dorsolateral surface; then scraping the material carefully away until the white surface of the corpus callosum is well exposed. By removing the corpus callosum the interior of the hemisphere may be examined. (a) (b) (c) (d) The lateral ventricle (ventriculus lateralis) is the extensive space enclosed by the hemisphere. It extends forward into the olfactory bulb and backward into the posterior free end of the hemisphere, passing a considerable distance behind the opening of the interventricular foramen. The excised portion of the hemisphere, forming the moder- ately thick roof and dorsolateral wall, consists largely of the peripheral grey cortex described as the pallium. The floor is formed by a greatly thickened mass of nervous matter, appearing from the interior of the ventricle in the form of two convex ridges. One of these, posterior and medial in position, is the hippocampus. The other is smaller, anterior and lateral in position, and is the corpus striatum. Between the two bodies the pigmented vascular tissue of the chorioid plexus of the lateral ventricle may be made out. On the medial wall, the thickened posterior portion forms the body of the fornix, immediately in front of which is the thinner portion of the wall, described above as the septum pellucidum. 8. The passage of the olfactory nerves to the ethmoturbinal surfaces may be traced by removing the nasal bones and working downward toward the cribriform plate, or the remaining portion of the skull con- taining the nasal region still intact may be divided vertically for a more extended examination of the nasal fossae. The features to be observed are largely those described in connection with the skeleton (pp. 83, 92). APPENDIX. THE PRESERVATION OF MATERIAL.* The method commonly used in the preparation of material for dissecting purposes consists in first embalming the body with suitable preserving fluids; afterwards filling the arteries with a colored injection mass, so that they are more easily traced. The objects served by embalming are: (1) preserving the body from decomposition for a sufficient length of time to complete the dissection; (2) keeping the body as nearly intact as possible; and (3) having the organs in good condition for study. The point last-mentioned is an important one, since much depends on having the parts of the animal in such condition that they are easily and comfortably handled, and also easily observed. The desired results are accomplished, first, by introducing the preserving fluid through the bloodvessels, instead of by immersing the animal, as was formerly the practice; secondly, by using in the preserving fluid such materials as will leave the organs in a condition as near the natural one as possible and at the same time keep them moist and flexible throughout dissection. A suitable fluid for the purpose is that recommended by Keiller} for the preservation of human subjects. The formula is as follows: FO TTA AIS sce ko tee knoe eye eee oe ee 1.5 parts CanbowiewA cid, fe eas 2) we ra et eer a wae 2: Oni tg GN Sere ob clea Genen my eran ae UREA ae a an PN ee, wat TN LOO Mine BWIA Mate ons cial eRe fn co aa tad, suk Oke tee 8600s. 100.0 A convenient method of making up the fluid, especially when em- balming the animals in numbers, is to prepare the mixture of formalin, carbolic acid and glycerin as a stock-solution, to be diluted for use by adding to each part of stock 6 parts of water. The amount required varies according to the size of the animal, the flow of the fluid in the vessels, the length of time during which the animal is left under the action *The methods here given apply only to the preservation of specimens for ordinary dissection, either singly, or in numbers for a laboratory course, with a few observations on the difficulties which are likely to be experienced. Especially in the matter of injections, the student who has acquired some knowledge of the vascular system will be able to make complete injections of the portal system and also satisfactory injections of the systemic veins, though the latter are somewhat more difficult on account of the presence of valves in the vessels. Finer vascular injections and injections of the lymphatic system according to the directions given in the anatomical textbooks may also be suggested. +Keiller, W. ‘‘On the Preservation of Subjects. etc.’’ (American Journal of Anatomy), Vol. II., 1902-3. THE PRESERVATION OF MATERIAL. ; 195 of the fluid, and the height of the pressure column. Not less than 1500 cc. should be allowed for each specimen. The apparatus needed for embalming includes a reservoir for the fluid, provided with an exit pipe to which a rubber tube may be at- tached; about 6 feet of rubber tubing to connect with the operating table; several three-way pieces to divide the stream in case several specimens are to be handled at the same time; selected rubber tubing of the size indicated below to attach the cannulae; clamps for the tubing; and, finally, glass or metal cannulae for insertion into the femoral artery. ; Glass cannulae suitable for the purpose are readily made by heating ordinary glass tubing over the Bunsen flame and drawing it out to the desired thinness. The tubing used for the purpose should be of about 6 mm. outside diameter. The cannula when completed should be about 7 cm. long; and its narrow end should have a uniform diameter of 1.5-2 mm. for about 2cm.at the tip. The tip should be touched lightly in the flame in order to round the margin by fusion, otherwise it might damage the wall of the vessel. The rubber tubing used to connect the cannula with the main tube should be of the best quality of soft rubber, and should have an inside diameter of 4mm., i.e., of proper size to slip on and off the cannula easily, but yet to retain its hold on the latter under moderate pressure. The reservoir for holding the embalming fluid may be an aspirator or irrigator bottle, an enamel fountain, percolator or ordinary funnel. It may have a capacity of one or two quarts. The capacity, however, is immaterial, so long as the operator keeps the fluid replaced. The reservoir is suspended in such a way that it may be moved up and down within a distance of four feet above the top of the operating table. At the time of beginning the embalming process the operator should have before him the reservoir, suspended at a height of about three feet, and a column of fluid, free from air-bubbles or foreign material to the tip of the cannula. This condition must be maintained throughout the operation. If at any time the pressure falls in the apparatus sufficiently to admit air, or allow coagulated blood to run back through the cannula, there is almost certain to be trouble, not only with the specimen under treatment, but also others which come after. The column of fluid is held back until the proper time by a clamp placed on the rubber tubing. The animal is killed by administering ether or illuminating gas. It is placed on its back on the table, with the head away from the operator. The skin is first divided by a small incision on the inner side of the right thigh.* By inserting the fingers well down into the incision, the skin may be torn backward and toward the ventral middle line, and at the same time the superficial epigastric vessels will be carried with the subcutaneous tissue well out of the operator’s way. Small portions of * The embalming may be done from the common carotid artery of the neck, a vessel much larger than the femoral artery and therefore easier of manipulation. This is not recommended, however, because of the damage done to various important structures of the cervical region. 