THE VAGUS NERVE I N THE DOMESTIC CAT (Felis clomestica). By T. B. ST0WELL>i A.M., Ph.D. (Read before the American Philosophical Society, July 15, 1881.) 1881.] 123 [Stowell The Vagus Nerve in the Domestic Cat (Felis ilomestica) By T. B. Stouten, AM., Ph.D. (Read before the American Philosophical Society, July 15, 1881.) The idea of using the cat as the basis of anatomical study is by no means arecentone. Straus-Durckheim’s “AnatomicduChat,” Dr. B. G. Wilder’s “ Anatomical Uses of the Cat,” and other papers published by the same author since 1877, and Mivart’s recent work on “The Cat,” present the general thought with more or less directness. I am not aware, however, that any one has made a study of the nerves of the cat in their detailed distribution. Having compared the vagus nerve in man, cat, dog, horse, ox, sheep, rabbit and frog, I am satisfied that the cat (Felis dornestica) pre- sents advantages over all others as a basis for comparative study. I ac- cordingly submit the accompanying figures and text to aid students who may be disposed to investigate Comparative Neurology. The cat, dog, and rabbit were injected with plaster, as recommended by Prof. Simon II. Gage, of Cornell University, in a paper published in The American Naturalist, vol. xii, p. 717. The figures are semi-diagrammatic; they were originally drawn to a scale, natural size; for the purpose of giving prominence to certain relations, to ramuli and anastomotic filaments, such modifications have been made as seemed necessar}'; where a nerve trunk is continuous, with no distinctive characters, it is shortened, e.g., thegastro- cardiac portions of the vagus (Fig. 9). The figure of the stomach is re- duced one-lialf (Fig. 13). For the sake of simplicity no attempt has been made to reproduce plexuses or the terminal ramification of filaments. The nomenclature used is largely that advocated by Dr. B. G. Wilder, before the American Association for the Advancement of Science, at Boston, J880, in a paper entitled “A Partial Revision of the Nomenclature of the- Brain,” and in a more detailed communication published in Science, March 19, and 20, 1881, entitled “A partial Revision of Anatomical Nomencla- ture, with especial reference to that of the Brain.” The simplicity and perspicuity of the nomenclature commend it alike to the lecture-room and the laboratory. [In cases wliere it was thought that any possible doubt might arise from using the new terminology, the new words are followed by their anthropotomica equivalents] The vagus nerve (N. vagus; N, pneumogastricus; Pars vaga; Par vagum; N. ambulatorius; N. sympathicus medius; Eighth pair, pneumogastric branch, Willis; Tenth pair, Sommering and Vicq-d’Azyr) presents the following marked characters, viz :— General Characters : N. vagus has the most extensive distribution and the longest course of the cranial nerves ; in its cephalic region princi- pal rami are derived from ganglia ; it forms by its frequent and complex anastomoses with N. sympathicus numerous plexuses, hence presents in- volved physiological and pathological complications ; its terminal fila- PROC. AMER. PHILOS. SOC. XX. 111. P. PRINTED MARCH 8r 1882. Stoweil.] 124 [July 15, ments supply the muscular substance and the mucous membranes of organs ; its development in relation with the development of, notably, the heart and adjacent blood-vessels, and the stomach, renders its distribu- tion somewhat asymmetrical, necessitating special anatomical study of its dextral and sinistral relations, and giving corresponding and distinctive physiological and pathological characters ; the relation of this nerve to organic life, to the automatic and the reflex phenomena of respiration, and to the so-called “inhibitory phenomena” gives importance to its study. Special anatomical characters: N. vagus and its rami are dis- tributed to the most important viscera, at least to viscera most intimately related to the functions of organic life, e. g., digestive—pharynx, oesopha- gus, stomach, liver, pancreas, intestines ; circulatory—heart, pulmonary arteries, pulmonary veins, systemic arteries and veins in the region of the heart; respiratory—larynx, trachea, bronchi, substance of lung. Special physiological characters: N. vagus is a sensory-motor nerve, having both sensitive and motor fibres ; it controls, regulates or modifies the movements and the secretory functions of the organs to which it is distributed, and upon it depend the sensory phenomena which •characterize the respective organs. DESCRIPTION : Origin and cervical portion—X. vagus in the cat (Felis doniestica) takes its superficial origin from two regions of t,lie medulla : by 12-14 filaments from the ventral border of corpus resti- forme and the depression line between cp. restiforme and the portion of medulla next laterad (Fig. 3, 4),* in a line caudad of (posterior to) the origin-filaments of N. glosso-pharyngeus (ninth pair of cranial nerves), (Fig. 2, 4), from which nerve it is sometimes separated by a small arterial twig of A. cerebellosa inferior ; and by 4-6 filaments imme- diately ventrad in the slight depression line ventrad of oliva and cepha- lad of the origin-filaments of the spinal portion of N". accessorius (Fig. 2. L). The