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A text-book of veterinary anatomy

Chapter 506: THE EIGHTH OR AUDITORY NERVE
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A comprehensive, systematically organized veterinary anatomy textbook presenting detailed descriptions and abundant photographic illustrations of skeletal, articular, muscular, and visceral structures of major domestic species (horse, ox, pig, dog). It emphasizes topographic relations alongside descriptive morphology, relies on modern preparation techniques to reflect natural organ shape, addresses nomenclature standardization while omitting embryology and histology for practicality, and provides guidance useful for students and practitioners.

Fig. 515.—Deep Dissection of Base of Cranium, Viewed from the Right and Below.

The tympanic cavity is opened and the paramastoid process is sawn off. a, Occipital condyle; b, body of sphenoid; c, external auditory meatus (part removed); d, malleus; e, incus; 1, stump of ophthalmic nerve; 2, maxillary nerve (cut off); 3, sphenopalatine plexus; 4, Vidian nerve; 4′, deep petrosal nerve; 4″, branch to tympanic plexus; 4‴, superficial petrosal nerve; 5, superficial temporal nerve (cut off); 6, masseteric nerve (cut off); 7, mandibular nerve (raised); 8, lingual nerve (cut off); 9, pterygoid nerve (cut); 10, nerve to tensor tympani; 11, otic ganglion; 12, facial nerve; 13, stapedial nerve; 14, chorda tympani; 15, glosso-pharyngeal nerve (cut); 16, tympanic nerve; 17, vagus (cut off); 18, auricular branch of vagus; 19, spinal accessory nerve (cut off); 20, hypoglossal nerve (cut off); 21, sympathetic nerve (cut off); 22, internal carotid artery. (After Ellenberger, in Leisering’s Atlas.)

Before its division the nerve gives off twigs to the guttural pouch, the parotid gland, the external ear, and the skin of the external auditory meatus and the membrana tympani. Branches from it concur with filaments from the cervical branch of the facial nerve in the formation of the auricular plexus.

6. The inferior alveolar or dental nerve (N. alveolaris mandibulæ) (Figs. 436, 437, 514) arises with the lingual by a common trunk which passes forward at first on the external pterygoid muscle, then inclines ventrally between the internal pterygoid and the ramus of the mandible. The lingual and alveolar separate at an acute angle, and the latter enters the mandibular foramen and courses in the canal within the ramus (Fig. 516). Emerging at the mental foramen, it terminates by dividing into six to eight inferior labial and mental branches, which ramify in the lower lip and chin. Before entering the bone, the nerve detaches the mylo-hyoid nerve (N. mylo-hyoideus), which runs downward and forward between the ramus and the mylo-hyoid muscle; it supplies that muscle, the anterior belly of the digastricus, and the skin of the anterior part of the submaxillary space. The dental and gingival branches detached from the nerve within the mandible are arranged like the corresponding nerves of the upper jaw.

7. The lingual nerve (N. lingualis) arises by a common trunk with the inferior alveolar or dental nerve (Figs. 436, 437). After separating from the latter it runs downward and forward, lying at first between the ramus of the mandible and the internal pterygoid muscle, then on the inner face of the mylo-hyoid. On reaching the root of the tongue it divides into superficial and deep branches. The superficial branch (Ramus superficialis) runs forward on the stylo-glossus and accompanies the submaxillary duct on the deep face of the sublingual gland. It supplies the mucous membrane of the tongue and the floor of the mouth. At the root of the tongue it gives off a recurrent branch to the isthmus faucium, which communicates with the lingual branch of the glosso-pharyngeal nerve. The larger deep branch (Ramus profundus) turns around the lower edge of the hyo-glossus, passes upward and forward between that muscle and the genio-glossus, and continues forward on the latter to the tip of the tongue. It gives branches to the mucous membrane and the fungiform papillæ of the tongue, and anastomoses with branches of the hypoglossal nerve and with the superficial branch. Minute ganglia occur on the finer branches of the lingual nerve. The chorda tympani branch of the facial joins the lingual nerve at the origin of the latter and is incorporated with it in its distribution to the tongue.

Fig. 516.—Part of Branch of Lower Jaw of Horse, Internal View.

The bone has been removed to show the vessels and nerves. 1, 1′, Inferior alveolar or dental nerve; 2, 2, branches to cheek teeth and gums; 2′, branch to canine and incisor teeth; 3, 3′, inferior alveolar or dental artery; 4, satellite vein. (After Leisering’s Atlas.)

