The anatomy of the peripheral nervous system


The peripheral nervous system (PNS) is composed of spinal and cranial nerves, their associated ganglia (autonomic and sensory) and their ramifications that transmit afferent and efferent information between the central nervous system (CNS) and the rest of the body. It also includes the peripheral part of the autonomic nervous system (ANS), notably the sympathetic trunks and the sympathetic and parasympathetic ganglia and their ramifications, and the enteric nervous system (ENS), composed of plexuses of nerve fibres and cell bodies in the wall of the gastrointestinal tract, and the visceral afferent system.

Cranial Nerves

Cranial nerves are the means by which the brain receives information from, and controls the activities of, the head and neck and the thoracic and greater part of the abdominal viscera. There are 12 pairs of cranial nerves, individually named and numbered (Roman numerals) in a rostrocaudal sequence (see Table 24.1 ). Unlike spinal nerves, only some cranial nerves are mixed in function, i.e. they carry both sensory and motor fibres; others are purely sensory or motor. The first cranial nerve (I; olfactory) has an ancient lineage and is derived from the forerunner of the cerebral hemisphere. It retains this unique position through the connections of the olfactory bulb, and is the only sensory cranial nerve that projects directly to the cerebral cortex rather than indirectly via the thalamus. The areas of cerebral cortex receiving olfactory input have a primitive cellular organization and are an integral part of the limbic system, which is concerned with the emotional aspects of behaviour. The second cranial nerve (II; optic) consists of the axons of second-order visual neurones and these terminate in the thalamus. Most of the component fibres of the other 10 pairs of cranial nerves originate from, or terminate in, named cranial nerve nuclei in the brainstem.

Olfactory nerve

The olfactory nerve is the first cranial nerve and, as its name suggests, is concerned with olfaction (the sense of smell). It is composed of bundles of very small unmyelinated axons derived from olfactory receptor neurones in the olfactory mucosa. The axons are in varying stages of maturity, reflecting the constant turnover of olfactory neurones that takes place in the olfactory epithelium. Bundles of axons are surrounded by olfactory ensheathing cells and form a plexiform network in the subepithelial lamina propria of the mucosa. The bundles unite into as many as 20 branches that cross the cribriform plate of the ethmoid bone in lateral and medial groups, and enter the overlying olfactory bulb, where they end in glomeruli. Each branch is ensheathed by dura mater and pia/arachnoid as it passes through the cribriform plate (see Fig. 39.13A ). The dura subsequently becomes continuous with the nasal periosteum, and the pia/arachnoid merges with the connective tissue sheaths that surround the nerve bundles, an arrangement that may facilitate the spread of infection into the cranial cavity from the nasal cavity.

The olfactory bulb is continuous posteriorly with the olfactory tract, through which the output of the bulb projects directly to the ipsilateral retrobulbar area (anterior olfactory nucleus), olfactory tubercle, prepiriform area, amygdala and rostral entorhinal cortex. There is a clear laminar structure in the olfactory bulb ( Fig. 32.32A ). Different odour molecules are represented by different patterns of spatial activity in the olfactory bulb ( ).

In severe injuries involving the anterior cranial fossa, the olfactory bulb may be separated from the olfactory nerves or the nerves may be torn, producing anosmia, i.e. loss of olfaction. Fractures may involve the meninges, so that cerebrospinal fluid may leak into the nose, resulting in cerebrospinal rhinorrhoea. These injuries open up avenues for intracranial infection from the nasal cavity.

Optic Nerve

The optic nerve is the second cranial nerve and contains the axons of retinal ganglion cells. The two optic nerves join to form the optic chiasma which is usually positioned over the diaphragma sellae and pituitary gland (normofixed), although it may be prefixed (above the tuberculum sellae) or postfixed (above the dorsum sellae). On entering the optic nerve, retinal ganglion cell axons initially maintain their relative retinal positions, with axons from the fovea forming a lateral wedge. This retinotopic mapping is largely maintained within the optic nerve, although nearer the chiasma the foveal axons take a position in the centre of the optic nerve while temporal fibres occupy their previous lateral location. A substantial rearrangement of axons takes place at the chiasma. Most axons arising from the nasal half of a line bisecting the fovea within each retina cross in the chiasma to enter the contralateral optic tract. Fibres from the temporal hemi-retinas do not generally cross in the chiasma.

Th axons within the optic tract were thought to maintain their topographic order and each tract was assumed to be a single representation of the contralateral hemifield. However, it is now clear that axons are mainly organized in functional groupings, larger superficial axons representing the magnocellular pathway and deeper axons originating from midget ganglion cells and forming the parvocellular pathway. This arrangement is chronotopic, the deeper axons developing earlier during axogenesis than the more superficial ones (Reese 1993).

Almost all (90%) of the retinal ganglion cell axons terminate on neurones in the lateral geniculate nucleus. Extrageniculate axons (10%) leave the optic tract before the lateral geniculate nucleus: they may leave the optic chiasma dorsally and project to the suprachiasmatic nucleus of the hypothalamus, or may branch off the optic tract at the superior brachium and project to the superior colliculus, pretectal areas and inferior pulvinar.

The optic nerve is usually described in four parts, intracranial (approximately 10 mm), intracanalicular (approximately 10 mm), intraorbital (25–30 mm), and intraocular (1 mm). The intracranial part arises from the optic chiasma in the suprasellar cistern. It is surrounded by the optic sheath and its dorsal surface is covered by the falciform ligament. The intracanalicular part passes through the optic canal together with the ophthalmic artery: this is the weakest point of the anterior visual pathway because the nerve is fixed to its bony surroundings by fibrous adhesions. The intraorbital part passes forwards, laterally and downwards, and pierces the sclera at the lamina cribrosa, slightly medial to the posterior pole: it has a somewhat tortuous course within the orbit that allows for movements of the eyeball ( ). The subarachnoid space surrounding the intracranial part is continuous with the subarachnoid space around the intracanalicular and intraorbital parts. The dural sheath surrounding the optic nerve originates at the level of the optic canal and accompanies the optic nerve to the sclera. The intraorbital part of optic nerve has important relationships with other orbital structures (see Fig. 44.4 ). As it leaves the optic canal, it lies superomedial to the ophthalmic artery, and is separated from lateral rectus by the oculomotor, nasociliary and abducens nerves, and sometimes by the ophthalmic veins. It is closely related to the origins of the four recti, whereas more anteriorly, where the muscles diverge, it is separated from them by a substantial amount of orbital fat. Just beyond the optic canal, the ophthalmic artery and the nasociliary nerve cross the optic nerve to reach the medial wall of the orbit. The central artery of the retina enters the substance of the optic nerve about halfway along its length. Near the back of the eyeball, the optic nerve is surrounded by the long and short ciliary nerves and vessels.

Oculomotor Nerve

The oculomotor nerve is the third cranial nerve and innervates levator palpebrae superioris and four of the extraocular muscles (superior, inferior and medial rectus and inferior oblique) (see Fig. 44.15 ). It also conveys preganglionic parasympathetic fibres from cell bodies in the Edinger–Westphal nucleus that relay in the ciliary ganglion.

Oculomotor nuclear complex

The nuclear complex from which the oculomotor nerve arises consists of several groups of large motor neurones that are collectively designated the oculomotor nucleus, and the smaller preganglionic parasympathetic Edinger–Westphal nucleus (see Figure 28.3, Figure 28.16 ). The nuclear complex lies at the ventromedial aspect of the periaqueductal grey matter, superior to the trochlear nuclei and is partially enclosed by the fibres of the median longitudinal fasciculi. It is approximately 5 mm long and is divisible into neuronal groups that correlate with the distribution of motor fibres within the oculomotor nerve. Afferent inputs to the oculomotor nuclear complex include fibres from the rostral interstitial nucleus of the medial longitudinal fasciculus and the interstitial nucleus of Cajal, both of which are involved in the control of vertical and torsional gaze. Other inputs come, directly or indirectly, from the nuclei of the posterior commissure and, via these nuclei, from the frontal eye fields, the superior colliculus, the dentate nucleus and cortical areas. The medial longitudinal fasciculus carries afferent connections from the trochlear, abducens and vestibular nuclei.

The fibres from the medial vestibular nucleus project to the medial rectus subnucleus. In order from dorsal to ventral, the oculomotor neurone groups innervate the ipsilateral inferior rectus, inferior oblique and medial rectus. A medially placed column of cells innervates the contralateral superior rectus through axons that decussate in the caudal part of this subnucleus. A median subnucleus of large neurones, designated the caudal central nucleus, lies at the caudal pole of the oculomotor nucleus adjacent to the superior rectus and medial rectus subnuclei. Fascicles of axons from these subnuclei course forwards in the midbrain, through the tegmentum, red nucleus and medial substantia nigra, to emerge as the oculomotor nerve in the interpeduncular fossa. The fascicles are most probably arranged from medial to lateral subserving the pupil, inferior rectus, medial rectus, levator palpebrae superioris and superior rectus, and inferior oblique.

Intra- and extracranial course

For descriptive purposes, the oculomotor nerve is usually divided into cisternal, cavernous and orbital portions: surgically oriented classifications derived from combined microsurgical and endoscopic cadaveric dissection studies have proposed five or seven parts ( , ).

The cisternal portion of the oculomotor nerve emerges from the midbrain, either through the medial sulcus of the crus cerebri in the caudal part of the interpeduncular fossa or through the medial or the middle part of the crus cerebri. It usually passes forwards and laterally through the interpeduncular cistern between the posterior communicating artery and the superior cerebellar artery: this portion of the nerve is supplied by the P1 and P2 segments of the posterior cerebral artery. The pupilloconstrictor fibres lie superficially on the dorsomedial surface of the cisternal portion of oculomotor nerve, a location that renders them vulnerable to aneurysmal compression ( , ), whereas a pupil-sparing palsy of the oculomotor nerve is pathognomonic of an ischaemic lesion, which tends to involve the central region of the nerve. The oculomotor nerve next passes inferolateral to the posterior communicating artery and inferomedial to the uncus and enters the roof of the cavernous sinus through the oculomotor trigone. This cavernous portion of the nerve passes along the lateral dural wall of the cavernous sinus, dividing into superior and inferior divisions that run beneath the trochlear and ophthalmic nerves. The two divisions enter the orbit through the superior orbital fissure, within the common tendinous ring of the recti, and separated by the nasociliary branch of the ophthalmic nerve.

The superior division of the oculomotor nerve passes above the optic nerve to enter the inferior (ocular) surface of superior rectus. It supplies this muscle and gives off a branch that runs to innervate levator palpebrae superioris. The inferior division of the oculomotor nerve divides into medial, central and lateral branches. The medial branch passes beneath the optic nerve to enter the lateral (ocular) surface of medial rectus; the central branch runs downwards and forwards to enter the superior (ocular) surface of inferior rectus; the lateral branch travels forwards on the lateral side of inferior rectus to enter the orbital surface of inferior oblique and also communicates with the ciliary ganglion to distribute parasympathetic fibres to sphincter pupillae and the ciliary muscle.