dorsal filaments form a somewhat curved line of superficial origin, measuring 3-4 mm. in caudo-cephalic direction, and presenting its convexity dorsad (Fig. 2. X) ; the cephalic filaments are most ventral and leave the medulla oblongata just caudad of A. cerebellosa inferior—a con- siderable branch of A. basilaris at right angles with the main trunk and * There is some difficulty in establishing satisfactorily the homologies of the medulla. There are reasons for regarding the third nerve tract irom the dor- simeson as the homologue of corpus olivarium: this is manifestly not the cp. ollvarium of Foster as given in his “ Practical Physiologyit should be noticed that the cephalic origin-filaments of N. accessorius become apparent in this depression line, while the caudal origin-fllaments appear along the depression line venirad of this tract. The elliptical area (Fig. 1, 3) laterad of ventripyra- mis (anterior pyramid) and the one still dorso-lateral have relations upon which homologies might be based, giving each one the name oliva (corpus olivarium). It is not proper in this connection to discuss homologies. I have made this allusion in apology for the indeflniteness of description of the origin-line of N. 'Jagus. Whatever homologies may be established and names assigned, the figures (Fig. 3, 4) designate the relation. 1881.] 125 [Stowell. given off 4-6 mm. cephalad of union of A A. vertebralos. These filaments unite about 1 mm. peripherad of their superficial origin into six or seven ramuli, which lie ventrad of plexus clioroideus lateralis (Fig. 2, PI. Ch.), and blend in foramen jugulare to form a single flattened nerve trunk, X. vagus. In the passage through the foramen 0 mm. peripherad of its origin. X. vagus is enclosed in common with X. accessorius (XI) in a sheath formed by a tubular prolongation of the dura mater and the arachnoid membrane, where it is also joined by X. glosso-pharyngeus (IX) ; but the sheath of the united XX. vagus (X) and accessorius (XI) may be readily dissected from that of X. glosso-pharyngeus (IX), which lies ectad and cephalad. Centrad of its foramen of exit—Foramen jugulare, (Foramen lacerum-posterius, Lacerum foramen posterius)—and 8-4 mm. peripherad of medulla oblongata, X. vagus presents a ganglionic enlarge- ment, ganglion jugulare, ganglion of the root. This ganglion is hemi- spherical in form, of a grayish color, and measures nearly 2 mm. in diam- eter ; it has relations with XX. facialis, glosso-pharyngeus, accessorius and sympathicus (Fig. 5, J). At G. jugulare, X. vagus is connected by a single twig with the adjacent petrous ganglion of X. glosso-pharyngeus (IX) the “ganglion of An- dersclv’’ (Fig. 5, Pe.); by a considerable trunk with X. accessorius (Fig. 6, 10); by ramus auricularis (Fig. 5, 2), with X. facialis (VII), from which ramus, a slender ramulus penetrates the petrous bone and joins a branch of X. facialis; a portion of the ramus continues across X. facialis to the cochlea (Fig. 5, 5), a filament from the auricular branch connects with a ganglionic plexus of X. sympathicus, entad of the gangliform plexus of X. vagus. Plexus The 5 mm. of 1ST. vagus immediately caudad of G. jugulare is involved in a somewhat intricate net-work, which seems to be allied to plexus gangliformis (Fig. 6., Px. gang.); the apposed trunks of XX. glosso-pharyngeus (IX), vagus (X), accessorius (XI) and liypo- glossus (XII), are embraced by interlacing filaments of X. sympathicus, with which nerve they sustain more or less intimate relations, through anastomotic filaments; X. glosso-pharyngeus is ectal in this group, and, together with its root-ganglion—G. Ehrenritteri, which lies upon the ectal surface of G. jugulare, but which does not seem to sustain anatomical re- lations with it—may be dissected from the ental trunk; XX. vagus and accessorius are most intimately related—their separation involving the rupture of interlacing fibre—and apparently constitute a single trunk; entad of this united trunk is X. liypoglossus. At the caudal border of this plexus X. accessorius is directed dorsad to be distributed to the muscles of the neck, and X. liypoglossus assumes ectal relations, crossing the ectal surface of X. vagus nearly at right angles, and takes its course ventrad, to the muscles of the tongue. As X. liypoglossus crosses X. vagus, it de- taches a filament to G. inferius (Fig. 5, 13). This region marks the origins of two other rami wTith whose terminal filaments X. vagus sustains inti- mate relations, XX. thyro-liyoideus and descendens noni. Fig-1- fir-2. %3. V- Hg.5. 1881.] 