THE SIXTH OR ABDUCENT NERVE

The abducent nerve (N. abducens) emerges from the brain behind the pons and just external to the pyramid (Fig. 499). It passes forward across the pons, pierces the dura mater, and accompanies the third and ophthalmic nerves, below which it emerges through the foramen lacerum orbitale. In the orbit it divides into two branches; the larger of these enters the external rectus, the smaller supplies the superior and external parts of the retractor muscle of the eyeball. Within the cranium it receives filaments from the carotid plexus of the sympathetic.

The fibers of the abducent nerve are axones of the large multipolar cells of the abducent nucleus which is situated beneath the eminentia teres of the floor of the fourth ventricle. The nucleus lies within the loop formed by the fibers of origin of the facial nerve. It is connected with (a) the anterior olive; (b) the oculomotor nucleus of the opposite side; (c) the motor area of the cerebral cortex by means of the pyramidal tract of the opposite side.

THE SEVENTH OR FACIAL NERVE

The facial nerve (N. facialis) has its superficial origin at the lateral part of the corpus trapezoideum, immediately behind the pons (Fig. 499). It passes outward in front of the eighth nerve and enters the internal auditory meatus. At the bottom of the meatus the two nerves part company, the facial coursing in the facial canal of the petrous temporal bone. The canal and nerve are at first directed outward between the vestibule and the cochlea, then curve backward and downward in the posterior wall of the tympanum to end at the stylo-mastoid foramen. The bend formed by the nerve is called the knee (Geniculum n. facialis) and bears at its highest point the round geniculate ganglion (G. geniculi).

The nerve consists of two parts, motor and sensory. The motor part constitutes the bulk of the nerve. Its deep origin is from the cells of the facial nucleus, which is situated in the medulla above the facial tubercle. On leaving the nucleus the root-fibers pass upward and inward, incline forward close to the median plane, and then bend sharply downward to the point of emergence. The highest point of the bend is subjacent to the gray matter of the eminentia teres in the floor of the fourth ventricle, and the abducent nucleus lies in the concavity of the curve. The small sensory part (N. intermedius) consists of axones of cells of the geniculate ganglion, which is interposed on the facial nerve as it bends downward in the facial canal. The fibers of this part after entering the medulla, pass to the nucleus of termination which it shares with the ninth and tenth nerves. The peripheral fibers from the geniculate ganglion constitute the chorda tympani.

After its emergence through the stylo-mastoid foramen (Fig. 514) the nerve passes downward, forward, and outward on the guttural pouch under cover of the parotid gland, and crosses between the origin of the superficial temporal and internal maxillary arteries internally and the superficial temporal vein externally. It then crosses the posterior border of the ramus of the mandible ventral to the transverse facial artery and about an inch and a half (ca. 3.5 to 4 cm.) below the articulation of the jaw. Emerging from beneath the parotid gland upon the masseter muscle, it receives the lower branch of the superficial temporal nerve, and divides into superior and inferior buccal branches. The following collateral branches are given off, the first five being detached within the facial canal, and the others between the stylo-mastoid foramen and the border of the jaw.

1. The great superficial petrosal nerve (N. petrosus superficialis major) arises from the geniculate ganglion.[203] It passes through the petrosal canal, contributes a filament to the tympanic plexus, receives the great deep petrosal nerve from the carotid plexus of the sympathetic, emerges through the foramen lacerum, and is continued as the Vidian nerve to the sphenopalatine plexus and ganglia (Fig. 515).

2. A delicate branch (R. anastomoticus cum plexu tympanico) emerges from the geniculate ganglion and unites with a filament issuing from the tympanic plexus to form the small superficial petrosal nerve (N. petrosus superficialis minor); this ends in the otic ganglion.

3. The stapedial nerve (N. stapedius) (Fig. 515) is a short filament detached from the facial nerve as it turns down in the facial canal. It innervates the stapedius muscle.

4. The chorda tympani (Fig. 515) is a small nerve which arises a little below the preceding and pursues a recurrent course in a small canal in the mastoid part of the temporal bone to reach the tympanic cavity. It traverses the latter, passing between the handle of the malleus and the long branch of the incus. Emerging through the petro-tympanic fissure, the nerve passes downward and forward, crosses beneath the internal maxillary artery, and joins the lingual nerve. It sends twigs to the submaxillary and sublingual glands, and through its incorporation with the lingual nerve furnishes fibers to the mucous membrane of the anterior two-thirds of the tongue which are believed to mediate the sense of taste.