196 ANATOMY OF THE RABBIT. the inner surface of the thigh and of the abdominal wall will be exposed. The white cord representing the inguinal ligament lies in the bottom of the inguinal furrow. Appearing from beneath the ligament in this position, and passing to the surface of the thigh are the femoral nerve, artery, and vein, covered by an exceedingly thin layer of muscle be- longing to the sartorius. The three structures may be separated from one another, and the muscle pulled away at the same time, by working lengthwise along the structures with the fine forceps. The artery must be thoroughly cleared for about 3 cm. from the inguinal ligament. Care must be exercised in this operation to avoid breaking its branches or the tributaries of the vein. The artery lies in front of the vein and is distinguishable by its smaller size, flattened or collapsed condition, and by its white coloration. The vein will be found greatly distended with blood. The nerve lies in front and partly on the lateral side of the artery. When the femoral artery has been fully exposed, a ligature of coarse thread, previously moistened, may be passed around its base, close to the inguinal ligament. An ordinary single knot may be placed on the ligature, but must be left loose until the cannula is inserted. By grasping the bare edge of the artery at about 2 cm. from the ligament, the operator may make a V-shaped incision in the vessel with fine scissors. The tips of the scissors are directed toward the ligament. The incision must be clean-cut, and care must be taken not to cut more than half- way through the vessel. By taking up the little angular flap with the fine forceps, the cannula may be worked into the vessel and pushed well down into it beyond the inguinal ligament. The knot is then tightened by a gentle even pull on the ends of the thread. The knot should never be pulled very tight or doubled. At the moment when the cannula is securely fastened into the vessel, the clamp is to be removed from the connecting tube and the fluid allowed to run in. At the beginning of the process a little care in arranging the animal will be amply rewarded by convenience in dis- section. The hind limb on the side opposite the incision should be drawn backward. The front limbs should be drawn apart, so that the breast is well exposed, and held in this position by a thick cord, or, better, a stout flexible wire, passing around the backof the animal. The body should be turned slightly to the operator’s left. The animal is sufficiently embalmed in two hours. About eight animals may be kept on the table by one operator, provided he has at his disposal a sufficient number of cannulae, one for each specimen, since the first may be taken off the apparatus after the eighth has been put on. Since small difficulties frequently occur in the process, especially in placing the cannulae and in keeping them clear of obstruction, a number of points may be mentioned which indicate to the operator just how the operation is succeeding. The entrance of the cannula into the artery, in the first place, is usually accompanied by a slight rise of blood into its tip. General muscle contractions in the recently killed animal are a safe indication of uniform flow of the fluid to these and also other parts ee ts THE PRESERVATION OF MATERIAL. 197 of the body, and no clogging of the vessels need be feared. The fluid may usually be observed running in the cannula, and, of course, falling in the reservoir. Finally, there are characteristic changes in the body. The abdomen becomes greatly distended, the subcutaneous tissue swollen, the eyes protrude, and there is usually more or less frothing at the nose. Leakage, either in the area of the incision or at the nose, is sometimes a sign of too much pressure. In the former case the leakage -is frequently behind the cannula, and may be stopped by artery forceps. In the latter case there is no recourse but to confine the fluid to the nasal cavity by tying the nostrils. After the embalming process the rubber tube is disconnected from the cannula, the latter being left carefully in place. The animal is then set aside for twenty-four hours in an upright position. After this it is ready for injection. The injection mass may be made by mixing ordinary starch and water to the consistence of thin cream; then adding a finely-ground coloring material, such as vermilion or a very small quantity of carmine. Thete is some advantage in using a 5% formalin instead of water alone in making up this mass, the arteries havide afterwards a brighter appear- ance, which is doubtless