Edinger–Westphal nucleus

The Edinger–Westphal nucleus lies dorsal to the main oculomotor nucleus. It is composed of small, multipolar, preganglionic parasympathetic (general visceral efferent, GVE) neurones. Afferent inputs to the nucleus come primarily from the pretectal nuclei bilaterally, mediating the pupillary light reflex, and from the visual cortex, mediating accommodation. Efferent fibres from the nucleus travel in the oculomotor nerve to synapse with postganglionic neurones in the ciliary ganglion ( ), and also descend through the medial longitudinal fasciculus to innervate the medial and descending vestibular nuclei ( ).

Trochlear Nerve

The trochlear nerve is the fourth cranial nerve and innervates superior oblique. It has the longest intracranial course and the smallest diameter of all of the cranial nerves, and is also the only cranial nerve to emerge from the dorsal surface of the brainstem, lateral to the lower edge of the inferior colliculus.

Trochlear nucleus

The trochlear nucleus is a small ovoid mass of grey matter just to the side of the midline (see Fig. 28.15 ). It is bounded by the periaqueductal grey matter dorsally and is enveloped by the fibres of the longitudinal fasciculi ventrally and laterally ( ). The nucleus is part of the somatic efferent column in line with the oculomotor, abducens and hypoglossal nuclei. Afferent inputs to the trochlear nucleus include fibres from the rostral intermediate nucleus of the medial longitudinal fasciculus and from the interstitial nucleus of Cajal, both of which are involved in the control of vertical and torsional gaze. Other inputs come from the vestibular, abducens and oculomotor nuclei and, directly or indirectly, from the nuclei of the posterior commissure (nuclei of Darkschewitsch and of Cajal) and, via these nuclei, from the superior colliculus, frontal eye fields and other cortical areas. Other afferents arrive from the nucleus prepositus hypoglossi. Efferent trochlear fibres first course dorsally and caudally, just deep to the outer perimeter of the periaqueductal grey matter and medial to the mesencephalic trigeminal nucleus. The trochlear nerves then decussate within the superior medullary velum of the fourth ventricle. A few fibres may remain ipsilateral.

Intra- and extracranial course

The nerves emerge from the dorsal surface of the brainstem just caudal to the inferior colliculus and to each side of the frenulum veli. At the point of emergence, a variable number of rootlets, usually two, unite to form the nerve trunks that curve ventrally through the perimesencephalic cisterns, between the superior cerebellar arteries below and the posterior cerebral arteries above. The cisternal portion of each nerve courses anterolaterally, winding round the cerebral peduncle toward the tentorium cerebelli through the quadrigeminal and ambient cisterns. The nerve passes anteriorly along the infratentorial groove of the tentorium cerebelli and penetrates the roof of the cavernous sinus at the junction of the anterior and posterior petroclinoid dural folds. The cavernous portion continues along the lateral wall of the cavernous sinus below the oculomotor nerve, crossing this nerve at the level of the optic strut to become the most superior structure in the cavernous sinus. The trochlear nerve enters the orbit through the superior orbital fissure outside the common tendinous ring, superior to levator palpebrae superioris and medial to the frontal and lacrimal nerves (see Fig. 44.16 ). The orbital portion runs medially across levator palpebrae superior and superior rectus to reach superior oblique: its branches typically end on the posterior one-third of the muscle, most often on the medial aspect ( ).

Trigeminal Nerve

The trigeminal nerve is the fifth cranial nerve and has extensive sensory, motor and autonomic components.

Sensory nuclei and central connections

The trigeminal nerve has three sensory nuclei (spinal tract, principal and mesencephalic) and one motor nucleus. On entering the pons, the fibres of the sensory root of the trigeminal nerve run dorsomedially towards the principal sensory nucleus. About 50% of the fibres divide into ascending and descending branches and the others ascend or descend without division. The descending fibres form the spinal tract of the trigeminal nerve, which terminates in the medially adjacent spinal nucleus of the trigeminal nerve. Some ascending trigeminal fibres, many of them heavily myelinated, synapse around the small neurones in the principal sensory nucleus, which lies lateral to the motor nucleus and medial to the middle cerebellar peduncle, and is continuous inferiorly with the spinal nucleus of the trigeminal nerve. The principal nucleus is considered to be mainly concerned with tactile stimuli. Other ascending fibres enter the mesencephalic nucleus, a column of unipolar cells with peripheral branches that may convey proprioceptive impulses from the masticatory muscles, and possibly also from the teeth, the facial and oculogyric muscles. Its neurones are unique in being the only primary sensory unipolar neurones that have cell bodies in the CNS. It is the relay for the ‘jaw jerk’, the only significant supraspinal monosynaptic reflex. Nerve fibres that ascend to the mesencephalic nucleus may give collaterals to the motor nucleus of the trigeminal nerve and to the cerebellum. Most fibres that arise in the trigeminal sensory nuclei cross the midline, ascend in the trigeminal lemniscus and end in the contralateral ventral posteromedial thalamic nucleus, from which third-order neurones project to the postcentral gyrus (areas 3, 1, 2).

Motor nucleus

The motor nucleus of the trigeminal nerve is ovoid in outline and lies in the upper pontine tegmentum, under the lateral part of the floor of the fourth ventricle. It lies medial to the principal sensory nucleus from which it is separated by fibres of the trigeminal nerve. It forms the rostral part of the pharyngeal (special visceral) efferent column. The motor nucleus contains characteristic large multipolar neurones interspersed with smaller multipolar cells, organized into a number of relatively discrete subnuclei that drive individual muscles. It receives fibres from both corticonuclear tracts. These fibres leave the tracts at the nuclear level or higher in the pons (aberrant corticospinal fibres), descend in the medial lemniscus and may end on motor neurones or interneurones. The motor nucleus receives afferents from the sensory nuclei of the trigeminal nerve, possibly including some from the mesencephalic nucleus, that form monosynaptic reflex arcs for proprioceptive control of the masticatory muscles. It also receives afferents from the reticular formation, red nucleus and tectum, the medial longitudinal fasciculus and possibly from the locus coeruleus. Collectively these represent pathways by which salivary secretion and mastication may be coordinated.

Tensor tympani and stapedius reflex

Loud sound elicits reflex contraction of tensor tympani and stapedius, attenuating movement of the tympanic membrane and the ossicular chain. Afferent impulses travel in the cochlear nerve to the cochlear nuclei in the brainstem. Efferent fibres to tensor tympani arise in the motor nucleus of the trigeminal nerve and travel in the mandibular division of the nerve. Efferent fibres to stapedius originate in the facial nucleus and travel in the facial nerve.

Jaw jerk reflex

The jaw jerk reflex is a deep tendon reflex, elicited by tapping on the midline of the mandible. Rapid stretching of the muscles that close the jaw (masseter, temporalis, medial pterygoid) activates muscle spindle afferents that travel via the mandibular division of the trigeminal nerve to the mesencephalic trigeminal nucleus. Collaterals project monosynaptically to the motor nucleus of the trigeminal nerve in the pons, establishing a two-neurone reflex arc.

Intra- and extracranial course

Embryologically, each division of the trigeminal nerve is associated with a developing facial process that gives rise to a specific area of the adult face: the ophthalmic nerve is associated with the frontonasal process, the maxillary nerve with the maxillary process, and the mandibular nerve with the mandibular process.

Ophthalmic nerve

The ophthalmic division of the trigeminal nerve is a sensory nerve. It arises from the trigeminal ganglion in the middle cranial fossa and passes forwards along the lateral dural wall of the cavernous sinus, giving off three main branches, the lacrimal, frontal and nasociliary nerves, just before it reaches the superior orbital fissure. These branches subsequently travel through the orbit to supply targets that are primarily in the upper part of the face (conjunctiva, skin over the forehead, upper eyelid and much of the external surface of the nose via the supraorbital, lacrimal, infratrochlear and external nasal nerves).

Lacrimal nerve

The lacrimal nerve is the smallest of the main ophthalmic branches. It enters the orbit through the superior orbital fissure, outside the common tendinous ring and lateral to the frontal and trochlear nerves. It passes forwards along the lateral wall of the orbit on the superior border of lateral rectus, and travels through the lacrimal gland and the orbital septum to supply the conjunctiva and skin covering the lateral part of the upper eyelid. The nerve also communicates with the zygomatic branch of the maxillary nerve, which may carry some postganglionic parasympathetic fibres from the pterygopalatine ganglion to the lacrimal gland. Occasionally, the lacrimal nerve is absent, in which case it is replaced by the zygomaticotemporal nerve; the relationship is reciprocal, and when the zygomaticotemporal nerve is absent it is replaced by a branch of the lacrimal nerve. The lacrimal nerve anastomoses with filaments of the facial nerve.

Frontal nerve

The frontal nerve is the largest branch of the ophthalmic nerve. It enters the orbit through the superior orbital fissure outside the common tendinous ring, and lies between the lacrimal nerve laterally and the trochlear nerve medially. It passes forwards towards the rim of the orbit on levator palpebrae superioris, and about halfway along this course it divides into the supraorbital and supratrochlear nerves.

Supraorbital nerve

The supraorbital nerve is the larger of the two terminal branches of the frontal nerve. It continues forwards along levator palpebrae superioris until it leaves the orbit through the supraorbital notch or foramen to emerge onto the forehead. It supplies the mucous membrane that lines the frontal sinus, the conjunctiva and the skin covering the upper eyelid via palpebral branches. It ascends on the forehead with the supraorbital artery, and divides into medial and lateral branches that supply the skin of the scalp nearly as far back as the lambdoid suture. These branches are at first deep to the frontal belly of occipitofrontalis. The medial branch perforates the muscle to reach the skin, while the lateral branch pierces the epicranial aponeurosis. The postganglionic sympathetic fibres that innervate the sweat glands of the supraorbital area probably travel in the supraorbital nerve, having entered the ophthalmic nerve via its communication with the abducens nerve within the cavernous sinus.

Supratrochlear nerve

The supratrochlear nerve is the smaller terminal branch of the frontal nerve. It runs anteromedially in the roof of the orbit, passes above the trochlea for the tendon of superior oblique, and supplies a descending filament to the infratrochlear branch of the nasociliary nerve. The nerve emerges between the trochlea and the supraorbital foramen at the frontal notch, curves up on the forehead close to the bone with the supratrochlear artery, and supplies the conjunctiva and the skin of the upper eyelid. It then ascends beneath corrugator and the frontal belly of occipitofrontalis before dividing into branches that pierce these muscles to supply the skin of the lower forehead near the midline.

Nasociliary nerve

The nasociliary nerve is intermediate in size between the frontal and lacrimal nerves. It is more deeply placed in the orbit, which it enters through the common tendinous ring, lying between the two rami of the oculomotor nerve. It crosses the optic nerve with the ophthalmic artery and runs obliquely below superior rectus and superior oblique to reach the medial orbital wall, where it gives off the anterior and posterior ethmoidal nerves. The nasociliary nerve has long ciliary and infratrochlear branches and a connection with the ciliary ganglion.

Anterior ethmoidal nerve

The anterior ethmoidal nerve passes through the anterior ethmoidal foramen and canal and enters the cranial cavity. It runs forwards in a groove on the upper surface of the cribriform plate beneath the dura mater, descends through a slit lateral to the crista galli into the nasal cavity, where it occupies a groove on the internal surface of the nasal bone, and supplies the roof of the nasal cavity. It gives off medial and lateral internal nasal branches that supply the anterior and upper parts of the septum and the anterior part of the lateral wall, respectively, before emerging at the inferior margin of the nasal bone as the external nasal nerve, which descends through the lateral wall of the nose and supplies the skin of the nose below the nasal bones to the nasal tip but excluding the alar portion around the external nares: damage following nasal trauma may result in paraesthesia of the nasal tip.