5. Anastomotic filaments unite with the auricular branch of the vagus near the stylo-mastoid foramen.

6. The posterior auricular nerve (N. auricularis posterior) arises from the facial at its emergence from the facial canal (Fig. 514). It runs upward and backward with the posterior auricular artery under cover of the parotid gland and supplies the posterior auricular muscles and the skin of the convex surface of the external ear. It anastomoses with branches of the first and second cervical nerves.

7. The internal auricular nerve (N. auricularis internus) springs from the facial close to or in common with the preceding (Fig. 514). It ascends in the parotid gland just behind the styloid process of the conchal cartilage, passes through an opening in the cartilage, and ramifies in the skin of the concave surface of the ear.

8. The digastric branch (R. digastricus) (Fig. 514) arises from the facial below the auricular nerves. Its branches innervate the posterior belly of the digastricus, the stylo-hyoideus, and the occipito-hyoideus. At its origin it gives off a small branch which forms a loop around the great auricular artery or its posterior branch and rejoins the trunk.

9. The auriculo-palpebral nerve (N. auriculo-palpebralis) (Fig. 514) arises from the upper edge of the facial near the posterior border of the ramus. It ascends in the parotid gland behind the superficial temporal artery, and terminates in anterior auricular and temporal branches. The anterior auricular branches form with branches of the trigeminus the anterior auricular plexus. They innervate the anterior auricular and parotido-auricularis muscles. The temporal branch runs forward and inward over the temporal muscle to the inner canthus of the eye, forms a plexus with the terminal branches of the ophthalmic nerve, and is distributed to the orbicularis oculi, corrugator supercilii, and levator nasolabialis.

10. The cervical branch (R. colli) (Fig. 435) arises from the ventral border of the facial opposite to the preceding nerve. It emerges obliquely through the parotid gland, passes downward and backward on or near the jugular vein, and anastomoses with the cutaneous branches of the cervical nerves. It gives branches to the parotido-auricularis and the cervical panniculus. In its course along the neck the nerve is reinforced by twigs from the cutaneous branches of the second to the sixth cervical nerves.

11. Small branches are detached to the guttural pouch and the parotid gland. The latter (Rami parotidei) concur with branches of the superficial temporal nerve in forming the parotid plexus.

The facial nerve usually terminates after a short course on the surface of the masseter by dividing into two buccal branches (Figs. 435, 461).

1. The superior buccal nerve (N. buccalis dorsalis) passes forward on the upper part of the masseter, dips under the zygomaticus, and continues along the lower border of the dilatator naris lateralis. It then runs under the last-named muscle and anastomoses with branches of the infraorbital nerve, and is distributed to the muscles of the cheek, upper lip, and nostril.

2. The inferior buccal nerve (N. buccalis ventralis) crosses the masseter obliquely and continues forward along the depressor labii inferioris. It is connected by variable anastomotic branches with the superior nerve. It gives collateral branches to the panniculus, buccinator, and depressor labii inferioris, and ramifies with the terminal branches of the inferior alveolar nerve in the lower lip.

The buccal nerves are subject to much variation in regard to their course, anastomoses, and relations to the sensory components derived from the superficial temporal nerve. Their distribution is constant. The point at which the branch of the superficial temporal nerve joins the facial is variable.

THE EIGHTH OR AUDITORY NERVE

The auditory nerve (N. acusticus) is connected with the lateral aspect of the medulla just behind and external to the facial (Fig. 499). It has two roots, vestibular and cochlear (Radix vestibularis et cochlearis).

The auditory nerve consists of two distinct parts which might well be regarded as separate nerves. The cochlear part mediates the sense of hearing, while the vestibular part is not auditory in function, but is concerned in the sense of the position of the body and the mechanism of equilibration.

The nerve passes outward to the internal auditory meatus, which it enters behind the facial nerve. In the meatus it divides into two nerves, of which the upper is the vestibular and the lower is the cochlear nerve.

1. The vestibular nerve (N. vestibuli) is distributed to the utriculus, the sacculus, and to the ampullæ of the semicircular canals, of the internal ear. In the internal auditory meatus the nerve is connected by filaments with the geniculate ganglion of the facial nerve. At the bottom of the meatus it bears the vestibular ganglion (G. vestibulare), from the cells of which the fibers of the nerve arise.

2. The cochlear nerve (N. cochleæ) detaches a filament to the sacculus, passes through the lamina cribrosa to the labyrinth, and is distributed to the organ of Corti in the cochlea.