due partly to better preservation and partly to the fixing of the starch in the vessels. The mass must be thoroughly strained before use, in order to avoid the presence in it of particles which are too large to go through the cannula. The injection is made with a syringe, the latter being provided with a rubber tube of the same kind as that used in the embalming process. The mass is sent in by applying a gentle, even pressure, and it is sometimes advantageous to allow the injection to run backward and forward in the tube, each time applying a little more pressure. When the vessels have been filled in this way, the tube is clamped. By drawing on one cord of the ligature the knot is loosened sufficiently to withdraw the cannula, and by keeping a finger pressed on the end of the vessel, the knot may then be drawn tight without loss of injection. It sometimes happens, despite ordinary precautions, that the cannula becomes clogged either with settled starch or with coagulated blood. In this case it may be easily removed, cleaned, and replaced. The same cannula should always be used. Material prepared according to the directions given above will keep indefinitely, provided, however, that precautions are taken to avoid con- tamination from the surface. These are especially necessary 1n view of the thick coating of hairs. It is a good plan, therefore, to sponge the animal with a preserving fluid which will penetrate the coat imme- diately, or if many specimens are being prepared, to immerse the whole animal for a moment. A suitable fluid for this purpose is formalin- alcohol, made by adding 2% of formalin to a mixture of equal parts of ordinary spirit and water. The alcohol ensures immediate penetration and assists the formalin in preservation. The fluid should be squeezed out of the coat as much as possible. The presence of a considerable quantity is not harmful, unless, after the dissection has begun, the fluid should gain access to the tissues and destroy the effect of the glycerin of the embalming fluid. 15 198 ANATOMY OF THE RABBIT. For the storage of material either before or during dissection no pre- caution is necessary except that of protecting the body from undue exposure to evaporation. A convenient plan for handling the material, however—one that is in use at the present time in the laboratories of the University of Toronto—is that of providing for each specimen a zinc- lined copper box, with sliding top, and of dimensions ample for the largest specimens—namely, 6 x 6 x 20 inches. This type of storage box was designed several years ago by Professor Ramsay Wright for the purpose, but is one which has proved useful for storing purposes in many other ways. ~ sss OPN A INDEX Abdomen, 111 Abdominal aorta, 133 cavity, 50, 116 wall, muscles of, 113 Abduction in limbs, 32 Accessory respiratory tracts, 43 See Nose. Acinous glands, 11 Acoustic meatus, external, 77, 88, 89, 111 internal, 81, 89 Adaptation, 5 Adduction in limbs, 32 Adipose tissue, 13 Afferent nerves, 22 Albinism, 14 Alveolar glands, 11 Anal aperture, 111 glands, 11, 130 Analogy, Anastomoses of intestinal vessels, 116 Anatomy defined, 3 Ankle, bones of, 108 joint, 156 Aorta, see Arteries. Aortic arches, branchial, 44 Aperture, anal, 111 ot auditory tube, 165 of larynx, 165 of mouth, 110 of nose, external, 111 internal, 82, 165 piriform, 88, 94 of thorax, inferior, superior, 74 urinogenital, 111 of uterine tube, internal, 131 of uterus, external, 132 Aponreuroses, 13 Appendicular skeleton, 32, 98 Arachnoidea, 184 Arch of aorta, 173 Arches, branchial aortic, 44 visceral, 28, 31 Arrectores pilorum, 10 Arterial ligament, 44, 176 Arteries, chief aorta, abdominal, 133 sarch of, 173 thoracic, 178 development of, dorsal, ventral, 29, 44 carotid, 160, 173 coeliac, 117, 118 diac, 133 intercostal, 133, 178 lumbar, 1338 mesenteric, inferior, 126 superior, 117, 125 pulmonary, 44, 176 renal, 127 spermatic, internal, 129, 131 subclavian, 173 See Bloodvessels. Arthrodia, 19 Articulations, nature of, 17 * of ankle, 156 of hip, knee, 155 Asymmetry, 24 in digestive tube, 40 in vascular