127 [Stowell 15 mm. caudad of G. jugulare and dorsad of the origin of A. carotidea interna, X. vagus receives a second ganglionic enlargement, ganglion inferius, ganglion of the trunk (Fig. 5,1.). This ganglion has a fusi- form outline 5-8 mm. in caudo-ceplialic diameter and 2 mm. in dorso- ventral; it is of a pinkish color; is located ectad of (superficial to) and dorso-caudad of the closely-apposed superior cervical ganglion of X. sympa- thicus, to which it is very intimately related through anastomotic filaments; its cephalic extremity is apposed to the middle of the superior cervical ganglion. G. inferius does not embrace or involve the main trunk of X. accessorius; it is however joined at its dorso-ceplialic border by a large ramus given off from X. accessorius just peripherad of Px. gangliformis (Fig. 5, 14)-, it is the superficial origin of a large ramus of X. vagus, viz., X. laryngeus superior; it communicates with X. glosso-pharyngeus (IX) (Fig. 5, 17), X. accessorius (XI) (Fig. 5, 14), X. hypoglossus (XII) (Fig. 5, 13), with the spinal nerves XX. vertebrales, in the loop which connects the first and second cervical nerves, and with X. sympathicus (entad of Px. gangliformis). In the cervical region X. vagus continues caudad from G. inferius asso- ciated with X. sympathicus in the sheath of A. carotidea primitiva. In the cephalic 20 mm. the trunk lies dorso-laterad of A. carotidea externa and A. carotidea primitiva, being concealed within the arterial sheath by the artery and by Y. jugularis interna. As the nerve approaches A. oc- cipitalis (?)* it lies laterad of A. carotidea primitiva and crosses the venter of A. occipitalis (?) at its origin; it resumes its dorso lateral relation 5-8 mm. caudad of A. occipitalis (?) until it enters the thorax. The trunk of N. vagus in its cervical region caudad of G. inferius gives otf several ramuli which anastomose with ramuli of N. sympathicus to constitute a more or less dense plexus around the trachea and oesophagus; this is especially marked in the caudal portion of the cervical region. The distinctive courses of the sinistral and the dextral nerves in the thorax require separate descriptions. The principal rami of the cervical portion ofX. vagus are Rm auricularis, X. pharyngeus, and X. laryngeus superior. Km. auricularis, is a large anastomotic branch and has its superficial origin in the dorso-ental border of G. jugulare; its course is curved dorso- laterad and cephalad, and it enters the periotic bone, follows a groove along the dorso-caudal border of the tympanic bulla, traverses the petrous por- tion of the bone and enters aqueductus Fallopii at a point 2 mm. centrad of the origin of chorda tympani; a portion of Km. auricularis continues to the opening where it meets the dorsal branch of X. facialis, to be distributed to the ear. A considerable fasciculus crosses X. facialis and maybe traced to the cochlea (Fig. 5, 5). 2 mm. peripherad of its origin Rm. auricularis receives a considerable twig from X. glosso-pliaryngeus (IX), and about * This artery, 25 mm. caudad ot foramen of exit and dorsad, or 1-3 mm. dorso- cephalad of A. thyreoidea superior, seems to be allied to A. princeps cervicis. There are some objections to this homology, but the measurements given in the text identify it beyond question. Stowell.] [July 15, the same distance, still peripherad, an anastomotic filament front X. sympa- tliicus (Fig. 5, 6). N. pharyngeus, the pharyngeal branch, takes its superficial origin from the ventro-ental surface of X. vagus just caudad of the united trunks of XX. vagus and accessorius, 7 mm. caudad of G.jugulare and entad of the point where X. hypoglossus (XII) lies ectad of X. vagus ; its origin is, therefore, involved in Px. gangliformis. A considerable accession is traceable through the plexus to the accessory branch of X. accessorius. Its course is ventrad, parallel with X. glosso-pliaryngeus (IX), and only 2-3 mm. caudad of that nerve ; it lies ectad of A. carotidea interna, and entad of V. jugularis interna and A. carotidea externa ; just peripherad of its origin it gives a twig caudad to the trunk, which may be traced to G. inferius (Fig. 5, 23). Opposite A. carotidea interna it divides into two rami (Fig. 5, 22, a, b), from which filaments are given to A. carotidea interna and to the adjacent Y. jugularis interna. From the cephalic ramus anastomotic filaments join N. glosso-pliaryngeus (IX) to form a plexus, from which filaments are distributed to the cephalic border of MM. pharyngis constrictor medius, and pharyngis constrictor superior ; others anastomose with filaments of N. sympathicus and form the pha- ryngeal plexus (Fig. 5), other filaments join X. hypoglossus (XII) in this plexus. The caudal ramus has its general course caudad ; 5 mm. peripherad of its origin, it subdivides into two ramuli, which may be designated, in view of their distribution, as the pharyngeal (Fig. 5-24), and the oesopha- geal (Fig. 5, 25) ; the pharyngeal ramulus is directed meso-dorsad and forms a loose network with the terminal filaments of the pharyngeal ramuli of X. glosso-pliaryngeus (IX)—Px. pharyngeus. The oesophageal ramulus gives filaments to MM. pharyngis constrictor medius and plia- ryngis constrictor inferior. 10-12 mm. caudad of the filaments to the mus- cles of the pharynx a considerable twig joins the cephalic ramus of X. laryngeus superior and receives an anastomotic filament from the caudal ramus of the same nerve. The oesophageal ramulus continues along the dorsum of the oesophagus caudad as far as the caudal third of the cervical portion, interlacing in the plexus around that viscus.