Posterior ethmoidal nerve

The posterior ethmoidal nerve leaves the orbit by the posterior ethmoidal foramen and supplies the ethmoidal and sphenoidal sinuses.

Infratrochlear nerve

The infratrochlear nerve leaves the orbit below the trochlea and supplies the skin of the eyelids, the conjunctiva, lacrimal sac, lacrimal caruncle and the side of the nose above the medial canthus.

Long ciliary nerves

Two or three long ciliary nerves branch from the nasociliary nerve as it crosses the optic nerve. They accompany the short ciliary nerves, pierce the sclera near the attachment of the optic nerve, and run forwards between the sclera and choroid. They supply the ciliary body, iris and cornea, and contain postganglionic sympathetic fibres for dilator pupillae from neurones in the superior cervical ganglion. An alternative pathway for the supply of the dilator pupillae is via the sympathetic root associated with the ciliary ganglion.

Ramus communicans to the ciliary ganglion

The ramus communicans to the ciliary ganglion usually branches from the nasociliary nerve as the latter enters the orbit lateral to the optic nerve. It is sometimes joined by a filament from the internal carotid sympathetic plexus or from the superior ramus of the oculomotor nerve as it enters the posterosuperior angle of the ganglion.

Maxillary nerve

The maxillary division of the trigeminal nerve is a sensory nerve. It leaves the skull via the foramen rotundum and enters the upper part of the pterygopalatine fossa where many of its extracranial branches are given off. It crosses the pterygopalatine fossa, giving off two large ganglionic branches that contain fibres destined for the nose, palate and pharynx and that all pass through the pterygopalatine ganglion without synapsing. The maxillary nerve then inclines sharply laterally on the posterior surface of the orbital process of the palatine bone and on the upper part of the posterior surface of the maxilla in the inferior orbital fissure (which is continuous posteriorly with the pterygopalatine fossa): it lies outside the orbital periosteum, and gives off zygomatic, and then posterior superior alveolar branches. About halfway between the orbital apex and the orbital rim, the maxillary nerve turns medially to enter the infraorbital canal as the infraorbital nerve. Named branches from the main trunk are meningeal, ganglionic, zygomatic, posterior, middle and anterior superior alveolar and infraorbital nerves. Named branches from the pterygopalatine ganglion are orbital, nasopalatine, posterior superior nasal, greater (anterior) palatine, lesser (posterior) palatine and pharyngeal nerves.

Meningeal nerve

The meningeal branch of the maxillary nerve arises within the middle cranial fossa and runs with the middle meningeal vessels. It contributes to the innervation of the dura mater (see Fig. 25.3B ).

Ganglionic branches

Two ganglionic branches usually connect the maxillary nerve to the pterygopalatine ganglion.

Zygomatic nerve

The zygomatic branch of the maxillary nerve leaves the pterygopalatine fossa through the inferior orbital fissure with the maxillary nerve and lies close to the base of the lateral wall of the orbit. Soon after entering the orbit the nerve divides into two branches, the zygomaticotemporal and the zygomaticofacial nerves, which run within the orbit for only a short distance before passing onto the face through the lateral wall of the orbit. They may enter separate canals within the zygomatic bone or the zygomatic nerve itself may enter the bone before dividing.

Zygomaticotemporal nerve

The zygomaticotemporal nerve traverses a canal in the zygomatic bone and emerges into the anterior part of the temporal fossa, where it ascends between the bone and temporalis before piercing the temporal fascia about 2 cm above the zygomatic arch to supply the skin of the temple. It communicates with the facial and auriculotemporal nerves. As it pierces the deep layer of the temporal fascia, it sends a slender twig between the two layers of the fascia towards the lateral angle of the eye. It was thought that the branch carried parasympathetic postganglionic fibres from the pterygopalatine ganglion to the lacrimal gland, but it is now believed that these fibres innervate the lacrimal gland directly.

Zygomaticofacial nerve

The zygomaticofacial nerve traverses the inferolateral angle of the orbit and emerges on the face through a foramen in the zygomatic bone. It perforates orbicularis oculi to supply the skin on the prominence of the cheek and forms a plexus with zygomatic branches of the facial nerve and palpebral branches of the maxillary nerve. The nerve is absent occasionally.

Infraorbital nerve

The infraorbital nerve can be regarded as the terminal branch of the maxillary nerve. It leaves the pterygopalatine fossa to enter the orbit at the inferior orbital fissure and at first it lies in the infraorbital groove on the floor of the orbit. As it approaches the rim of the orbit it runs into the infraorbital canal through which it passes to emerge onto the face at the infraorbital foramen, where it lies between levator labii superioris and levator anguli oris. It gives off palpebral, nasal and superior labial branches. The palpebral branches ascend deep to orbicularis oculi, pierce the muscle to supply the skin in the lower eyelid, and join the facial and zygomaticofacial nerves near the lateral canthus. Nasal branches supply the skin of the side of the nose, the vestibule and the movable part of the nasal septum, and join the external nasal branch of the anterior ethmoidal nerve. Superior labial branches, which are large and numerous, descend behind levator labii superioris to supply the skin of the anterior part of the cheek and upper lip. They are joined by branches from the facial nerve to form the infraorbital plexus.

Orbital branches

Fine orbital branches enter the orbit through the inferior orbital fissure and supply orbital periosteum. Some fibres also pass through the posterior ethmoidal foramen to supply the sphenoidal and ethmoidal sinuses. The orbital branches probably join branches of the internal carotid nerve to form a ‘retro-orbital’ plexus from which orbital structures such as the lacrimal gland and orbitalis receive an autonomic innervation.

Posterior superior nasal nerves (lateral and medial)

The posterior superior alveolar nerves enter the back of the nasal cavity through the sphenopalatine foramen. Lateral posterior superior nasal nerves innervate the mucosa covering the posterior part of the superior and middle nasal turbinates (conchae) and lining the posterior ethmoidal sinuses. Two or three medial posterior superior nasal nerves cross the nasal roof below the opening of the sphenoidal sinus to supply the mucosa of the posterior part of the roof and of the nasal septum.

Nasopalatine nerve

The nasopalatine nerve leaves the pterygopalatine fossa through the sphenopalatine foramen and enters the nasal cavity. It passes across the cavity to the back of the nasal septum, runs downwards and forwards on the septum in a groove in the vomer, and then turns down through the incisive fossa in the anterior part of the hard palate to enter the roof of the mouth. When an anterior and a posterior incisive foramen exist in this fossa, the left nasopalatine nerve typically passes through the anterior foramen, and the right nerve passes through the posterior foramen. The nasopalatine nerve supplies the inferior part of the nasal septum and the anterior part of the hard palate, where it communicates with the greater palatine nerve.

Surgical division of the nasopalatine nerve, for example during the removal of an ectopic canine tooth, causes no obvious sensory deficit in the anterior part of the palate, which suggests that either the territory of the greater palatine nerve reaches as far forwards as the gingivae lingual to the incisor teeth or that the nasopalatine nerve has significant regenerative potential.

Palatine nerves (greater and lesser)

The greater and lesser palatine nerves pass downwards from the pterygopalatine ganglion through the greater palatine canal. The greater palatine nerve descends through the greater palatine canal, emerges on the hard palate from the greater palatine foramen and runs forwards in a groove on the inferior surface of the bony palate almost to the incisor teeth. It supplies the gingivae, mucosa and glands of the hard palate and also communicates with the terminal filaments of the nasopalatine nerve. In the greater palatine canal it gives off posterior inferior nasal branches that emerge through the perpendicular plate of the palatine bone and ramify over the inferior nasal turbinate (concha) and walls of the middle and inferior meatuses. As it leaves the greater palatine canal, it gives off branches which are distributed to both surfaces of the adjacent part of the soft palate.

The lesser (middle and posterior) palatine nerves are much smaller than the greater palatine nerve. They descend through the greater palatine canal, from which they diverge low down to emerge through the lesser palatine foramina in the tubercle or pyramidal process of the palatine bone. They innervate the uvula, tonsil and soft palate.

Fibres conveying taste impulses from the palate probably pass via the palatine nerves to the pterygopalatine ganglion. They pass through the ganglion without synapsing, and leave via the greater petrosal nerve. Their cell bodies are located in the facial ganglion and their central processes pass via the sensory root of the facial nerve (nervus intermedius) to the gustatory nucleus in the nucleus of the tractus solitarius.

Pharyngeal nerve

The pharyngeal branch of the maxillary nerve leaves the pterygopalatine ganglion posteriorly. It passes through the palatovaginal canal with the pharyngeal branch of the maxillary artery and supplies the mucosa of the nasopharynx behind the pharyngotympanic tube.

Superior alveolar nerves

The teeth in the upper jaw are supplied by the three superior alveolar (dental) nerves that arise from the maxillary nerve either in the pterygopalatine fossa or in the infraorbital groove and canal.

The posterior superior alveolar (dental) nerve leaves the maxillary nerve in the pterygopalatine fossa and runs anteroinferiorly to pierce the infratemporal surface of the maxilla, at which point it is possible to perform a local anaesthetic block that will anaesthetize the pulps of the premolar and molar ipsilateral teeth; it then descends under the mucosa of the maxillary sinus. After supplying the lining of the sinus, the nerve divides into small branches that link up as the molar part of the superior alveolar plexus, supplying twigs to the molar teeth. It also supplies a branch to the upper molar gingivae and the adjoining part of the cheek.

The middle superior alveolar (dental) nerve arises from the infraorbital nerve as it runs in the infraorbital groove, and runs downwards and forwards in the lateral wall of the maxillary sinus. It ends in small branches that link up with the superior dental plexus, supplying small rami to the upper premolar teeth. This nerve is variable; it may be duplicated, triplicated, or absent.

The anterior superior alveolar (dental) nerve leaves the lateral side of the infraorbital nerve near the midpoint of its canal and traverses the canalis sinuosus in the anterior wall of the maxillary sinus. It curves first under the infraorbital foramen, then passes medially towards the nose, and finally turns downwards and divides into branches to supply the incisor and canine teeth. It assists in the formation of the superior dental plexus and gives off a nasal branch that passes through a minute canal in the lateral wall of the inferior meatus to supply the mucous membrane of the anterior area of the lateral wall as high as the opening of the maxillary sinus, and the floor of the nasal cavity near the anterior nasal spine. It communicates with the nasal branches of the pterygopalatine ganglion and finally emerges near the root of the anterior nasal spine to supply the adjoining part of the nasal septum.