The fibers of the vestibular nerve arise from the vestibular ganglion as central processes (axones) of the bipolar cells of the ganglion. The peripheral processes (dendrites) of the cells form arborizations about the deep ends of the hair cells of the maculæ and cristæ acusticæ of the utriculus, sacculus, and semicircular canals. The fibers enter the medulla, pass between the restiform body and the spinal tract of the trigeminus, and spread out to end in the vestibular nucleus in the floor of the fourth ventricle. Among the central connections of the vestibular nerve are: (1) fibers which connect its nucleus with centers in the cerebellum (chiefly of the opposite side); (2) the vestibulo-spinal tract, which conveys impulses to the motor cells of the ventral columns of the spinal cord; (3) fibers which connect the nucleus with those of the abducent nerve of the same side, the third and fourth nerves, and the motor part of the trigeminus of both sides.

The fibers of the cochlear nerve are the central processes of the bipolar cells of the spiral ganglion of the cochlea. The peripheral processes of these cells end in relation to the hair cells of the organ of Corti. Some of the nerve-fibers enter the ventral cochlear nucleus in the medulla close to the superficial origin of the nerve; others end in the dorsal nucleus of the tuberculum acusticum at the lateral angle of the floor of the fourth ventricle. From the ventral nucleus fibers pass in the corpus trapezoideum to the anterior olivary nucleus of the same and of the opposite side. Thence tracts pass to the nuclei of the motor nerves of the eye, and through the lateral fillet to the posterior quadrigeminal body and the internal geniculate body. The axones of the cells of the dorsal nucleus pass largely (as the striæ acusticæ) over the restiform body and across the floor of the fourth ventricle toward the median plane. They then turn ventrally, cross to the opposite side, and are continued by the lateral fillet. From the mid-brain a tract proceeds to the cortex of the temporal lobe of the hemisphere.

THE NINTH OR GLOSSO-PHARYNGEAL NERVE

The glosso-pharyngeal nerve (N. glossopharyngeus) is attached to the anterior part of the lateral aspect of the medulla by several filaments (Fig. 499). The root-bundles enter the furrow ventral to the restiform body; they are separated by a short interval from the origin of the facial nerve, but are not marked off behind from the roots of the vagus. The bundles converge laterally to form a nerve which perforates the dura mater and emerges through the foramen lacerum posterius just in front of the tenth nerve (Fig. 515). As it issues from the cranium the nerve bears a considerable ovoid gray enlargement, the petrous ganglion (G. petrosum).[204] It then curves downward and forward over the guttural pouch and behind the great cornu of the hyoid bone, crosses the deep face of the external carotid artery, and divides into pharyngeal and lingual branches (Fig. 437). The collateral branches are as follows:

1. The tympanic nerve (N. tympanicus) (Fig. 515) arises from the petrous ganglion and passes upward between the petrous and tympanic parts of the temporal bone to reach the cavity of the tympanum. Here it breaks up into branches to form, along with branches from the carotid plexus of the sympathetic, the tympanic plexus. From the plexus branches pass to the mucous membrane of the tympanum and the Eustachian tube. The continuation of the nerve issues from the plexus and unites with a filament from the geniculate ganglion of the facial to form the small superficial petrosal nerve; this runs forward and ends in the otic ganglion.

Filaments also connect the petrous ganglion with the jugular ganglion of the vagus nerve and with the superior cervical ganglion of the sympathetic.

2. A considerable branch runs backward on the guttural pouch, contributes filaments to the pharyngeal plexus, and concurs with twigs from the vagus and the sympathetic in forming the carotid plexus on the terminal part of the carotid artery and on its chief branches. In this plexus is the small ganglion intercaroticum.

The pharyngeal branch (R. pharyngeus) (Fig. 437) is the smaller of the two terminal branches. It runs forward across the deep face of the great cornu of the hyoid bone and concurs with the pharyngeal branches of the vagus and with sympathetic filaments in forming the pharyngeal plexus; from this branches pass to the muscles and mucous membrane of the pharynx.

The lingual branch (R. lingualis) is the continuation of the trunk (Fig. 437). It runs along the posterior border of the great cornu of the hyoid bone in front of the external maxillary artery and dips under the hyo-glossus muscle. It gives collateral branches to the soft palate, isthmus faucium, and tonsil, and ends in the mucous membrane of the posterior part of the tongue, where it supplies gustatory fibers to the vallate papillæ. A considerable branch unites with a twig from the lingual nerve.