* N. laryngeus superior, the superior laryngeal branch, is consider- ably larger than X. pharyngeus ; it takes its superficial origin from the ventral border of the middle region of G. inferius ; its course is imme- diately ventrad—occasionally it is directed caudad apposed to the main trunk and ectad of X. sympathicus, 8-10 mm., at which point it turns ventrad—and passes entad of A. carotidea primitiva, where it bifurcates into a cephalic, ental, ramus, N. laryngeus internus (Fig. 5, 7, 28) and a caudal, ectal, ramus, JY. laryngeus externus (Fig. 7. 29). N. laryngeus internus is much larger than X. laryngeus externus ; it accompanies A. laryngea superior, and with the artery perforates the hyo-thyroid membrane at the ventro-caudal border of the cephalic cornu * It sometimes occurs that the cancial ramus is detached caudad ot the cephalic ramus of N. pharyngeus : in this case it constitutes a second pharyngeal nerve ; this arrangement does not change its distribution. 129 1881.] [Stowell of Ctl. thyroidea—A. laryngea superior is given off from A. carotidea externa, just caudad of the origin of A. carotidea interna ; its course is, therefore, at right angles with A. carotidea externa, N. laryngeus externus lies ectad of the larynx; it sends a pharyn- geal ramulus cephalad and entad of A. laryngea superior (Fig. 7, 30), which is distributed to M. constrictor pharyngis inferior, anastomoses with the oesophageal ramus of N. pharyngeus (Fig. 7, 31), gives terminal fila- ments to Px. pharyngeus and the pharyngeal mucous membrane; fila- ments of this ramulus anastomose with N. crico-thyroideus and unite in plexiform relation with N. sympathicus around A. thyroidea superior ; a twig of this pharyngeal ramulus detached just dorsad of the liyo-thyroid foramen, passes ventro-caudad to terminate in M. crico-thyroideus. This ramus sends a laryngeal ramulus ventro-caudad which lies entad of M. sterno-thyroideus, to which a few filaments are distributed, and is apposed to A. laryngea inferior, which artery it accompanies to the cricoid membrane as far as the ventrimeson (Fig. 7, 32) ; from this ramulus fila- ments are given to M. crico-thyroideus. A caudal ramulus is also de- tached whose course is dorsad of M. sterno-thyroideus and parallel with its dorsal border to Cp. tliyroideum (Fig. 7, 33) ; it gives filaments to M. crico-thyroideus and anastomotic filaments to N. descendens noni, which it joins opposite the origin of A. thyroidea superior. N. laryngeus internus enters the liyo-tliyroid foramen and divides into a cephalic and a caudal offset: the cephalic offset (Fig. 7, 34) accom- panies the ental portion of A. laryngea superior, pierces M. tliyro-arytse- noideus to which numerous filaments are distributed, takes its cephalic course obliquely toward the ventrimeson and perforates the thyro-hyoid membrane 2 mm. laterad of the meson. Terminal filaments of this offset supply the epiglottis (Fig. 7, 36), the aryheno-epiglottidean folds (Id. 37), the laryngeal glands (Id. 35), and the mucous membrane of the larynx (Id. 35, a). A twig given from the cephalic offset, 2 mm. periplierad of the foramen, passes entad of the apposed artery (A. laryngea superior entalis) and joins the caudal offset 2 mm. cephalad of the caudal border of Ctl. thyroidea (Fig. 7 and 8, 39). The caudal offset lies closely apposed to the ental surface of Ctl. thyroidea (Fig. 7 and 8, 38); its first twig is sent dorso-caudad and termi- nates upon M. crico-arytaenoideus lateralis, M. arytaenoideus and M. con- strictor pharyngis inferior (Fig. 8, 40); anastomotic filaments join in plex- iform relation with its dextral homologue and with the pharyngeal and oesophageal plexuses. The principal portion of the caudal offset at its union with the twig from the cephalic offset gives off radiating filaments upon the ectal surface of the arytamoid muscles which constitute a multiple palmate plexus (Fig. 8, 41.); a twig passes dorsad of the articular facet of Ctl. cricoidea and joins N. laryngeus inferior (Fig. 8. 42). Near the union of the twig and the offset, entad of cephalic border of Ctl. cricoidea several filaments penetrate M. crico-arytsenoideus and are distributed upon the mucous membrane of the larynx. /Pl.Phar. fig: 7 Fig.^. Pg.ll (ifft 131 1881.] [Stowell. Thoracic division of N. vagus sinister: The main trunk of X. vagus sinister enters the thorax dorsad of V. innorainata sinistra at tlie union of V. subclavia and V. jugularis externa, meso-dorsad of Y. verte- bralis and laterad of A. carotidea primitiva sinistra. In the thorax cephalad of the arch of A. aorta the nerve lies between AA. carotidea and subclavia meso-dorsad of A. sternalis until it reaches a point 10-15 nun. cephalad of A. aorta, at which point it rests upon the ventral surface of A. subclavia and crosses the arch ventro-laterad of the origin of A. subclavia. Opposite the origin of A. sternalis the nerve is crossed by X. phrenicus and lies dorso-laterad of this nerve in the area between the two arteries aforenamed. As X. vagus enters the thorax, two fasciculi from the middle and caudal areas respectively of the middle cervical ganglion of X. sympatlvicus, G. thyreoideum, connect X. vagus with X. sympathicus; these commissural fasciculi are about 3 mm. apart. Between a point opposite A. sternalis and the arch of A. aorta the trunk of X. vagus sustains intimate relations with X. sympathicus, X. cardiacus magnus and X. cardiacus minor through numerous anastomotic filaments which constitute aplexiform net- work around the arteries, trachea and oesophagus in this region of the thorax—AA. subclavise, carotidese primitives and innominata. Opposite the caudal border of the arch of A. aorta a considerable fasciculus from the main trunk about 5 mm. in length joins X. laryngeus inferior. 