system, 24, 45 Atria of heart, 43, 176 Auditory epithelium, 12 ossicles, 77, 90 tube, 79, 89, 165 Axial skeleton, 30, 68 in chordates, 8 Axis, basicranial, 31, 77 basifacial, 31 Axis—continued. longitudinal, of body, 23, 30 position of organ systems with respect to, » Axillary fossa, 111 lymph nodes of, 138 vessels and nerves of, 137 Ball and socket joints, 18 Basicranium, 31, 77 Biceps muscles, 20 Bilateral symmetry, 238 Bile duct, 121 3iology, 3 Bipennate muscles, 20 Bladder, urinary, 128 Bloodvessels, general structure of, 21 lining membranes of, 12 of abdomen, 133; wall, 113, 115, 174 of brain, 190 of limbs, anterior, 137, 142 posterior, 149, 153 of neck, 160, 162 of orbit, 168 of pelvis, 133 _ See Arteries, Veins. 3ody, divisions of, 110 posture of, 23, 24 pituitary, 35, 188 Bones, structure, development, types of, 14 articulations of, 17 Prain, general divisions, 34 dissection, 186 flexures of, 38 olfactory, 34, 186 Pranchiomerism, 27, 28 Bronchi, 177 Bulb, olfactory, 34, 186 Pulbourethral gland, 131, 132 Caecum, 42, 123 Capillary vessels, blood, 43 lymphatic, 46 Capsules of joints, 18 of sense organs, 31 Carpus, bones of, 102 Cartilage, structure of, 14 bones, 15; of skull, 31 Caudal vertebra, 73 Caval veins, inferior, 117, 134, 176 superior, 174, 176 Cavity, abdominal, 50, 116 of central nervous system, 34 of larynx, 166 of mouth, 42, 164 of nose, 48, 82, 92, 193 orbital, 75 pericardial, 49, 175 peritoneal, 50, 116 pleural, 49, 177 serous, 49 of skull, 79 of thorax, 74 tympanic, 89, 90, 171 Cell, structure of, 8 Central nervous system, general divisions, 34 brain, 186 spinal cord, 184 Cerebral cranium, 31 hemispheres, 34, 186 nerves, see Nervous System. Cerebellum, 36, 188 Cerebrum, 36 Cervical vertebrae, 70 Characters of animals, 5 Chondrocranium, 31 200 INDEX. Chordates, characters of, 8 Chorioid plexus, structure of, 36 of lateral ventricles, 193 of third ventricle, 187 Circulation, organs of, 26 See Vascular and Lymphatic Systems. pulmonary, 43 portal, 45 systemic, 43 Classification, purpose of, 6 \ of organ systems, 26 Clavicle, 100 Clitoris, 111, structure of, 132 Cloaca in lower vertebrates, 46 Cochlea, 171 Ccelom, 49, 116 Colon, 41, 124 Coloration of skin, etc., 13 Column, vertebral, 68 Comparative anatomy defined, 3 method of, 4 Conjugation in Protozoa, 9 Connective tissues, 12 Convergence, 5 Condyles of femur, 106 occipital, 76, 85 of tibia, 107 Cord, spermatic, 129 spinal, 34, 184 vocal, 166 Corpora quadrigemina, 188 Corpus callosum, 186, 193 Coxal bone, 103 Cranial cavity, 79 nerves, see Nervous System. Craniota, characters of, 8 Cranium, cerebral, visceral, 31 See Skeleton. Deferent ducts, 129 Dental formula, 42 Dentition, in rabbit, 42 in rodents, 7 Descent of testis, 48 Descriptive anatomy, 3 terms, 23 Diaphragm, 179 Diarthrosis, 18 Digestive tube, wall of, 10, 21, 119 Digestive system, 40 glands of, 11; anal, 130; oral, infraorbi- tal. 167; parotid, 157; submaxillary, liver, 120; pancreas, 118, 122 intestines. small, 122; large, 123 mouth, 42. 164 pharynx, 43, 164 oesophagus, 41. 164, 178; connection with stomach, 116 stomach, 41. 116 Digits, anterior limb, 103, 112 posterior limb, 109; 112 Dissection, method of, 3 of rabbit, 110 Ducts, of liver, 121 nasopalatine, 165 of oral glands, infraorbital, 167 parotid, 157; submaxillary, 159 of pancreas, 122 of prostate, 159 thoracic, 46 urinogenital, 46, 127, 129, 131 Ductus arteriosus, 44 deferens, 129 Duodenum, 41, 122 Duplicidentata, 7 Dura mater, 184 Ear, external, 111 middle, 89, 90, 171 internal, 171; capsule of, 31 Iifferent nerves, 22 Egg, fertilized, 9 Egg-laying mammals, 7 Elastic fibres of connective tissue, 12 Elbow, position of, 33, 111 Embryo, organ systems in, 29 Embryonic kidney, 47 Embryology, 3 method of, 4 ' Enarthrosis, 18 Encephalon, see Brain. Endothelia, 12 Environment, relations of animals to, 5 Epicardium, 175 Epicondyles, of femur, 106 of humerus, 100 Epidermis, 10 Epididymis, 129 Epiglottis, 165 Epiphyses of bones, 17 Epistropheus, 70 Epithelial, tissues, 9 Erect posture in man, 24 Eustachian, tube, see Auditory Tube. Extension