Mandibular nerve

The mandibular division of the trigeminal nerve is the largest division of the trigeminal nerve and is a mixed nerve. Its sensory branches supply the mandibular teeth and gingivae, the skin in the temporal region, part of the auricle and the lower lip, the lower part of the face and the mucosa of the anterior two-thirds (presulcal part) of the tongue and the floor of the oral cavity. The motor branches innervate the muscles of mastication, tensor veli palatini, tensor tympani, anterior belly of digastric and mylohyoid. The large sensory root emerges from the lateral part of the trigeminal ganglion and exits the cranial cavity through the foramen ovale. The small motor root passes under the ganglion and through the foramen ovale to unite with the sensory root just outside the skull. As it descends from the foramen ovale, the mandibular nerve is usually around 4 cm from the surface and a little anterior to the neck of the mandible. It immediately passes between tensor veli palatini, which is medial, and lateral pterygoid, which is lateral, gives off a meningeal branch and the nerve to medial pterygoid from its medial side, and then divides into a small anterior and large posterior trunk. The anterior trunk gives off branches to the four main muscles of mastication and a buccal branch which is sensory to the cheek. The posterior trunk gives off three main sensory branches, the auriculotemporal, lingual and inferior alveolar nerves, and motor fibres to supply mylohyoid and the anterior belly of digastric.

Meningeal branch (nervus spinosus)

The meningeal branch re-enters the middle cranial fossa through the foramen spinosum with the middle meningeal artery. It divides into anterior and posterior branches that accompany the main divisions of the middle meningeal artery and supply the dura mater in the middle cranial fossa and, to a lesser extent, in the anterior fossa and calvaria (see Fig. 25.3B ).

Nerve to medial pterygoid

The nerve to medial pterygoid is a slender ramus that enters the deep aspect of the muscle. It supplies one or two filaments that pass through the otic ganglion without interruption to supply tensor tympani and tensor veli palatini.

Anterior trunk of mandibular nerve

The anterior trunk of the mandibular nerve gives rise to the buccal nerve, which is sensory, and the masseteric, deep temporal and lateral pterygoid nerves, which are all motor.

Buccal nerve

The buccal nerve passes between the two heads of lateral pterygoid, giving off motor fibres to the muscle as it passes through it. The nerve then descends deep to the tendon of temporalis, passes laterally anterior to masseter, and anastomoses with the buccal branches of the facial nerve. It may give off the anterior deep temporal nerve. The buccal nerve supplies sensation to the skin over the anterior part of buccinator, the corresponding buccal mucous membrane and the posterior part of the buccal gingivae opposite the second and third molar teeth. It may also give off a branch that passes through a small retromolar foramen to supply the molar teeth.

Nerve to masseter

The nerve to masseter passes laterally above lateral pterygoid onto the skull base, anterior to the temporomandibular joint and posterior to the tendon of temporalis. It crosses the posterior part of the mandibular notch with the masseteric artery and ramifies on and enters the deep surface of masseter. It provides articular branches to the temporomandibular joint.

Deep temporal nerves

There are usually two deep temporal nerves, anterior and posterior, although there may be a middle branch. They pass above lateral pterygoid to enter the deep surface of temporalis. The anterior branch may arise as a branch of the buccal nerve or from the nerve to masseter; the small posterior nerve sometimes arises in common with the nerve to masseter. Branches from the buccal nerve and the nerve to masseter may supply the inferior aspect of temporalis independently from the deep temporal nerves.

Nerve to lateral pterygoid

The nerve to lateral pterygoid enters the deep surface of the muscle. It may arise separately from the anterior division of the mandibular nerve or from the buccal nerve.

Posterior trunk of mandibular nerve

The posterior trunk of the mandibular nerve is larger than the anterior and is mainly sensory, although it receives fibres from the motor root for the nerve to mylohyoid. It divides into auriculotemporal, lingual and inferior alveolar (dental) nerves.

Auriculotemporal nerve

The auriculotemporal nerve usually has two roots that encircle the middle meningeal artery. It runs back under lateral pterygoid on the surface of tensor veli palatini, passes between the sphenomandibular ligament and the neck of the mandible, and then runs laterally behind the temporomandibular joint related to the upper part of the parotid gland. Emerging from behind the joint, it ascends over the posterior root of the zygoma, posterior to the superficial temporal vessels, and divides into superficial temporal branches. It communicates with the facial nerve and otic ganglion. The rami to the facial nerve, usually two, pass anterolaterally behind the neck of the mandible to join the facial nerve at the posterior border of masseter. Filaments from the otic ganglion join the roots of the auriculotemporal nerve close to their origin. The auriculotemporal nerve emerges onto the face behind the temporomandibular joint within the superior surface of the parotid gland, ascends posterior to the superficial temporal vessels, passes over the posterior root of the zygoma, and divides into superficial temporal branches. The cutaneous branches of the auriculotemporal nerve supply the tragus and part of the adjoining auricle of the ear and the posterior part of the temple. The nerve may be damaged during parotid gland surgery, resulting in impaired sensation over the tragus and temple. The auriculotemporal nerve communicates with the temporofacial division of the facial nerve, usually by two rami that pass anterolaterally behind the neck of the mandible. These communications anchor the facial nerve close to the lateral surface of the condylar process of the mandible and limit its mobility during surgery. Communications with the temporal and zygomatic branches of the facial nerve loop around the transverse facial and superficial temporal vessels.

Lingual nerve

The lingual nerve is sensory to the mucosa of the floor of the mouth, mandibular lingual gingivae and mucosa of the presulcal part of the tongue (excluding the circumvallate papillae, which are supplied by the glossopharyngeal nerve). It also carries postganglionic parasympathetic fibres from the submandibular ganglion to the sublingual and anterior lingual glands. It arises from the posterior trunk of the mandibular nerve and at first runs beneath lateral pterygoid and superficial to tensor veli palatini, where it is joined by the chorda tympani branch of the facial nerve, and often by a branch of the inferior alveolar nerve. Emerging from under cover of lateral pterygoid, the lingual nerve then runs downwards and forwards on the surface of medial pterygoid, and is thus carried progressively closer to the medial surface of the mandibular ramus. It becomes intimately related to the bone a few millimetres below and behind the junction of the vertical ramus and horizontal body of the mandible. Here it lies anterior to, and slightly deeper than, the inferior alveolar (dental) nerve. It next passes below the mandibular attachment of the superior pharyngeal constrictor and pterygomandibular raphe, closely applied to the periosteum of the medial surface of the mandible, until it lies opposite the posterior root of the lower third molar tooth, where it is covered only by the gingival mucoperiosteum. At this point it usually lies approximately 3 mm below the alveolar crest with a mean distance from the bone of 2 mm: it may be in direct contact with the bone (25%) or lie above the alveolar crest (15%). The nerve next passes medial to the mandibular origin of mylohyoid, and this carries it progressively away from the mandible, separating it from the alveolar bone covering the mesial root of the lower third molar tooth. It then passes downwards and forwards on the deep surface of mylohyoid (i.e. the surface nearer the mucosa covering the floor of the mouth), crossing the lingual sulcus beneath the mucosa. In this position, the lingual nerve lies on the deep portion of the submandibular gland, which bulges over the top of the posterior border of mylohyoid. The nerve passes below the submandibular duct, which crosses it from medial to lateral, and curves upwards, forwards and medially to enter the tongue by medial and lateral branches. Within the tongue, the medial branch sends small branches to the medial part of the ventrolateral tongue, and the lateral branch runs along the lateral border of the tongue and sends larger branches to innervate the mucosa of the anterior tip of the tongue. The lingual nerve is connected to the submandibular ganglion by two or three branches. It forms connecting loops with twigs of the hypoglossal nerve at the anterior margin of hyoglossus ( ) and within the tongue ( ).

The lingual nerve is at risk during surgical removal of (impacted) lower third molars: patients occasionally develop lingual sensory disturbance postoperatively but this is rarely persistent ( ). The nerve is also at risk during operations to remove the submandibular salivary gland because the submandibular duct must be dissected from the lingual nerve, and also because its connection to the submandibular ganglion pulls it into the operating field.

Inferior alveolar (dental) nerve

The inferior alveolar nerve descends behind lateral pterygoid. At the lower border of the muscle the nerve passes between the sphenomandibular ligament and the mandibular ramus and enters the mandibular canal via the mandibular foramen. Just before entering the mandibular canal, the inferior alveolar nerve gives off a small mylohyoid branch that also carries fibres that innervate the anterior belly of digastric. Below lateral pterygoid, the nerve is accompanied by the inferior alveolar artery, a branch of the first part of the maxillary artery, which enters the canal with its associated veins. In the mandibular canal, the inferior alveolar nerve runs downwards and forwards, generally below the apices of the teeth until it is below the first and second premolars, where it divides into terminal incisive and mental branches. The incisive branch continues forwards in a bony canal or in a plexiform arrangement, giving off branches to the first premolar, canine and incisor teeth, and to the associated labial gingivae. The lower central incisor teeth receive a bilateral innervation, fibres probably crossing the midline within the periosteum to re-enter the bone via numerous canals in the labial cortical plate. The mental nerve passes upwards, backwards and outwards to emerge from the mandible via the mental foramen between and just below the apices of the premolar teeth. It immediately divides into three branches, two of which pass upwards and forwards to form an incisor plexus labial to the teeth that supplies the labial gingivae (and probably the periosteum). From this plexus and from the dental branches, fibres turn downwards and then lingually to emerge on the lingual surface of the mandible on the posterior aspect of the mental symphysis or opposite the premolar teeth, probably communicating with the lingual or mylohyoid nerves. The third branch of the mental nerve passes through the intermingled fibres of depressor anguli oris and platysma to supply the skin of the lower lip and chin. Branches of the mental nerve communicate with terminal filaments of the mandibular branch of the facial nerve.

Nerve to mylohyoid

The nerve to mylohyoid is given off from the inferior alveolar nerve just before that nerve enters the mandibular canal. It pierces the sphenomandibular ligament and enters a shallow groove on the medial surface of the mandible, following a course roughly parallel to its parent nerve. It passes below the origin of mylohyoid to lie on the superficial surface of the muscle, between it and the anterior belly of digastric, both of which it supplies. Although primarily a motor nerve supplying mylohyoid and the anterior belly of digastric, the mylohyoid nerve carries some sensory fibres that may enter the mandible via one or more small retromental foramina in the vicinity of the genial tubercles to supply some sensation to the anterior teeth, and it may also give a few filaments to supply the skin over the point of the chin.

Dental extractions

Variations in the fascicular organization of the inferior alveolar nerve are clinically important when extracting impacted third molars. The nerve may appear as a single bundle lying a few millimetres below the roots of the teeth, or it may lie much lower and almost reach the lower border of the bone, so that it gives off a variable number of large rami that pass anterosuperiorly towards the roots before dividing to supply the teeth and interdental septa. Only rarely is it plexiform. The nerve may lie on the lingual or buccal side of the mandible (slightly more commonly on the buccal side). Even when the third molar tooth is in a normal position, the nerve may be so intimately related to it that it grooves its root. Exceptionally, the nerve may be similarly related to the second molar tooth.

Failure to achieve complete anaesthesia to the mandibular teeth following an inferior alveolar nerve block (which should anaesthetize the nerve before it enters the mandibular canal) suggests that there are alternative pathways for sensory nerves to innervate the teeth. Knowledge of these pathways is important in helping clinicians to obtain complete dental anaesthesia. There are anatomical variations in the distribution of the inferior alveolar nerve, the buccal branch of the mandibular nerve, the mylohyoid nerve, the lingual nerve, the deep temporal nerves and the cervical plexus. The inferior alveolar nerve may give multiple branches before entering the mandibular canal that may re-enter the mandible via small accessory foramina in the retromolar region to supply sensation to the molar teeth: one or more of these branches may escape anaesthesia following inferior alveolar nerve block. If a successful inferior alveolar nerve block does not completely anaesthetize teeth in a jaw quadrant, additional local injections of anaesthetic solution around the tooth in question may be necessary to block alternative pathways. There is also the possibility of overlap across the midline involving the incisive and mylohyoid nerves. In children, there is evidence to suggest that the lingula is variable in position and its position from the occlusion plane increases with age: it is therefore not considered an appropriate landmark for inferior alveolar nerve block.