The glosso-pharyngeal is a mixed nerve, containing both motor and sensory fibers. The latter constitute the bulk of the nerve and include those which mediate the special sense of taste. They are processes of the cells of the petrous ganglion. The central processes of the ganglion cells enter the medulla, pass upward and inward through the formatio reticularis, and end in the nucleus of termination in the floor of the fourth ventricle. The motor fibers arise from dorsal and ventral efferent nuclei in the medulla. The glosso-pharyngeal shares these nuclei with the vagus and has practically the same central connections as that nerve (q. v.).

THE TENTH, VAGUS, OR PNEUMOGASTRIC NERVE

The vagus or pneumogastric (N. vagus) is the longest and most widely distributed of the cranial nerves; it is also remarkable for the connections which it forms with adjacent nerves and with the sympathetic. It is attached to the lateral aspect of the medulla by several filaments which are in series with those of the ninth nerve in front and the eleventh nerve behind (Fig. 499). The bundles converge to form a trunk which passes outward, pierces the dura mater, and emerges from the cranium through the foramen lacerum posterius (Fig. 515). In the foramen the nerve bears on its lateral aspect the elongated flattened jugular ganglion (G. jugulare).

The ganglion communicates with (a) the tympanic nerve, (b) the petrous ganglion of the ninth nerve, (c) the spinal accessory, and (d) the hypoglossal. It also gives off the auricular branch (R. auricularis), which runs forward below the petrous ganglion and passes through a small canal in the petrous temporal bone to gain the facial canal. Here it gives filaments to the facial and emerges with that nerve through the stylo-mastoid foramen. It ascends behind the external auditory meatus, dips under the rotator longus muscle, and passes through a foramen in the conchal cartilage to ramify in the integument which lines the meatus and the adjacent part of the ear.

Beyond the ganglion the vagus runs backward and downward with the spinal accessory in a fold of the guttural pouch (Fig. 437). Then the two nerves separate, allowing the hypoglossal to pass between them, and the vagus descends with the internal carotid artery and crosses the inner face of the origin of the occipital artery. Here it is joined by the cervical trunk of the sympathetic, and the two nerves continue along the dorsal aspect of the common carotid artery in a common sheath (Fig. 433). At the root of the neck the vagus separates from the sympathetic, and from this point backward the relations of the right and left vagi differ somewhat and must be described separately.

Fig. 517.—Cross-section of Neck of Horse, Passing through Posterior Part of Atlas.

The head and neck were extended. 1, Dorsal arch of atlas; 2, odontoid process; 3, odontoid ligament; 4, vertebral sinuses; 5, dura mater; 6, spinal cord; 7, vertebral artery; 8, wing of atlas; 9, atlanto-axial joint cavity; 10, œsophagus; 11, 11, recurrent nerves; 12, 12, ventral branches of spinal accessory nerves. By an oversight the obliquus cap. post. (above wing of atlas) is unmarked.

The right vagus (Fig. 429) enters the thorax in the angle of divergence of the right brachial artery and the truncus bicaroticus. It then passes backward and slightly upward, crossing obliquely the outer surface of the brachiocephalic artery and the right face of the trachea. Reaching the dorsal surface of the latter near the bifurcation, it divides into dorsal and ventral branches.

The left vagus (Fig. 428) enters the thorax on the ventral face of the œsophagus, crosses obliquely under the left brachial artery, and passes back on the external surface of that vessel in company with a large cardiac nerve.[205] Separating from the latter, the vagus continues backward on the left face of the aorta, inclines to the upper surface of the left bronchus, and divides into dorsal and ventral branches.

The dorsal and ventral branches unite with the corresponding branches of the opposite nerve, thus forming dorsal and ventral œsophageal trunks (Truncus (œsophageus dorsalis, ventralis). These run backward in the posterior mediastinum, above and below the œsophagus respectively, and enter the abdominal cavity through the hiatus œsophageus; they supply branches to the œsophagus and anastomose with each other. The dorsal trunk receives the major part of its fibers from the right vagus. After entering the abdomen it passes to the left of the cardia, gives branches to the visceral surface of the stomach, and ends in the cœliac and subsidiary plexuses. The ventral trunk passes to the lesser curvature of the stomach and ramifies on the parietal surface of the stomach; it forms here the anterior gastric plexus, from which branches are supplied also to the first part of the duodenum and to the liver.