7 mm. peripherad of this fasciculus, where X. largyneus inferior bends around the arch of A. aorta, a ramulus is given oft' whose interlacings with rami from XX. vagus and sympathicus constitute a plexiform network which is related with the cardiac plexus. In its course caudad of arch of A. aorta. X. vagus passes dorsad of the root of the left lung. Pulmonary Kami: Between a point opposite the cephalic border of A. pulmonaris and 15 mm. caudad, X. vagus gives several ramuli meso- ventrad to anastomose with terminal filaments of X. cardiacus minor and X. laryngeus inferior in the formotion of the ectal (superficial) cardiac and the ventral (anterior) pulmonary plexuses. From the same region of the main trunk filaments are directed meso-dorsad, which interlace in a dense network with filaments of offsets detached from the main trunk of the area above named, and with terminal filaments of X. cardiacus minor, and other filaments from X. sympathicus to form on the ventral aspect of the trachea just cephalad of its bifurcation, a large plexus, the ental car- diac—Px. profundus magnus—from which filaments ramify upon the bronchi and have intimate relations with the plexiform network which is formed by filaments from the offsets named and accessory offsets from thoracic ganglia of X. sympathicus around the bronchi,—dorsal pulmonary plexus. Offsets from this plexus may be traced along the air tubes into the substance of the several lobes and upon the broncho-pulmonary mu- cous membrane of the sinistral lung. The ramuli which form the dorsal pulmonary plexus are noticeably larger than those given to the ventral plexus. (Esophageal Rami : Cephalad and caudad of the pulmonary rami PROC. AMER. PHILOS. SOC. XX. 111. Q. PRINTED MARCH 8, 1882. Stowell.] 132 [July 15, numerous filaments are directed dorsad, by whose anastomoses and union with N. sympathicus is formed the oesophageal plexus which embraces the entire length of the thoracic oesophagus. Caudad of the ramuli given to the pulmonary plexus, 15-20 mm. caudad of the caudal border of the arch of A. aorta, N. vagus sinister divides into sinistral or lateral and dex- tral or mesal rami (Fig. 9, 46) which lie respectively upon the sinistral dorsum and venter of the adjacent oesophagus. The lateral ramus trends dorso-caudad, and 50-60 mm. peripherad of its origin it unites with the lateral ramus of N. vagus dexter in a median line upon the dorsum of the (esophagus, to constitute a single dorsal trunk for about 25 mm. (Fig. 9, 47) Numerous anastomotic filaments from the two rami of N. vagus sinister and the rami of N. vagus dexter interlace in the oesophageal plexus from which filaments are given to the muscular tissue and mucous mem- brane of the oesophagus. The united dorsal trunk perforates the dia- phragm and enters the abdomen as the gastric nerve. The mesal ramus of N. vagus sinister trends ventro-caudad, and 20-25 mm. peripherad of its origin is joined by its dextral homologue (Fig. 9, 48) and theke two mesal rami constitute a united ventral trunk which lies in the caudal mediastinum upon the venter of the oesophagus and perfo- rating the diaphragm near the meson, lies on the venter of the cardia (Fig. 13). A slight twig connects the two mesal rami 2 mm. peripherad of their origins. From the thoracic portion of the ventral trunk anastomotic filaments are given to its homologue in the formation of the oesophageal plexus. The thoracic portion of N. vagus dexter lies ventrad of A. sub- clavia and mesad of A. sternalis; at the caudal border of A. subclavia it bends slightly dorsad to pass mesad of V. vertebralis at its junction with V. innominata, it continues laterad of the trachea, entad of V. azygos and dorsad of the root of the right lung. As the main trunk enters the thorax it sustains intimate relations through anastomotic twigs with N. cardiacus magnus, N. cardiacus minor and the inferior cervical ganglion of N. sympathicus (Fig. 10). 15 mm. caudad of A. subclavia. a considerable ramus is directed meso-caudad and accompanies a large ramus detached entad of V. azygos ; these cardiac rami pass meso-ventrad around the base of the right pulmonary artery and to the right auricle (Px. cardiacus ectalis). Three or four ramuli are given off between A. subclavia and Y. azygos whose ramifications interlace the plexus of the trachea and oesophagus. From the 12-14 mm. of the trunk dorsad of the lung, numerous filaments are directed mesad and ventrad to join the pulmonary plexus (Fig. 9). Caudad of this point and opposite the bifurcation of its sinistral homologue the dextral nerve bifurcates into lateral and.mesal rami (Fig. 9, 49); cau- dad of the bifurcation the lateral ramus trends dorso-caudad until it joins its sinistral homologue already described. The mesal ramus gives re- current ramuli cephalad to the dextral border of the pulmonary plexus. Several other anastomotic filaments are detached from the ramus between the root of the lung and the union with its fellow which terminate in the oesophageal plexus (Fig. 9). 