in limbs, 32 Fye, 111; dissection, 166 Eyelids, 111 Facets, articular, of vertebra, 70, 71 Fascia, 13 Fat, 13 Femur, 105 Fertilized egg, 9 Fibres, of connective tissue, 12 muscle, 19 nerve, 22 Fibrocartilage, 14 Fibrous connective tissue, 12 Fibula, 107 Flexion in limbs, 32 Flexures of brain, 38 Flocculus of cerebellum, 188 Follicles, of hairs, 10, 113 ; lymphatic, 45; of intestine, 123; of tonsil, 165 ovarian, 1381 Foot, 112 skeleton of, 108 See Hand. Foramina of skull, chief, 78 Forearm, 111 muscles of, 140 nerves and vessels of, 142 skeleton of, 101 Fore brain, 34, 186 Formula, dental, 42 Function, relation of structure to, 4 Functional organs, 5 Gall bladder, 120 Ganglia spinal, 38, 184 sympathetic, of head, 170 prevertebral; cceliac, 117 oi inferior mesenteric, 126; superior mesenteric, 117 of trunks, cervical, 164, 175 thoracic, 178; lumbar, sacral, caudal, 134 Gastric glands, 11 General anatomy, 3 Genital organs, external, 111 female, 131 male, 128 Ginglymus, 18 Girdle, pectoral, 98 pelvic, 103 Clana ie ; ue igestive system, 7 al: 130; gastric’ 11 -ssornalS 15% 1595 167; liver, 120, pancreas, 118, 122 ANE EPID NE Glands—continued. epithelial, 10 of lymphatic system, 45 axillary, 138; inguinal, 113 mesenteric, 116 of orbit, Harderian, lacrimal, 167 of skin, 11 inguinal, 11, mammary, 7, 11, 113 suprarenal, 117 thymus, 172 thyreoid, 160 of urinogenital system, bulbourethral, prostate, 131 Gliding joint, 19 Gonads, 48 Grey substance of nervous system, 23 Gross anatomy, 3 Gubernaculum, 48, 129 Gustatory epithelium, 12; organs, 165 Hairs, structure of, 10 Hand, 111 skeleton of, 102 Hard palate, 42, 165 Hares and rabbits, distinction of, 6 Head, dissection of, 157 skeleton of, 75 Heart, 43, 175 Hemispheres of brain, 34, 186 of cerebellum, 188 Hepatic portal system, 45 See Portal Vein. Heredity, 5 Hind brain, 36 Hinge joints, 18 Hip joint, 155 Histology, 3 Homogeny, 5 Homoplasy, 5 Humerus, 100 Hyoid apparatus, 97 arch, 28 Hypophysis, 35, 188 Tleum, 123 Ilium, 104 Incisors in rodents, 7 Inguinal furrow, 111; lymph nodes of, 113 glands, 11 Insertion of muscles, 20 Integument, see Skin. Interarticular cartilages, 14 Intercellular substance, 12 Interpretation of structure, 4 Intestines, 40, 41 divisions of, 122 large, 123 lymphatics of, 116, 123 muscle layers of, 21 serous coat of, 116 small, 122 Involuntary muscles, 19, 20 Ischium, 105 Jejunum, 123 Joints, structure of, 18 ankle, 156; hip, knee, 155 Jugular veins, external, 158; internal, 160 Kidney, 127 embryonic, 47 position of, 48 Knee joint, 18, 155 position of, 33 Labyrinth of ear, 171 ethmoidal, 92 Lacrimal apparatus, Harderian and lacrimal glands, 167 nasolacrimal canal, 82, 94; duct, 166 Lacteal vessels, 116 Larynx, 165 ) INDEX. 201 Leg, 112 muscles of, 150 nerves and vessels of, 153 skeleton of, 107 Leporide, characters of, 6 Lepus, genus defined, 6 Ligaments, 13 Limbs, general position of, 32 anterior, divisions, 111 dissection of, 135 skeleton, 98 posterior, divisions, 112 dissection of, 144 skeleton, 103 Lips, 110 Tjiver, 11, 120 Long axis of body, 23, 30 Lumbar vertebrae, 71 Lumbosacral plexus, 144, 154 Lungs, 43, 177 Lymph, 9 Lymphatic system, 45 follicles of sacculus rotundus, 123 vermiform process, 124; tonsil, 165 glands, axillary, 1388; cervical, 160 inguinal, 113; intestinal, 123; mesen- teric, 116, 123 vessels of mesentery, 116 thoracic duct, 46 Mammalia, characters of, 7 Mammary glands, 7, 11, 113 nipples of, 111 Mandible, 84, 96 Mandibular arch, 28 Marsupial mammalia, 7 Meatus, acoustic, external, 77, 88, 89, 111 internal, 81, 89 Medulla oblongata, 37 Membrane bone, 15; of skull, 31 mucous, 10 tympanic, 171 Meninges of central nervous system, 184 Mesencephalon, 36, 188 Mesenchyme, 12 Mesenterial small intestine, 41, 123 Mesentery, 116, 123 structure of, 50 Mesoderm, 27 Metacarpus, 103 Metatarsus, 109 Microscopic anatomy, 3 Mid-brain, 36, 188 Middle ear, see Ear. : Mixed nerves, 22 Monotremata, 7 Morphology, 3 Morphological aspect of structure, 4 Motor nerves, 22 Mouth, cavity of, 42, 164 glands of, see Digestive System. Mucous membrane, 10 : X tunic