The collateral branches of the vagus are as follows:

1. The pharyngeal branch (R. pharyngeus) is given off in relation to the superior cervical ganglion, turns around the internal carotid artery, and runs downward and forward on the guttural pouch to the dorsal wall of the pharynx (Fig. 437). Here its branches concur with the pharyngeal branch of the ninth nerve and with filaments from the spinal accessory and the sympathetic in forming the pharyngeal plexus. This supplies numerous twigs to the pharynx, and a larger branch which passes along the side of the œsophagus and ramifies in its cervical part.

According to Ellenberger and Baum the pharyngeal plexus receives filaments also from the digastric, superior laryngeal, hypoglossal, and first cervical nerves. The branches of the plexus form secondary intermuscular and submucous plexuses, in which there are numerous minute ganglia.

2. The superior or anterior laryngeal nerve (N. laryngeus cranialis) is larger than the preceding and arises a little behind it (Fig. 437). It crosses the deep face of the origin of the external carotid artery, runs downward and forward over the lateral wall of the pharynx behind the hypoglossal nerve, and passes through the foramen below the anterior cornu of the thyroid cartilage. Its terminal branches ramify in the mucous membrane of the larynx, the floor of the pharynx, and the entrance to the œsophagus; they anastomose with those of the recurrent. At its origin the nerve gives off its small external branch (R. externus); this descends to the crico-thyroid muscle, which it supplies, and sends filaments to the crico-pharyngeus also. It may arise from the trunk of the vagus or from the pharyngeal branch.

At the point of origin of the superior laryngeal nerve there is a plexiform widening which is regarded by some authors as the homologue of the ganglion nodosum of man. From it a filament arises which, after a short course, rejoins the vagus or enters the sympathetic trunk. Stimulation of its central end causes a reduction of the blood-pressure, and it is therefore termed the depressor nerve (N. depressor).

3. The recurrent nerve (N. recurrens), also termed the inferior or posterior laryngeal nerve, differs on the two sides in its point of origin and in the first part of its course. The right nerve (Fig. 429) is given off opposite the second rib, turns around the dorso-cervical artery from without inward, runs forward on the lower part of the lateral surface of the trachea, and ascends in the neck on the ventral face of the common carotid artery. The left nerve (Fig. 428) arises from the vagus where the latter begins to cross the aortic arch. It passes back over the ligamentum arteriosum, winds around the concavity of the aortic arch from without inward, runs forward on the lower part of the left face of the trachea, and continues in the neck in a similar position to the right nerve.

It is worthy of note that the left nerve passes beneath the bronchial lymph glands as it winds around the aorta; also that in the next part of its course it lies between the left surface of the trachea and the deep face of the aorta, and is then related to lymph glands which lie along the ventral aspect of the trachea. The left recurrent is often incorporated in part of its course in the anterior mediastinum with a deep cardiac nerve. Further, the left nerve lies at first ventral to, and then upon, the œsophagus in the neck. The right recurrent is given off from or in common with a considerable trunk which connects the vagus with the first thoracic ganglion of the sympathetic. The arrangement here is commonly more or less plexiform, and from it one or two cardiac nerves arise.

The terminal part of each nerve (Fig. 517) lies on the dorsal surface of the trachea, in relation to the œsophagus internally and the carotid artery above.[206] It passes between the crico-arytenoideus posterior and the crico-pharyngeus; the terminal branches supply all the muscles of the larynx except the crico-thyroid, and communicate with branches of the superior laryngeal nerve. Collateral branches are given off to the cardiac plexus (Rr. cardiaci), to the trachea (Rr. tracheales), to the œsophagus (Rr. œsophagei), and to the inferior cervical ganglion of the sympathetic.

4. Cardiac branches (Rr. cardiaci), usually two or three in number, are given off from each vagus within the thorax (Figs. 428, 429). These concur with the cardiac branches of the sympathetic and recurrent nerves to form the cardiac plexus, which innervates the heart and great vessels.

5. Small tracheal and œsophageal branches (Rr. tracheales et œsophagei) are given off from both vagi in the thorax. These concur with branches from the recurrent nerves and the inferior cervical and anterior thoracic ganglia of the sympathetic in forming the posterior tracheal and œsophageal plexuses, from which twigs go to the trachea, œsophagus, heart, and large vessels.

6. Bronchial branches (Rr. bronchiales) are detached at the roots of the lungs and unite with sympathetic filaments in forming the pulmonary plexuses. From the latter numerous branches proceed in a plexiform manner along the bronchi and vessels into the substance of the lungs.