1881.] 133 NX. laryngei inferiores, recurrent or inferior branches of N. vagus, tracheal recurrents, have the following general characters in com- mon, viz.: their general cephalic direction; their disposition along the dorso lateral border of the trachea; the anastomotic character of their ramuli; the distribution of the terminal filaments; the sensor}7 function of the fibre. Distinctive characters: their origin; their length; their disposi- tion in the thorax; the relative number of anastomotic filaments; the num- ber of tracheo-oesophageal ramuli. Special description : N. laryngeus inferior sinister, the sinistral re- current nerve, branches from the mesal aspect of the main trunk of N. vagus, 1-3 mm. cephalad of the arch of A. aorta (Fig. 9, 45); * its course is caudad, apposed to the mesal side of the main trunk as far as the root of A. subclavia sinistra where the main trunk crosses the arch of A. aorta. Upon the ventral aspect of the arch of A. aorta, N. laryngeus inferior sep- arates from the main trunk upon the mesal side, and twisting around the concave aspect of the arch about 1 mm. sinistrad of the obliterated “ductus arteriosus,” it trends meso-dorsad, and returns cephalad along the dorso-lateral border of the trachea, between the trachea and the oeso- phagus, as a “recurrent nerve’’ (Fig. 9, 50). At ihe caudal border of the larynx N. laryngeus detaches several ectal filaments to M. crico-thyroideus (Fig. 8), passes entad of a caudal twig of A. thyroidea superior, bends dorsad around the articular facet ofCtl. cricoidea (Fig. 8) and enters the larynx as an ental nerve. A slender anastomotic twig passes ectad of the arterial twig named and may be traced dorsad of the nerve trunk until it joins a corresponding twig from the caudal division of N. laryngeus su- perior (Fig. 7, ‘29, a). Pharyngeal ramuli from the ental nerve are dis- tributed to M. constrictor pliaryngis inferior; other dorsal filaments supply M. arytsenoideus posterior and M. arytamoideus; ventral filaments supply MM. crico arytsenoideus lateralis and thyro-arytsenoideus, while terminal filaments reach the sub-glottic mucous membrane. Upon the ectal surfaces of MM. crico-arytamoideus posterior and crico-arytamoideus lateralis a multiple palmate plexus is formed by anastomotic filaments of NN. laryngeus superior and laryngeus inferior (Fig. 8, J)l). N. laryngeus inferior dexter is detached from Ihe main trunk of N. vagus, 12 mm. cephalad of the origin of A. subclavia, where the main trunk is disposed upon the ventral aspect of A. subclavia (Fig. 10); N. laryngeus dexter is immediately directed caudad over the venter of the artery, is reflected around the caudal aspect, and assumes a meso-dorsal direction to the dextral side of the trachea, and is disposed like its sinistral liomologue, with the exception of having fewer anastomotic filaments. Peripherad of the origin of N. laryngeus inferior dexter, dorsad of A. sub- clavia, ramuli are given to the deep cardiac and the posterior pulmonary plexuses; another ramulus cephalad joins its sinistral fellow', a third, the thoracic cardiac, is directed caudad by the side of the main trunk of N. vagus dexter, and terminates in the dextral bronchial plexus. As N. * An occasional origin is 8-10 mm. cephalad of cephalic border ot arch of A. aorta. [Stowell Stowell.] [July 15, Fs.io. Fig.?. Fig-13 18S1.] 135 [Stowell laryngeus inferior dexter bends around A. subclavia, just dorsad of A. sternalis, a branch is given off caudad, which, 10-12 mm. from its origin, joins N. cardiacus minor (Fig. 10), and these apposed trunks are joined 5 mm. peripherad by N. cardiacus magnus dexter, and .the trunk thus con- stituted passes dorsad of Y. cava descendens and A. innominata to the dorso-caudal border of the arch of A. aorta, where it terminates in Px. mag- nus profundus, from which filaments proceed to the ventral and dorsal coronary and the pulmonary plexuses. Tracheo-cesophageal ramuli of N. laryngeus inferior (Fig. 11, 12). General characters: these ramuli of the sinistral and dextral nerves have in common the following characters—their origin; general direction; numerous terminal filaments; the plexiform relation of these filaments; their mode of entering larynx; their distribution upon its mucous mem- brane; distribution of the dorsal filaments to oesophagus. Distinctive characters : the smaller number of ramuli from the dextral side than from the sinistral; the homologue of the first sinistral nerve is always found as a ramulus from the main trunk caudad of the origin of N. laryngeus in- ferior dexter (Fig. 12); the terminal filaments of the dextral side are less numerous than those of the corresponding nerves of the sinistral side.