Neural connections between branches of trigeminal and other nerves

Communications exist between the inferior alveolar and lingual nerves. If the lingual nerve is not anaesthetized during an inferior alveolar nerve block, this pathway could account for any remaining sensation in the teeth. Although the deep temporal nerves are primarily motor, it has been suggested that terminal branches may pass from the substance of temporalis to enter the mandible through foramina in the retromolar region to supply some sensation to molar teeth. In the cervical plexus, the great auricular nerve, formed by the anterior primary rami of the second and third cervical spinal nerves, supplies skin over the angle of the mandible: it is possible that this nerve may provide a branch that penetrates the mandible to supply one or more of the cheek teeth.

Abducens Nerve

The abducens nerve is the sixth cranial nerve and innervates the ipsilateral lateral rectus.

Abducens nucleus and central connections

The abducens nucleus lies in the floor of the fourth ventricle, just lateral to the midline, immediately subjacent to the facial colliculus and slightly lateral to the medial longitudinal fasciculus (see Fig. 28.4 ). The abducens nucleus is composed of approximately 6500 neurones that align with the trochlear, oculomotor and hypoglossal nuclei to form the general somatic efferent column (GSE). Within the nucleus, large motor neurones are intermixed with small multipolar interneurones, the latter being most heavily concentrated in the lateral and ventral aspects of the nucleus. The abducens nucleus receives afferent connections from multiple sources, including corticonuclear fibres (principally contralateral, some of the fibres are aberrant corticospinal fibres that descend from the midbrain to this level in the medial lemniscus); the medial longitudinal fasciculus (connecting the abducens nucleus to oculomotor, trochlear and vestibular nuclei); the tectobulbar tract (from the deep layers of the superior colliculus); the paramedian pontine reticular formation (rostral and caudal to the nucleus); the nucleus prepositus hypoglossi; and the contralateral medullary reticular formation.

Neural Control of Gaze

Although the detailed anatomical substrates for the different types of eye movement differ, they share common neural circuitry that lies mainly in the pons and midbrain, for horizontal and vertical gaze movements, respectively (see Fig. 44.12 ). The common element for all types of horizontal gaze movements is the abducens nucleus. It contains motor neurones that innervate the ipsilateral lateral rectus and interneurones that project via the medial longitudinal fasciculus to the contralateral oculomotor nucleus, which controls medial rectus. A lesion of the abducens nucleus leads to a total loss of ipsilateral horizontal conjugate gaze. A lesion of the medial longitudinal fasciculus produces slowed or absent adduction of the ipsilateral eye, usually associated with jerky movements (nystagmus) of the contralateral abducting eye, a syndrome called internuclear ophthalmoplegia (Leigh and Zee 2015). The gaze motor command involves specialized areas of the reticular formation of the brainstem, which receive a variety of supranuclear inputs. The main region for the generation of horizontal saccades is the paramedian pontine reticular formation, located on each side of the midline in the central paramedian part of the tegmentum, and extending from the pontomedullary junction to the pontopeduncular junction. Each paramedian pontine reticular formation contains excitatory neurones (referred to as ‘burst’ neurones) that discharge at high frequencies just prior to and during ipsilateral saccades. Excitatory burst neurones make monosynaptic connections with the ipsilateral abducens nucleus. Omnipause neurones in the pontine nucleus raphe interpositus, located in the midline between the rootlets of the abducens nerves, discharge tonically during fixation but stop firing immediately prior to a saccade. They appear to exert an inhibitory influence on the burst neurones and act as a switch to change from fixation to saccadic mode (Ramat et al 2007).

The tonic activity of neurones in the nucleus prepositus hypoglossi and medial vestibular nucleus is thought to provide an eye position signal to maintain the eccentric position of the eye against the viscoelastic forces in the orbit. These forces tend to move the eyeball back to looking straight ahead, i.e. the primary position, after a saccade. Vestibular nuclei and the perihypoglossal complex project directly to the abducens nuclei: these projections probably also carry smooth pursuit signals, via the cerebellum.

The final common pathway for vertical gaze movements is formed by the oculomotor and trochlear nuclei. The rostral interstitial nucleus of the medial longitudinal fasciculus contains both excitatory burst neurones that discharge in relation to up-and-down vertical saccadic movements and project to motor neurones involved in vertical gaze, and omnipause neurones that act similarly to those for horizontal gaze. The rostral interstitial nucleus of the medial longitudinal fasciculus projects through the posterior commissure to its equivalent on the other side of the mesencephalon for upgaze, as well as directly to the ipsilateral oculomotor and trochlear nuclei for downgaze (see Fig. 44.12 ). Neurones in and around the interstitial nucleus of Cajal, which lies slightly caudal to the rostral interstitial nucleus of the medial longitudinal fasciculus, provide signals for vertical gaze holding. Vertical gaze palsies can affect upgaze, downgaze or both. Lesions within the posterior commissure predominantly give rise to disturbances in upgaze, associated with other signs of dorsal midbrain syndrome, e.g. pupillary abnormalities (light–near dissociation). Discrete lesions placed more ventrally in the region of the rostral interstitial nucleus of the medial longitudinal fasciculus may cause mixed up-and-down, or mainly downgaze disturbances (Leigh and Zee 2015).

Intra- and extracranial course

The axons of the motor neurones in the abducens nucleus spread diffusely from the ventromedial aspect of the nucleus. Fifteen to twenty fascicles pass ventrally, laterally and caudally, directly through the intervening reticular formation, trapezoid body, medial lemniscus, pontocerebellar fibres, pontine nuclei and descending corticospinal tracts to emerge from the brainstem at the pontomedullary sulcus about 1–2 cm lateral to the midline. In their course, the rootlets typically coalesce to emerge from the brainstem as a compact bundle of fibres. Occasionally all or some of the rootlets may emerge rostral to the pontomedullary sulcus. The nerve runs upward, anteriorly and slightly laterally across the prepontine cistern between the pons and clivus, pierces the inner layer of dura mater and runs through the inferior venous compartment of the petroclival venous confluence in a bow-shaped canal, Dorello’s canal. It then bends sharply across the upper border of the petrous part of the temporal bone to enter the cavernous sinus, where it lies lateral to the cavernous segment of the internal carotid artery but medial to the oculomotor, trochlear, ophthalmic and maxillary nerves that also course through the sinus (but merely invaginate the lateral dural wall of the sinus). The abducens nerve enters the orbit through the inferomedial part of the superior orbital fissure, within the common tendinous ring, at first below, and then between, the two divisions of the oculomotor nerve and lateral to the nasociliary nerve (see Fig. 44.15 ). It passes forwards to enter the medial (ocular) surface of lateral rectus, most often in the posterior one-third of the muscle ( ).

Facial nerve

The facial nerve is the seventh cranial nerve. It contains special visceral/branchial efferent (SVE) motor axons that supply muscles derived from the second branchial arch, namely the mimetic facial muscles, buccinator, stapedius, platysma, the posterior belly of digastric and stylohyoid. The craniofacial muscles innervated by the facial nerve typically lack muscle spindles: proprioceptive information may be mediated by modified cutaneous mechanoreceptors or capsular corpuscle-like structures within the muscles. The facial nerve contains two, possibly three, types of sensory axon. Special visceral afferent (SVA) axons carry taste from the anterior two-thirds of the tongue via the chorda tympani, and general sensory afferent (GSA) axons carry cutaneous sensation, including pain, from the posterior aspect of the external acoustic meatus, with the caveat that the innervation of this region is complex and largely contributed by Arnold’s nerve: these axons are thought to be responsible for coughing during cerumen removal, idiopathic otalgia, Hitselberger’s sign and to be the site of vesicular eruptions in Ramsay Hunt syndrome.

General visceral efferent (GVE) fibres in the facial nerve arise from preganglionic parasympathetic neurones in the superior salivatory nucleus and pass via either the chorda tympani to the submandibular ganglion or the greater petrosal nerve and the nerve of the pterygoid canal to the pterygopalatine ganglion. The GVE, SVA, GSA axons form the nervus intermedius (intermediate nerve, nerve of Wrisberg).

Facial nucleus and central connections

The facial (motor) nucleus is situated caudal, ventral and lateral to the abducens nucleus. It lies within the caudal pontine reticular formation, ventromedial to the spinal tract and nucleus of the trigeminal nerve (see Figure 28.11, Figure 28.13 ) ( ). Groups of neurones form columns that innervate individual muscles or that correspond to branches of the facial nerve. The facial nucleus receives corticonuclear fibres for volitional control of the facial musculature. Within the nucleus, neurones that innervate the muscles of the scalp and upper face are innervated by bilateral corticonuclear fibres. They lie dorsal to the neurones that innervate the muscles of the lower face, which receive a predominately contralateral innervation. Clinically, therefore, upper and lower motor neurone lesions affecting the facial nerve can be differentiated because upper motor neurone lesions cause paralysis that is confined to the contralateral lower face (supranuclear facial palsy), whilst lower motor neurone lesions cause complete ipsilateral facial paralysis (Bell’s palsy). The facial nucleus also receives ipsilateral afferents from the nucleus solitarius and from the sensory nucleus of the trigeminal nerve, establishing important reflex connections.

Efferent fibres that arise from large motor neurones of the facial nucleus form the motor root of the facial nerve. At first, the facial fibres course dorsomedially towards the floor of the fourth ventricle as a diffuse spread of fibres that pass dorsal and medial to the abducens nucleus. They then coalesce into a defined fibre bundle that re-curves dorsally and laterally over the rostral pole of the abducens nucleus, creating a bulge in the floor of the fourth ventricle, the facial colliculus (see Fig. 28.11 ). The fibres descend ventrolaterally through the reticular formation and pass between their nucleus of origin medially and the nucleus of the spinal tract of the trigeminal nerve laterally. The facial nerve emerges from the brainstem at the supraolivary fossette, between the olive and restiform body, and passes laterally through the cerebellopontine angle cistern to the porus acusticus of the internal acoustic meatus (internal auditory canal).