The vagus and glosso-pharyngeal nerves are so closely associated in origin and central connections that they may be described together in this respect.

The sensory fibers arise from the petrous and jugular ganglia, and their central parts enter the lateral aspect of the medulla and divide into anterior and posterior branches like the fibers of the dorsal roots of the spinal nerves. Most of the fibers end in arborizations about the cells of the vago-glosso-pharyngeal nucleus of termination, which consists of two parts. Of these the dorsal sensory nucleus (Nucleus alæ cinereæ) is situated in the posterior part of the floor of the fourth ventricle and in the adjacent part of the closed portion of the medulla near the median plane. The other part is termed the nucleus of the solitary tract, and is so named because its cells are grouped about the bundle (Tractus solitarius) formed by the posterior divisions of the afferent nerve-fibers. It ends about the level of the pyramidal decussation. The secondary central connections are similar to those of the sensory part of the trigeminus.

The motor fibers (and those of the medullary part of the accessory) arise from the dorsal motor nucleus and the ventral motor nucleus. The cells of the former lie in groups along the ventro-medial side of the dorsal sensory nucleus. The latter, also termed the nucleus ambiguus, is situated more deeply in the lateral part of the formatio reticularis.

THE ELEVENTH OR SPINAL ACCESSORY NERVE

The spinal accessory nerve (N. accessorius) is purely motor. It consists of two parts which differ in origin and function.

The medullary part arises from the lateral aspect of the medulla by several rootlets which are behind and in series with those of the vagus (Fig. 499). The spinal part arises from the cervical part of the spinal cord by a series of fasciculi which emerge between the dorsal and ventral roots. The bundles unite to form a trunk which is very small at its origin at the fifth segment of the cord, but increases in size when traced toward the brain, since it continually receives accessions of fibers. It passes through the foramen magnum and joins the medullary part. The trunk thus formed sends its medullary fibers to the tenth and ninth nerves and emerges through the foramen lacerum posterius. It then runs backward and downward with the vagus in a fold of the guttural pouch, separates from that nerve, crosses the deep face of the submaxillary gland and the occipital artery, and divides in the recessus atlantis into dorsal and ventral branches.

Fig. 518.—Neck of Horse, after Removal of Part of Panniculus and Trapezius.

a, a′, Mastoido-humeralis; b, anterior superficial pectoral muscle; c, cervical panniculus; d, sterno-cephalicus; e, omo-hyoideus; f, sterno-thyro-hyoideus; g, trachea; h, h′, h″, tendons of splenius, mastoido-humeralis, and trachelo-mastoideus; i, trapezius cervicalis; k, supraspinatus; l, anterior deep pectoral muscle; m, rhomboideus cervicalis; n, serratus cervicis; o, splenius, upper and lower borders of which are indicated by dotted lines; p, parotid gland; q, parotido-auricularis muscle; r, wing of atlas; s, spine of scapula; 1, external maxillary vein; 2, 3, jugular vein; 4, carotid artery; 5, descending branch of inferior cervical artery; 6, cephalic vein; 7–11, ventral branches of second to seventh cervical nerves; 12, cutaneous branch of second cervical nerve; 13, cervical branch of facial nerve; 14, terminal branches of dorsal divisions of cervical nerves; 15, dorsal branch of spinal accessory nerve. (After Ellenberger-Baum, Top. Anat. d. Pferdes.)

It is connected by anastomotic branches with the vagus and hypoglossal nerves and the superior cervical ganglion of the sympathetic, and contributes a branch to the pharyngeal plexus.

The dorsal branch (R. dorsalis) (Figs. 433, 517, 518) receives a twig from the second and third cervical nerves and turns around the atlantal tendon of the splenius under cover of the mastoido-humeralis. It then passes obliquely through the latter muscle and continues backward on the splenius and the cervical part of the serratus, inclines upward across the anterior deep pectoral and the supraspinatus, and enters the deep face of the trapezius, in which it ramifies.

The ventral branch (R. ventralis) (Fig. 436) is smaller. It enters the sterno-cephalicus muscle behind the cervical angle of the parotid gland.

The fibers of the spinal part of the accessory arise from the ventro-lateral cells of the ventral gray column of the cord as far back as the fifth cervical segment. The fibers of the medullary part come chiefly from the nucleus ambiguus in common with the motor fibers of the vagus.