* Special description: opposite the cephalic border of the arch of A. aorta the first tracheal ramulus is detached (Fig. 11, l°rm.); a considerable offset is directed caudad from the origin to Px. magnus profundus; 2 mm. peripherad of origin an anastomotic filament joins N. vagus 8 mm. caudad of origin of IS. laryngeus inferior; 6 mm. peripherad of origin the ramulus bifurcates, the longer division is distributed upon the dorsum of the trachea 30 mm. cephalad of the arch of A. aorta; the shorter or caudal division sends filaments to Px. cardiacus ventralis, to Px. magnus profundus and to the bronchioli. Five mm. cephalad of the first ramulus a second is given to the venter and the sides of the trachea over that portion corresponding to the distribution of the cephalic division of the first ramus upon the dorsum. Ten mm. cephalad of the second ramulus and nearly opposite the origin of A. sternalis, the longest ramulus is detached ; this divides into three offsets, the caudal is distributed to the venter of the oesophagus, the median to the sides of the trachea, the cephalic lies just laterad of the ventrimeson and gives two considerable fasciculi, whose terminal filaments supply the walls of the trachea ; the terminal filaments of the ramulus are traceable nearly to Ctl. cricoidea. Opposite the sixth cervical vertebra the fourth ramulus is detached, whose filaments anastomose with the preceding ramulus, and supply the dorsum of the trachea and venter of adjacent oesophagus along the entire cervical region from the thorax to the larynx. The fifth tracheal ramulus takes its origin 10mm. caudad of Ctl. cricoidea * The double ramuli sometimes occur with separate origins; this apparent increase of ramuli may be regarded as a modification and not a violation of the plan. In the special description the details of measurements of a single speci- men are given. Stowell. 136 [July 15, (Fig. 8 and 11, 5°). This ramulus is largely if not exclusively oeso- phageal and joins in Px. pharyngeus; the caudal or recurrent portion is reflected caudad upon the oesophagus. Gastric nerves: Caudad of the diaphragm the dorsal gastric nerve splits into several terminal ramuli, the longest of which terminates in ganglion semi-lunare of the great solar plexus, Px. Solaris ; near the cardia numerous filaments are distributed to the cardia ; offsets supply the lesser curvature of the stomach, the plexus around A. coronaria ventriculi, and the dorsal surface of the stomach ; ramuli may be traced to the plexus around A. liepatica (Px. hepaticus), A. splenica (Px. splenicus), A. mesen- terica superior (Px. mesarseicUs). At the cardia, terminal filaments of the ventral trunk are distributed to the lesser curvature of the stomach, a few join terminal filaments of the dorsal trunk (Fig. 13), and others still maybe traced to the great solar plexus, from which ramuli enter the gastro-hepatic omentum and join the hepatic plexus. This anastomosis of the dorsal and ventral trunks in the solar plexus constitutes the “memor- able loop of Wrisberg.” SUMMARY. A. Anatomical. 1. Origin—12-14 filaments along a line ventro- laterad of Cp. restiforme, and by 4-6 filaments ventrad of oliva. 2. Foramen of exit—foramen lacerum posterius. 3. Ganglia—G. jugulare, in the proximal end of foramen of exit— G. inferius, 15mm. peripherad. 4. Relations of ganglia—G. jugulare, with NN. facialis (VII), glosso-pharyngeus (IX), accessorius (XI), sympatliicus ; G. fnferius, with NX. glosso-pharyngeus (IX), accessorius (XI), hypoglossus (XII), pharyngeus, laryngeus superior, sympathicus. 5. Px. gangliformis—the 5 mm. of trunk peripherad of G. jugu- lare ; it is formed by accessory portion of N. accessorius, anastomotic filaments between NN. vagus and accessoi'ius, filaments to N. pharyngeus, and N. sympathicus. 6. Principal rami—respective origins and general distribution : Km. auricularis, G. jugulare to N. facialis; N. pharyngeus, Px. gangli- formis to Px. pharyngeus and oesopliagams ; N. laryngeus superior, G. inferius to larynx ; N. laryngeus inferior, N. vagus near arch of A. aorta to trachea and oesophagus ; Rm. cardiaci, trunk of N. vagus proximad of base of heart to Px. cardiaci; Rm. pulmonares, trunk of N. vagus proximad of root of lungs to Px. pulmonares ; anastomotic filaments to N. sympathicus. 7. Bifurcation—dorso-laterad from roots of lungs into lateral and mesal rami. 8. Formation of nerve trunks—dorsal trunk by union of lateral rami = dorsal gastric nerve (N. gastricus dorsalis)—ventral trunk by union of mesal rami = ventral gastric nerve (N. gastricus ventralis). 9. Termination—ganglia semi-lunaria of Px. Solaris in loop of W risberg. 137 18S1.] [Stowell. R. Physiological—sensibility of mucous membrane of pharynx, larynx, trachea, bronchi, bronchioli—motion of pharynx, larynx ; reflex movements of broncho pulmonary passages, oesophagus and stomach— action upon secretions, e. g., gastric juice, biliary products, etc.