Intracranial and intratemporal course

The facial nerve has intracranial (cisternal), intratemporal and extratemporal portions. The intratemporal portion is further divided into meatal, labyrinthine, tympanic (horizontal) and mastoid (vertical) segments (see Fig. 42.12 ). The cisternal portion emerges from the ventrolateral aspect of the caudal border of the pons. The meatal portion enters the porus acusticus of the internal acoustic meatus accompanied by the nervus intermedius, the vestibulocochlear nerve and the labyrinthine vessels. In this part of its course, the facial nerve lacks a fibrous sheath or endoneurium and is surrounded by a thin layer of arachnoid. The motor root, which supplies the muscles of the face, and the nervus intermedius, which contains sensory fibres concerned with the perception of taste and parasympathetic (secretomotor) fibres to various glands, are still separate components; they usually merge within the meatus. The labyrinthine segment runs from the fundus of the internal acoustic meatus to the geniculate ganglion, where the nerve makes the first bend or genu (see Fig. 42.12 ). The labyrinthine segment is the shortest and narrowest part of the facial nerve; its lack of anastomosing arterial cascades renders it susceptible to vascular compression. The tympanic part initially curves around the oval window niche, then lies just anterior and inferior to the lateral semicircular canal and bends again at the second genu to become the vertical or mastoid part. The mastoid part is the longest of the petrous segments and runs from the pyramidal process to the stylomastoid foramen. The tympanic and mastoid segments of the facial nerve are supplied by the facial arch, formed by the petrosal branch of the middle meningeal artery and the stylomastoid branch of the posterior auricular artery.

The branches that arise from the facial nerve within the temporal bone can be divided into those that come from the geniculate ganglion (greater petrosal nerve) and those that arise within the facial canal (nerve to stapedius and chorda tympani).

Greater petrosal nerve

The greater (superficial) petrosal nerve, a branch of the nervus intermedius, is the main branch from the geniculate ganglion. It passes anteriorly, receives a branch from the tympanic plexus and traverses a hiatus on the anterior surface of the petrous part of the temporal bone. It enters the middle cranial fossa, runs forwards in a groove on the bone above the lesser petrosal nerve, and then passes beneath the trigeminal ganglion to reach the foramen lacerum. Here it is joined by the deep petrosal nerve from the internal carotid sympathetic plexus, to become the nerve of the pterygoid canal (Vidian nerve). The greater petrosal nerve contains parasympathetic fibres destined for the pterygopalatine ganglion, and taste fibres from the palate. The geniculate ganglion also communicates with the lesser petrosal nerve.

Nerve to stapedius

The nerve to stapedius arises from the facial nerve in the facial nerve canal behind the pyramidal eminence of the posterior wall of the tympanic cavity and passes forwards through a small canal to reach the muscle.

Chorda tympani

The chorda tympani (see Figure 42.11, Figure 42.12 ) leaves the facial nerve some 6 mm above the stylomastoid foramen and runs anterosuperiorly in a canal to enter the tympanic cavity via the posterior canaliculus. It then curves anteriorly in the substance of the tympanic membrane between its mucous and fibrous layers, and crosses medial to the upper part of the handle of the malleus to the anterior wall of the tympanic cavity, where it enters the anterior canaliculus. It exits the skull and enters the infratemporal fossa by passing through the medial end of the petrotympanic fissure posterior to the capsule of the temporomandibular joint. It next descends medial to the spine of the sphenoid bone, which it sometimes grooves, lying posterolateral to tensor veli palatini. It is crossed medially by the middle meningeal artery, the roots of the auriculotemporal nerve and the inferior alveolar nerve (see Fig. 38.23 ) and joins the posterior aspect of the lingual nerve at an acute angle. The chorda tympani carries taste fibres from the anterior two-thirds of the tongue (but not the from the circumvallate papillae) and efferent preganglionic parasympathetic secretomotor fibres destined for the submandibular ganglion in the floor of the mouth.

Dehiscences of the facial nerve canal

The facial nerve may be somewhat variable in its anatomical course through the temporal bone ( , ). It may split into two or three strands, starting at the geniculate ganglion, and then make its way across the promontory to the stylomastoid foramen, or pass a few millimetres posteriorly to its second genu, before it turns inferiorly posterior to the fossa incudis, a position where it is particularly vulnerable during surgical exploration of the mastoid antrum. The more proximal the division into strands, the more bizarre is the subsequent course. More distal bifurcations pass either side of the fenestra vestibuli. It may be dehiscent, particularly in its second part, when it occasionally overhangs the stapes, or run inferior to the stapedial superstructure, a position that renders it vulnerable during surgery to the stapes ( ). The motor fibres to the face may be carried through the chorda tympani, which is then enlarged. When this is the case, the distal facial nerve dwindles to a fibrous strand in a narrowed stylomastoid foramen. In chronic bone disease in the tympanic cavity, the facial nerve may be exposed in its canal. Inflammation may lead to facial paralysis of the infranuclear or lower motor neurone type.

Extratemporal course

The facial nerve emerges from the base of the skull at the stylomastoid foramen and almost immediately gives off the nerves to the posterior belly of digastric and stylohyoid and the posterior auricular nerve, which supplies the occipital belly of occipitofrontalis and some of the intrinsic auricular muscles (see Fig. 36.23 ). The nerve next enters the parotid gland high up on its posteromedial surface and passes forwards and downwards behind the mandibular ramus. Within the substance of the gland, it branches into superior (temporofacial) and inferior (cervicofacial) trunks, usually just behind and superficial to the retromandibular vein. The trunks branch further to form a parotid plexus (pes anserinus). Five main terminal branches arise from the plexus, diverge within the gland and leave by its anteromedial surface, medial to its anterior margin, to supply the muscles of facial expression (see Fig. 36.23 ). Six distinctive anastomotic patterns were originally classified by and these are illustrated in Fig. 36.24 . Numerous microdissection studies have demonstrated considerable individual variation in branching patterns and anastomoses between branches, both within the parotid and on the face (e.g. ): the account that follows is therefore an overview. In surgical terms, these anastomoses are important and presumably explain why accidental or deliberate division of a small branch often fails to result in the expected facial nerve weakness.

The temporal branch usually divides into anterior and posterior rami soon after piercing the parotidomasseteric fascia below the zygomatic arch; there is often a middle (frontal) ramus. These rami cross the arch in subcutaneous tissue, lying in the subgaleal space: their course is extremely variable.

Twigs supply intrinsic muscles on the lateral surface of the auricle and the anterior and superior auricular muscles. They communicate with the zygomaticotemporal branch of the maxillary nerve and the auriculotemporal branch of the mandibular nerve. The more anterior branches supply the frontal belly of occipitofrontalis, orbicularis oculi and corrugator and join the supraorbital and lacrimal branches of the ophthalmic nerve.

Zygomatic branches

Zygomatic branches are generally multiple. They cross the zygomatic bone to the lateral canthus of the eye and supply orbicularis oculi and may also supply muscles innervated by the buccal branch. Twigs communicate with filaments of the lacrimal nerve and the zygomaticofacial branch of the maxillary nerve.

Buccal nerve

The buccal branch of the facial nerve is usually single. It has a close relationship to the parotid duct for about 2.5 cm after emerging from the parotid gland and typically lies below the duct. Superficial branches run beneath the subcutaneous fat and the superficial musculo-aponeurotic system (SMAS). Some branches pass deep to procerus and join the infratrochlear and external nasal nerves. Upper deep branches supply zygomaticus major and levator labii superioris and form an infraorbital plexus with the superior labial branches of the infraorbital nerve. They also supply levator anguli oris, zygomaticus minor, levator labii superioris alaeque nasi and the small nasal muscles (these branches are sometimes described as lower zygomatic branches). Lower deep branches supply buccinator and orbicularis oris and communicate with filaments of the buccal branch of the mandibular nerve.

Marginal mandibular nerve

There are usually two marginal mandibular nerves. They run forwards towards the angle of the mandible under platysma, and then turn upwards across the body of the mandible to pass under depressor anguli oris. The branches supply risorius and the muscles of the lower lip and chin. Filaments communicate with the mental nerve. The marginal mandibular branch has an important surgical relationship with the inferior border of the mandible.

Cutaneous branches

Cutaneous branches of the facial nerve accompany the auricular branch of the vagus and probably supply small areas on both aspects of the auricle, in the depression of the concha and over its eminence.

The facial nerve has intra- or extracranial connections with the cutaneous branches of all three divisions of the trigeminal nerve (including branches of the auriculotemporal, buccal, mental, lingual, infraorbital, zygomatic and ophthalmic nerves); with branches of the vestibulocochlear, glossopharyngeal and vagus nerves; and with branches of the cervical plexus (including the great auricular, greater and lesser occipital and transverse cervical nerves). These cutaneous connections are significant in facilitating the perineural spread of tumours that arise either within the parotid or on the face and explains why some patients with perineural spread from cutaneous or parotid malignancies may present with vocal cord palsy.

Cervical branch

The cervical branch of the facial nerve emerges from the lower part of the parotid gland and runs anteroinferiorly under platysma to the anterior neck. Typically single, it supplies platysma and communicates with the transverse cutaneous cervical nerve.

Lesions of the facial nerve

Facial paralysis may be due to an upper motor neurone lesion (when frontalis is partially spared because of the bilateral cortical representation of the muscles of the upper part of the face) or a lower motor neurone lesion (when all branches may be involved). Lower motor neurone facial paralysis is most commonly idiopathic (Bell’s palsy). There are a large number of other recognized causes that include geniculate herpes zoster (Ramsay Hunt syndrome), and malignant tumours of the parotid gland and temporal bone. Iatrogenic damage also accounts for a significant number of palsies, most often acquired in vestibular schwannoma, parotid and middle ear or mastoid surgery.

Bell’s palsy

Bell’s palsy is an idiopathic lower motor neurone facial palsy. It may be complete or partial, and is characterized by a flaccid paralysis of the ipsilateral mimetic facial muscles (muscles of facial expression); decreased lacrimation in the ipsilateral eye (which is controlled by neurones in the greater petrosal nerve); and hyperacusis or decreased tolerance of loud noises in the ipsilateral ear as a result of paralysis of stapedius. Its cause remains the subject of speculation but is most probably a herpes virus infection ( ). Magnetic resonance imaging studies suggest that there are inflammatory changes in the labyrinthine and perigeniculate segments of the facial nerve. In the vast majority of cases, spontaneous full recovery takes place after a few weeks.

Vestibulocochlear Nerve

The vestibulocochlear nerve is the eighth cranial nerve. As its name suggests, it consists of two components, the vestibular nerve, concerned with balance, and the cochlear nerve, concerned with hearing.

Vestibular nuclei and central connections

The vestibular nuclear complex is situated beneath the floor of the fourth ventricle. It has traditionally been divided into four cellular masses, the lateral nucleus (of Deiter), medial nucleus (of Schwalbe), superior nucleus (of Bechterew) and inferior (spinal, descending) vestibular nucleus (see Figure 28.8, Figure 28.11 ) ( ). The nucleus prepositus hypoglossi, located medial to the medial vestibular nucleus, may also be regarded as a vestibular nucleus. Cell groups X and Y have been identified as accessory nuclei of the vestibular complex.