THE TWELFTH OR HYPOGLOSSAL NERVE

The hypoglossal nerve (N. hypoglossus) is purely motor and innervates the muscles of the tongue (Fig. 437). Its root-fibers arise from the ventral face of the medulla in linear series about 3 to 4 mm. lateral to the posterior half of the pyramid (Fig. 499). The filaments converge to three or four bundles which perforate the dura mater and unite to form the trunk. The latter emerges through the hypoglossal foramen (Fig. 515) and runs downward and backward between the guttural pouch and the capsule of the atlanto-occipital articulation for a distance of a little less than an inch (ca. 2 cm.). It then passes between the tenth and eleventh nerves, turns downward and forward, crosses the external face of the external carotid artery, and continues over the pharynx parallel with the great cornu of the hyoid bone and behind the external maxillary artery. It then crosses beneath the artery, runs forward on the external face of the hyo-glossus muscle, and divides into its terminal branches (Rami linguales). The smaller branch supplies the stylo-glossus, hyo-glossus, and lingualis. The larger branch ramifies on the genio-glossus and supplies the remaining muscles. Anastomoses occur with branches of the lingual nerve.

In the first part of its course the nerve communicates with the superior cervical ganglion and with the ventral branch of the first cervical nerve, and gives filaments to the pharyngeal branch of the vagus and the pharyngeal plexus.

The fibers of the nerve arise from the hypoglossal nucleus, an elongated group of large multipolar cells situated chiefly under the posterior part of the floor of the fourth ventricle close to the median plane. The two nuclei are connected by commissural fibers. The other central connections include: (a) communications by the medial longitudinal fasciculus with the nuclei of termination of other cranial nerves; (b) cortico-nuclear fibers which come from the cortex by way of the internal capsule and the pyramids and go largely to the nucleus of the opposite side; (c) fibers which join the dorsal longitudinal bundle of Schütz, a tract which underlies the floor of the fourth ventricle and is traceable forward below the cerebral aqueduct.

The Spinal Nerves

The spinal nerves (Nervi spinales) are arranged in pairs, of which there are usually forty-two in the horse. They are designated according to their relations to the vertebral column as cervical (8), thoracic (18), lumbar (6), sacral (5), and coccygeal (5). Each nerve is connected with the spinal cord by two roots, dorsal and ventral (Fig. 497).

The dorsal (or superior) root (Radix dorsalis) is the larger of the two. Its fibers (Fila radicularia) spread out in fan shape and join the cord in a linear series along the dorso-lateral groove. Laterally the fibers converge to form a compact bundle, on which is a gray nodular enlargement, the spinal ganglion (Ganglion spinale). Beyond the ganglion the dorsal root joins the ventral root to constitute the nerve. The ganglia are external to the dura mater, and are situated in the intervertebral foramina, except in the case of the sacral and coccygeal nerves, the ganglia of which lie within the vertebral canal. Those of the coccygeal nerves are intradural.

The ganglia vary greatly in size; that of the first cervical nerve is scarcely as large as a hemp seed, while that of the eighth cervical is about 2 cm. long and 1 cm. wide. On the large roots connected with the cervical and lumbar enlargement of the cord there are multiple ganglia of varying sizes interposed in the course of the root-bundles. The fibers of the dorsal roots arise from the cells of the spinal ganglia; each cell gives off a fiber which enters the spinal cord and another which passes into the nerve.

The ventral (or inferior) root (Radix ventralis) contains fewer fibers than the dorsal root, except in the case of the first cervical nerve. It arises from the ventral surface of the spinal cord (Fig. 495) by means of numerous small bundles of fibers which do not form a linear series but emerge from the cord over an area three to five millimeters in width (ventral root zone). The fibers are processes of the large cells of the ventral gray columns of the spinal cord. There is no ganglion on the ventral root.

In the cervical, thoracic, and anterior lumbar regions the bundles of both roots pass through separate openings in linear series in the dura mater before uniting into a root proper. Further back the bundles of each root unite within the dura. In the anterior part of the cervical region and in the thoracic part of the cord there are intervals of varying length between adjacent roots, but in some places the fibers of adjacent roots overlap and an exchange of fibers may be observed. Many of the roots are directed almost straight outward or incline slightly backward, but the posterior lumbar, sacral, and coccygeal roots and nerves run backward to reach the foramina through which they emerge. The distance thus to be traversed increases from before backward, so that these nerves form a tapering sheaf around the conus medullaris and filum terminale in the last lumbar vertebra and the sacrum which is known as the cauda equina.