*—indirect influence upon phenomena of respiration and of “inhibition.” Explanation of the Xumbeks and Abbreviations used in thi Figures. A. bas., A. basilaris ; A. cb., A. cerebralis posterior; A. cbl., A. cere- bellosa inferior; A. ver., Arteria vertebralis ; Ar. el., area elliptiea (possibly related to olivary body) ; ?, elongated, pyriform area lateral from Ar. el., whose homology is not determined ; Cb., cerebrum ; Cbl., cerebellum ; Ch., chiasma; dpy., dorsipyramis (posterior pyramid) ; Ehr., G. Ehren- ritteri ; liph., hypophysis ; I., G. inferius ; J., G. jugulare ; mtc., meta- coelia (fourth ventricle) ; O., oliva, corpus olivarium (?) ; olf., lobus olfactorius ; opt., X. opticus ; Pe., G. petrosum ; Px.cli., plexuschoroideus lateralis ; Px. phar., plexus pharyngeus ; Pn., Pons Varolii; Iif., corpus restiforme; Ypy., ventripyramis (anterior pyramid) ; II., X. opticus; III., X. motor oculi ; Y., XX. trigemini ; VI., N. abducens ; YII., X. faci- alis ; VIII., X. auditorius, Portio mollis ; IX., X. glosso-pharyngeus ; X. X. vagus ; XI., X. accessorius 5 XII., X. hypoglossus : 1, accessory filament from X. glosso-pharyngeus ; 2, Rm. auricularis ; 3, anastomotic twig from J. to Pe. ; 4, filament from origin line of IX. to 2 ; 5, ramulus from 2 to cochlea ; 6, anastomotic twig to X. sympathicus ; 7, chorda tympani ; 8, anastomotic twig from Pe. to X. ; 10, Rm. accessorius from XI. ; 11, second accession from XI. ; 12, anastomotic filaments between X. and XI. ; 13, filament from XII. to I. ; 14, Rm. from XI. to I. ; 15, superior cervical ganglion of X. sympathicus ; 1(5, pharyngeal ramus from IX. at Pe. ; 17, anastomotic filament from 16 to I. ; 18, anastomotic fila- ment from 16 to Px. phar. ; 19, filament from 16 to X. laryngeus superior; 20, cephalic ramus of IX. ; 21, caudal ramus of IX. ; 22, X. pharyngeus : 22 a, cephalic ramus ; 22 b, caudal ramus ; 23, filament from 22 to I. ; 24, pharyngeal ramus of 22 b ; 25, oesophageal ramus of 22 b ; 26, filament from 25 to 22 a ; 27, X. laryngeus superior ; 28, cephalic = ental ramus; 29, caudal = ectal ramus ; a, twig to 50 ; 30, pharyngeal ramus of 29 ; 31, filament from 30 to 25 ; 32, Rm. of 29 to Mb. crico-thyroidea ; 33, to Cp. thyroideum, a, to descendens noni ; 34, cephalic offset of 28 ; 35, fila- ments to interior of larynx ; 36, to epiglottis ; 37, to arytamo-epiglottidean folds ; 38, caudal offset of 28 ; 39, twig from 34 to 38 ; 40, twig from 38 to M. arytamoideus, etc. ; 41, palmate plexus ; 42, ramus to 50 ; 43, X. car- diacus magnus sinister ; 44, X. cardiacus minor ; 45, origin of 50 ; 46, division of X. vagus sinister ; 47, union of lateral rami ; 48, union of mesal rami ; 49, division of X. vagus dexter ; 50, X. laryngeus inferior. *The extent to which secretions and excretions may be referred directly to N. vagus is questionable. 138 Stowell.] Description of the figures. [July 15, Fig. l.—General view of venter of brain ; special reference to venter of medulla, area post pontilis, showing relations of lines of origin-filaments of NN. glosso-pharyngeus, vagus, and liypoglossus ; also ectal relations, ventripyramis (vpy.), area elliptica (ar. el.)> and the lateral tract (?). Pig. 2.—View of sinistral surface of brain, special reference to curved line of origin-filaments of X. vagus and to origin line (L) ventrad, and their relations ; the cephalic filaments of X. accessorius (XI) are in the depression line ventral from Ilf., while the caudal filaments have their origin in the depression line lateral from O. Pig. 3.—Diagram to show the origin of X. vagus ventro-lateral to Rf. ; also that X. accessorius (XI) has its cephalic filaments from the same de- pression line, and its caudal filaments from the depression line ventro- lateral to O ; X. liypoglossus (XII) is dorso-lateral to ar. el. Fig. 4.—Dorsal aspect of metencephalon (medulla) showing origins of XX. IX, X, XI ; metaccelia (mtc.), dorsipyramis (clpy.), corpus resti- forme (Rf.), oliva (0), and the lateral tract (?). Fig. 5.—To show relations of origin-filaments ; of Rm. auricularis ; of G. jugulare ; of G. petrosum ; of G. inferius. G. Ehrenritteri is removed from its normal relation which is ectal to G. jugulare, and is placed ceplialad to expose the parts. X. XII, is reflected dorsad to expose origin of X. pharyngeus and anastomotic ramus from Pe. The dotted lines represent XX. liyo-thyroideus and desce.ndens noni. Px. pliar. = pharyngeal plexus. Fig. 6. is Fig. 5, dissected to show Rm. accessorius given to J., and the second accession to the trunk peripheral to J. ; X. XII, is omitted as are the anastomotic filaments of Px. gangliformis ; the dotted line shows the direction of the filaments from X. XI, to X. pharyngeus. Fig. 7.—X. laryngeus superior ; origin; division ; distribution of ental or cephalic and ectal or caudal rami ; anastomotic relation between pharyngeal ramulus of the ectal ramus and the oesophageal ramus of X. pharyngeus. Dig. 8.—X. laryngeus inferior. To show the laryngeal relations of X. laryngeus inferior ; entad of Ctl. tliyroideus ; the palmate plexus ; the anastomotic filaments of XX. laryngeus superior and laryngeus inferior ; the pharyngeal ramus of X. laryngeus inferior (5°). Fig. 0.—X. laryngeus inferior sinister. To show its origin ; relations with A. aorta and adjacent plexus ; relations of X. vagus with X. sympatliicus ; division of X. vagus dorso-caudad of root of lungs ; the relations of the lateral and the mesal rami ; the dorsal and the ventral pulmonary plexus ; the formation and the relations of the dorsal and the ventral nerve trunks. Fig. 10.—X. laryngeus inferior dexter. To show its origin ; its relations with A. subclavia ; relations of X. vagus with X. sympatliicus. Fig. 11 and 12.—Tracheo-oesophageal ramuli of X. laryngeus inferior sinister and dexter respectively. Fig 13.—Distribution and relations of the ventral gastric nerve and the ramus which terminates in the dextral G. semilunare of Px. Solaris.