The medial vestibular nucleus is the largest of the vestibular nuclei. It lies lateral and superior to the visceromotor nuclei of the glossopharyngeal and vagus nerves, is crossed dorsally by the striae medullares of the fourth ventricle and is continuous with the nucleus intercalatus below. The superior vestibular nucleus is small and lies above the medial and lateral nuclei. It extends higher into the pons than the other subdivisions and occupies the upper part of the vestibular area. The inferior vestibular nucleus is the smallest of the group. It lies between the medial vestibular nucleus and the restiform body, from the level of the upper end of the nucleus gracilis to the pontomedullary junction, and is traversed by descending fibres of the vestibular nerve, from the cerebellum and the vestibulospinal tract. The lateral vestibular nucleus lies ventrolateral to the upper portion of the medial nucleus, just above the inferior nucleus and ascends almost to the level of the abducens nucleus. Its rostral end is continuous with the caudal end of the superior nucleus. It projects to the spinal cord through the lateral vestibulospinal tract ( ). The lateral vestibular nucleus can be subdivided into dorsal and ventral components. The dorsal portion contains giant cells and is located within the juxtarestiform body: it receives no direct vestibular afferents, but receives Purkinje cell axons from the anterior vermis, and extends efferent connections to the cerebellum. The ventral portion contains cells of various sizes and is located medial to the juxtarestiform body

Vestibulocerebellar fibres (primary and secondary) travel via the juxtarestiform body mainly to the flocculus and nodule. Cerebellovestibular fibres pass to the nuclei, also via the juxtarestiform body: they arise mainly in the flocculus and nodule (posterior lobe), but some fibres are derived from the anterior lobe and the fastigial nucleus. A few vestibular fibres enter the cerebellum directly through the juxtarestiform body and end in the fastigial nucleus, flocculonodular lobe and uvula.

The vestibular nuclei receive afferent fibres from multiple sources, including the vestibular portion of the vestibulocochlear nerve, cerebellum, brainstem reticular formation and spinal cord. Myelinated fibres in the vestibular nerve enter the medulla between the restiform body and the spinal tract of the trigeminal nerve and bifurcate into ascending and descending branches. Most ascending vestibular fibres end in the superior vestibular nucleus. Descending vestibular fibres traverse the juxtarestiform body and give off extensive transverse collaterals to the inferior and medial (but not the lateral) vestibular nuclei. Additional afferent fibres descend from the cerebral cortex to innervate the vestibular nuclei bilaterally.

Afferent fibres arising from the interstitial nucleus of Cajal descend through the medial longitudinal fasciculus to synapse in the ipsilateral superior and medial vestibular nuclei and the nucleus prepositius hypoglossi. Afferent fibres from the Edinger–Westphal nucleus project to the medial and inferior vestibular nuclei, which also receive root fibres from the upper cervical nerve roots (C2, C3). The lateral vestibular nucleus and inferior portions of the medial and inferior vestibular nuclei receive ipsilateral spinovestibular fibres that ascend from lower spinal levels through the dorsal spinocerebellar tract.

The vestibular nuclei project extensively to the cerebellum and also receive fibres from the cerebellar cortex and the fastigial nuclei. Vestibulocerebellar fibres pass through the juxtarestiform body to the cerebellum ( ). A few vestibular nerve fibres bypass the vestibular nuclei and traverse the juxtarestiform body to terminate directly in the fastigial nucleus, the flocculonodular lobe and uvula.

Evidence suggests that the vestibular apparatus is spatially represented in these nuclei. Efferents from the lateral vestibular nucleus descend to the spinal cord through the ipsilateral lateral vestibulospinal tract and extend along the full length of the spinal cord to exert a facilitatory influence on spinal reflex activity and on extensor muscle tone. Vestibular nuclear efferents form a major part of the medial longitudinal fasciculus, through which interconnections between the vestibular and oculogyric nuclei and the pontine reticular formation enable conjugate eye movements that compensate for head movements perceived by the labyrinths. The reflex arcs stabilize visual fixation despite changing head position. Vestibular efferents project to the thalamus (probably via the posterior portions of the ventroposterior complex and the medial pulvinar), which in turn projects to the primary vestibular cortex in the parietal lobe at the junction of the intraparietal and postcentral sulci, adjacent to the head portion of the postcentral gyrus. There may be additional representation of the vestibular system in the superior temporal gyrus near to the auditory cortex.

Vestibular ganglion

The somata of the bipolar neurones that form the vestibular nerve are in the vestibular (Scarpa’s) ganglion located in the trunk of the nerve within the lateral end of the internal acoustic meatus (see Fig. 43.12B ). They vary considerably in size, with circumferences ranging from 45 to 160 μm ( ): no topographically ordered distribution relating to size has been found. The somata are notable for their abundant granular endoplasmic reticulum, which in places forms Nissl bodies, and prominent Golgi complexes. They are covered by a thin layer of satellite cells and are often arranged in pairs, closely abutting each other so that only a thin layer of endoneurium separates the adjacent coverings of satellite cells. This arrangement has led to speculation that ganglion cells may affect each other directly by electrotonic spread (ephaptic transmission: see ). Two distinct sympathetic components have been identified in the vestibular ganglion, a perivascular adrenergic system derived from the stellate ganglion, and a blood vessel-independent system derived from the superior cervical ganglion.

The central processes of the vestibular ganglion cells travel via the vestibular nerve, enter the brainstem at the cerebellopontine angle and terminate in the vestibular nuclear complex (see below). Neurones in this complex project to motor nuclei in the brainstem and upper spinal cord, and to the cerebellum and thalamus. Thalamic efferent projections pass to a cortical vestibular area that is probably located near the intraparietal sulcus in area 2 of the primary somatosensory cortex.

The peripheral processes of the vestibular ganglion cells are aggregated into definable nerves, each with a specific distribution. The main nerve divides at and within the ganglion into superior and inferior divisions connected by an isthmus. The superior division, the larger of the two, passes through the small holes in the superior vestibular area at the fundus of the internal acoustic meatus (see Fig. 43.3 ) and supplies the ampullary crests of the lateral and anterior semicircular canals via the lateral and anterior ampullary nerves, respectively. A secondary branch of the lateral ampullary nerve supplies the macula of the utricle; however, the greater part of the utricular macula is innervated by the utricular nerve, which is a separate branch of the superior division. Another branch of the superior division supplies part of the saccule. The inferior division passes through small holes in the inferior vestibular area (see Fig. 43.3 ) to supply the remainder of the saccule and the posterior ampullary crest via saccular and singular branches, respectively; the latter passes through the foramen singulare. Occasionally, a very small supplementary or accessory branch innervates the posterior crest; it is probably a vestigial remnant of the crista neglecta, an additional area of sensory epithelium found in some other mammals but seldom in humans.

Afferent and efferent cochlear fibres are also present in the inferior division of the vestibular nerve but leave at the anastomosis of Oort to join the main cochlear nerve (see review by ). Another anastomosis, the vestibulofacial anastomosis, is situated more centrally between the facial and vestibular nerves and is the point at which fibres originating in the intermediate nerve pass from the vestibular nerve to the main trunk of the facial nerve.

There are approximately 20,000 fibres in the vestibular nerve, of which 12,000 travel in the superior division and 8000 travel in the inferior division. The distribution of fibre diameters is bimodal, with peaks at 4 μm and 6.5 μm. The smaller fibres go mainly to the type II hair cells and the larger fibres tend to supply the type I hair cells. In addition to the afferents, efferent and autonomic fibres have been identified. Efferent fibres synapse exclusively with the afferent calyceal terminals around type I cells and usually with the afferent boutons on type II cells, although a few are in direct contact with the cell bodies of type II cells. The autonomic fibres do not contact vestibular sensory cells but terminate beneath the sensory epithelia.

Cochlear nuclei and central connections

The primary afferents of the auditory pathway are the central processes of type I and II sensory neurones in the spiral ganglion of Corti in the cochlea. They travel in the cochlear division of the vestibulocochlear nerve and enter the brainstem just caudal to the vestibular division of the vestibulocochlear nerve at the cerebellopontine angle. The axons bifurcate and terminate in the dorsal (posterior) and ventral (anterior) cochlear nuclei lying along the lateral surface of the restiform body. The dorsal cochlear nucleus forms a bulge, the auditory tubercle (auditory eminence) on the posterior surface of the restiform body and also forms the floor of the lateral recess of the fourth ventricle; it is continuous medially with the vestibular area in the floor of the fourth ventricle (see Fig. 28.9 ). The ventral cochlear nucleus lies between the cochlear and vestibular fibres of the vestibulocochlear nerve. The dorsal and ventral cochlear nuclei are separated only by a thin stratum of nerve cell bodies and fibres.

The axons of type 1 spiral ganglion cells enter the ventral cochlear nucleus and bifurcate within its centre, dividing it into anterior and posterior ventral subnuclei. The fibres are organized into dorsoventrally oriented sheets in which fibres conveying low frequency sounds from apical spiral ganglion cells terminate ventrally and fibres conveying high frequency sounds from the basal turn of the spiral ganglion terminate dorsally. Posterior processes of type I ganglion cells pass to the caudal pole of the ventral nucleus, from which they enter the dorsal cochlear nucleus. Thin unmyelinated axons of type 2 spiral ganglion cells contact clusters of small interneurones scattered through the cochlear nuclei.

The axons that emerge from the cochlear nuclei are aggregated into three, mostly transverse, fibre bundles, the dorsal and intermediate acoustic striae and the trapezoid body. The dorsal cochlear nucleus projects via the dorsal acoustic stria to the contralateral inferior colliculus. The dorsal acoustic stria curves dorsally and medially, dorsal to the restiform body, traverses the reticular formation ventral to the striae medullares, crosses the midline, passes to the contralateral superior olivary complex, and ascends in the contralateral lateral lemniscus. The ventral cochlear nucleus projects via the trapezoid body or the intermediate acoustic stria to relay centres in either the superior olivary complex, the nuclei of the lateral lemniscus, or the inferior colliculus. Initially, the intermediate acoustic stria accompanies the dorsal acoustic stria as they pass medially, immediately dorsal to the restiform body, but it then separates from the dorsal acoustic stria, angles sharply ventrally in the ipsilateral tegmentum, passes through the spinal trigeminal nucleus, joins the trapezoid body, and crosses the midline to join and ascend with the contralateral lateral lemniscus.

The cochlear nerve also conveys efferent fibres that pass outward from the brainstem to terminate on the hair cells and on the cells of the spiral ganglion.

Intratemporal and intracranial course

The vestibulocochlear nerve emerges from the cerebellopontine angle (see Figure 28.4, Figure 28.5 ). It courses through the posterior cranial fossa in close association with the facial nerve, nervus intermedius and labyrinthine vessels. Together with these structures, it enters the petrous temporal bone via the porus acusticus of the internal acoustic meatus (see Fig. 43.3 ), and divides into an anterior trunk, the cochlear nerve, and a posterior trunk, the vestibular nerve. Both contain the centrally directed axons of bipolar neurones, together with a smaller number of efferent fibres that arise from brainstem neurones and terminate on cochlear and vestibular sensory neurones. In humans, the intratemporal portion of the vestibulocochlear nerve consists of two histologically distinct portions: a central glial zone adjacent to the brainstem, and a peripheral or non-glial zone ( ). In the glial zone, the axons are supported by central neuroglia, whereas in the non-glial zone they are ensheathed by Schwann cells. The non-glial zone sometimes extends into the cerebellopontine angle medial to the internal acoustic meatus in human vestibulocochlear nerves. During development, a gap of several weeks has been reported between the onset of Schwann cell myelination distally and glial myelination proximally; it has been suggested that the gap may coincide with the time of the final maturation of the organ of Corti. For further details about the development of the human cochlear nerve, see ).

In audiological practice, it is important to distinguish between intratemporal and intracranial lesions. However, this surgical distinction does not correlate with the precise anatomical description of peripheral and central portions of the auditory and vestibular systems. Clinically, the term ‘peripheral auditory lesion’ is used to describe lesions peripheral to the spiral ganglion, and the term ‘peripheral vestibular disturbance’ includes lesions of the vestibular ganglion and the entire vestibular nerve.