of stomach, 119; of intestine, 10 Multicellular organisms, 9 - Muscles, structure and types of, 19 of abdominal wall, 113 of face. 158 f of limbs, anterior, 135; posterior, 144 of mastication, 161 of neck, 159, see vertebral occipital, 182 of skin, 113, 135, 157 of tongue, 162 vertebral, 180 Nasal cavity, 43, 838, 92, 193 Nasopalatine ducts, 165 Nasopharynx, 164 Neck, dissection of, 158 Nerves, structure and types of, Nervous system, general divisions, 34 central, brain, 186; spinal cord, 184 99 aa 202 INDEX. Nervous system—continued. peripheral, cranial nerves, 39 I. olfactory, 186 ; INI Koyo Htey ANY AUCH mS III. oculomotor, 169, 188 IV. trochlear, 169, 189 * V. trigeminal, 162,7169, 170, 189 x VI. abducent, 169, 190 VII. facial, 157, 170, 190 VIII. acoustic, 190 IX. glossopharyngeal, 163, 164, 1 90 X. vagus, 119, 160, 164, 174, 178, 190 XI. accessory, 164, 190 XII. hypoglossal, 160, 163, 164, 190 spinal nerves, composition of, 38 cervical, 135, 138 : lumbar, sacral, 144, 154 thoracic, 178 plexuses of, brachial, cervical, 38; lumbosacral, 144, 154 sympathetic system, 38 ganglia, coeliac, 117; mesenteric, inferior, 126; superior, 117; of head, 170, see trunks. plexuses, abdominal, 118, 126 cardiac, 175 trunks, cervical, 160, 164; lumbar, sacral, 134; thoracic, 178 Nervous tissues, 21 Nose, apertures of, external, 111; piriform, 83, 94 internal, 83, 164 cavity of, 43, 83, 92, 193 Notochord, in chordates, 8; in rabbit, 29, 30 Ochodontide, 7 Oesophagus, 41, 164, 178 Olfactory brain, 34, 186 epithelium, 11 Omenta, greater, lesser, 117 Ontogeny, 4 Orbicular muscles, 20 Orbit, 75; structures of, 166 Organs, definition of, 8 relative development of, 5 visceral, 49 Organ systems, see Systems. Origin of muscles, 20 Ossicles of ear, 77, 90 Osteocranium, 31 Osteology of rabbit, 68 Ovary, 131; position of, 49 Oviduct, 47, 131 Ovum, 9 Palate, 42, 165 Pancreas, 11, 118, 122 Parotid gland, 11, 157 Pectoral girdle, 98 Pelvic girdle, 103 Penis, 111; structure of, 129 Pericardium, 49, 175 Perichondrium, 16 Periosteum, 15 Peritoneal cavity, 50, 116 Peritoneum, general relations of, 50; parietal, 115; visceral; 116 of bladder, 128 of intestines, 116, 122, 123, 124, 125 of kidney, 127 of liver, 120 of ovary, 131 of stomach, 117 of testis, 129 Phalanges, digital, of hand, 103, 112; of foot, 09 2 a Pharynx, 43, 164 Phylogeny, 4 Physiological aspect of structure, 4 Physiology, 3 Pia mater, 184 Pigmentation of skin, etc., 13 Pineal body, 36, 186 Pituitary body, 35, 188 Placenta, 7 Placental mammals, 7 Planes of body, 23 Plantigrade foot, 7, 33 Pleura, pleural cavity, 49, 177 Plexuses, of spinal nerves, brachial, 138 cervical 138; lumbosacral, 144, 154 of sympathetic nerves, abdominal aortic, 126; cardiac, 175; cceliac, 118, hypogas- tric, 126; mesenteric, inferior, ' 126; superior, 118; renal, spermatic, 126 Pons, 38, 189 Popliteal fossa, 112 Portal system, hepatic, 45, see Portal. Vein. renal, 45 , Portal vein, 119, 121 Posture of body, in man, 24 in quadrupeds, 23, 24 Prevertebral ganglia, 38 Pronation, 33 Prone position of body, 23 Prosencephalon, 84, 186- Prostate, 131 Protozoa, cells of, 9 Pubis, 105 - Pulmonary artery, 176; circulation, 43 Quadriceps muscles, 20 Rabbit, races of, 6 Radius, 101 Rami communicantes, 38, 184 Recapitulation, law of, 4 Rectum, 123 -’ Reduced organs, 5 Regional sections, 51 Renal portal system, 45 Replacing bones, 15 Reproduction in Protozoa, 9 Reproductive organs, 46 female, 131 male, 128 Respiration, branchial, pulmonary, 44 relation of ribs to, 74; diaphragm, 179 Respiratory system, 43 ¥ accessory respiratory tracts, see Nose. lungs, 177 respiratory tracts, 160, 177 Retrogressive organs, 5 Rhombencephalon, 34, 36, 188 Ribs, 73 Rodents, characters of, 7 Roots of spinal nerves, 38, 184 Sacculus rotundus, 122 Sacral vertebre, 72 Sacrum, 72 Scapula, 98 Scrotum, 111, 128 Sebaceous glands, 11 Sections, regional, 51 Segmentation of body, 27 Seminal vesicle, 130 Sense organs, special, capsules of, 31 epithelium of, 11 See Ear, Eye, Nose. Sensory epithelia, 11 hairs, 10, 110 nerves, 22 Septum of nose, 83 of thorax, 175