Glossopharyngeal Nerve

The glossopharyngeal nerve is the ninth cranial nerve (see Figure 35.8, Figure 35.11, Figure 35.19 ). It contains motor fibres to stylopharyngeus; parasympathetic secretomotor fibres to the parotid gland (derived from the inferior salivatory nucleus); sensory fibres to the tympanic cavity, pharyngotympanic tube, fauces, tonsils, nasopharynx, uvula and posterior (postsulcal) third of the tongue; and gustatory fibres from the circumvallate papillae.

Nucleus ambiguus

The nucleus ambiguus is a complex group of large motor neurones situated deep in the medullary reticular formation (see Figure 28.3, Figure 28.9 ). Rostrally, it extends as far as the rostral end of the vagal nucleus and caudally it is in line with, but not continuous with, the nucleus of the accessory nerve. A small retrofacial nucleus separates the upper end of the nucleus ambiguus from the facial nucleus. The nucleus ambiguus contains cellular subgroups that exhibit some topographical representation. Relatively discrete groups of cells in the caudal region innervate individual laryngeal muscles; neurones in the intermediate area innervate the pharynx; neurones in the rostral area innervate the oesophagus and soft palate.

The nucleus ambiguus contributes special visceral efferent fibres to the glossopharyngeal nerve and may give general visceral efferent fibres to the vagus nerve. Fibres emerging from the nucleus ambiguus pass dorsomedially, then curve laterally. Rostral fibres join the glossopharyngeal nerve and caudal fibres join the vagus nerve and are distributed to the pharyngeal constrictors, intrinsic laryngeal muscles and striated muscles of the palate and upper oesophagus. A specific caudal pool of laryngeal motor neurones gives rise to the recurrent laryngeal nerve.

Extracranial course

The glossopharyngeal nerve leaves the skull through the anteromedial part of the jugular foramen, anterior to the vagus and accessory nerves, and in a separate dural sheath. In the foramen, it is lodged in a deep groove leading from the cochlear aqueductal depression, separated by the inferior petrosal sinus from the vagus and accessory nerves. The groove is bridged by fibrous tissue, which is sometimes calcified. After leaving the foramen, the nerve passes forwards between the internal jugular vein and internal carotid artery, and then descends anterior to the latter, deep to the styloid process and its attached muscles, to reach the posterior border of stylopharyngeus. It curves forwards on stylopharyngeus and either pierces the lower fibres of the superior pharyngeal constrictor or passes between it and the middle constrictor to be distributed to the palatine tonsil, the mucosae of the pharynx and postsulcal part of the tongue, the vallate papillae and oral mucous glands.

Two ganglia, superior and inferior, are situated on the glossopharyngeal nerve as it traverses the jugular foramen. The superior ganglion is in the upper part of the groove occupied by the nerve in the jugular foramen. It is small, has no branches and is usually regarded as a detached part of the inferior ganglion. The inferior ganglion is larger and lies in a notch in the lower border of the petrous part of the temporal bone. Its cells are typical unipolar neurones, their peripheral branches convey gustatory and tactile signals from the mucosa of the tongue (posterior third, including the sulcus terminalis and vallate papillae) and general sensation from the oropharynx, where the glossopharyngeal nerve is responsible for initiating the gag reflex.

The glossopharyngeal nerve has tympanic, carotid, pharyngeal, muscular, tonsillar and lingual branches.

Communicating branches

The glossopharyngeal nerve communicates with the sympathetic trunk and vagus and facial nerves. Its inferior ganglion is connected with the superior cervical sympathetic ganglion. Two filaments from the inferior ganglion pass to the vagus, one to its auricular branch and the other to its superior ganglion. A branch to the facial nerve arises from the glossopharyngeal nerve below the inferior ganglion and perforates the posterior belly of digastric to join the facial nerve near the stylomastoid foramen.

Tympanic nerve

The tympanic nerve leaves the inferior ganglion, ascends to the tympanic cavity through the tympanic canaliculus and divides into branches that contribute to the tympanic plexus on the promontory. The lesser petrosal nerve is derived from the tympanic plexus.

Carotid branch

The carotid branch is often double. It arises just below the jugular foramen and descends on the internal carotid artery to the wall of the carotid sinus and to the carotid body. The nerve contains primary afferent fibres from chemoreceptors in the carotid body and from the baroreceptors lying in the carotid sinus wall. It may communicate with the inferior ganglion of the vagus, or with one of its branches, and with a sympathetic branch from the superior cervical ganglion (see Fig. 35.20 ).

Pharyngeal branches

The pharyngeal branches are three or four filaments that unite with the pharyngeal branch of the vagus and the laryngopharyngeal branches of the sympathetic trunk to form the pharyngeal plexus near the middle pharyngeal constrictor. They constitute the route by which the glossopharyngeal nerve supplies sensory fibres to the mucosa of the pharynx.

Muscular branch

The muscular branch supplies stylopharyngeus.

Tonsillar, lingual and lesser petrosal branches and the tympanic plexus

The tympanic plexus supplies branches to the mucosa of the tympanic cavity, pharyngotympanic tube and mastoid air cells. The nerves that constitute the tympanic plexus ramify on the surface of the promontory on the medial wall of the tympanic cavity. They are derived from the tympanic branch of the glossopharyngeal nerve and the caroticotympanic nerves. The former arises from the inferior ganglion of the glossopharyngeal nerve and reaches the tympanic cavity via the tympanic canaliculus for the tympanic nerve. The superior and inferior caroticotympanic nerves are postganglionic sympathetic fibres derived from the carotid sympathetic plexus that traverse the wall of the carotid canal to join the plexus.

The lesser petrosal nerve, which may be regarded as the continuation of the tympanic branch of the glossopharyngeal nerve, traverses the tympanic plexus. It occupies a small canal below that for tensor tympani. It runs past, and receives a connecting branch from, the geniculate ganglion of the facial nerve. The lesser petrosal nerve emerges from the anterior surface of the temporal bone via a small opening lateral to the hiatus for the greater petrosal nerve and then traverses either the foramen ovale or the sphenopetrosal fissure or the innominate canal (of Arnold) to join the otic ganglion. Postganglionic secretomotor fibres leave this ganglion in the auriculotemporal nerve to supply the parotid gland. The tympanic plexus sends a branch to the greater petrosal nerve via an opening anterior to the fenestra vestibuli.

The tonsillar region is innervated by tonsillar branches of the maxillary and glossopharyngeal nerves. The fibres from the maxillary nerve pass through, but do not synapse in, the pterygopalatine ganglion; they are distributed through the lesser palatine nerves and form a plexus (the circulus tonsillaris) around the tonsil together with the tonsillar branches of the glossopharyngeal nerve. Nerve fibres from this plexus are also distributed to the soft palate and the region of the oropharyngeal isthmus. The tympanic branch of the glossopharyngeal nerve supplies the mucous membrane lining the tympanic cavity. Infection, malignancy and postoperative inflammation of the tonsil and tonsillar fossa may therefore be accompanied by pain referred to the ear.

Lesions of the glossopharyngeal nerve

There are anastomoses between the glossopharyngeal, vagus, greater auricular and other branches of the cervical plexus that may be significant when treating patients with skull base pathology. Damage to the glossopharyngeal nerve is rarely seen without associated involvement of other lower cranial nerves. Transient or sustained hypertension may follow surgical section of the nerve and indicates involvement of the carotid branch. Isolated lesions of the glossopharyngeal nerve lead to loss of sensation over the ipsilateral soft palate, fauces, pharynx and posterior third of the tongue, although this is difficult to assess clinically, and confirmation requires galvanic stimulation. The palatal and pharyngeal (gag) reflexes are reduced or absent and salivary secretion from the parotid gland may also be reduced. Weakness of stylopharyngeus cannot be tested individually. Glossopharyngeal neuralgia consists of episodic brief but severe pain, often precipitated by swallowing and experienced in the throat, behind the angle of the jaw and within the ear. Superior jugular bulb thromboses (e.g. in otitis media) and jugular foramen syndrome (associated with nasopharyngeal carcinoma or a jugular paraganglioma) may cause lesions of the adjacent glossopharyngeal, vagus and accessory nerves, with associated weakness in the muscles supplied in the pharynx and larynx.

Vagus nerve

The vagus is the tenth cranial nerve. It is a large mixed nerve, containing branchiomotor nerve fibres, general visceral afferents and preganglionic parasympathetic fibres. The vagus has a more extensive course and distribution than any other cranial nerve, running through the neck, thorax and abdomen. The proportion of efferent parasympathetic nerve fibres it contains varies at different levels but is small relative to its sensory and sensorimotor content. The branchiomotor fibres arise from neurones in the nucleus ambiguus in the medulla oblongata and are distributed to the constrictor muscles of the pharynx and the intrinsic muscles of the larynx. The preganglionic parasympathetic nerves arise from cell bodies in the vagal dorsal nucleus in the medulla oblongata, travel in the nerve and in its pulmonary, cardiac, oesophageal, gastric and intestinal branches and relay/synapse in minute ganglia in the visceral walls (see Fig. 56.6 ) ( ).

The general visceral afferent fibres carry sensation from thoracic and abdominal viscera and from aortic arch baroreceptors and chemoreceptors and end in the nucleus solitarius of the medulla. Their cell bodies are in the superior and inferior vagal ganglia. Their peripheral processes are distributed to terminals in the pharyngeal and oesophageal walls where, acting synergistically with glossopharyngeal visceral afferents in the pharynx, they are concerned with swallowing reflexes. Vagal afferents are also believed to innervate the thyroid and parathyroid glands. In the heart, vagal afferents innervate the walls of the great vessels, the aortic bodies and pressor receptors, where they are stimulated by raised intravascular pressure. In the lungs they are distributed via the pulmonary plexuses: they supply bronchial mucosa, where they are probably involved in cough reflexes; bronchial muscle, where they encircle myocytes and end in tendrils, which are sometimes regarded as muscle spindles and which are believed to be stimulated by change in the length of myocytes; interalveolar connective tissue, where their knob-like endings, together with terminals on myocytes, may evoke Hering–Breuer reflexes; the adventitia of pulmonary arteries, where they may be pressor receptors; and the intima of pulmonary veins, where they may be chemoreceptors. Vagal visceral afferent fibres also end in the gastric and intestinal walls, digestive glands and the kidneys. Fibres ending in the gut and its ducts respond to stretch or contraction. Gastric impulses may evoke sensations of hunger and nausea.

About 5% of the vagal afferents project directly to, and terminate in, the upper cervical spinal cord (C1–C2), where they are believed to contribute to referred sensations, as well as to propriospinal mechanisms of nociceptive modulation. Almost all the central processes of vagal and glossopharyngeal afferent fibres end in the nucleus solitarius of the medulla.

Dorsal motor nucleus of the vagus

The dorsal motor nucleus of the vagus lies slightly dorsolateral to the hypoglossal nucleus, from which it is separated by the nucleus intercalatus (see Fig. 28.8 ). The vagal nucleus extends caudally to the first cervical spinal segment and rostrally to the open portion of the medulla subjacent to the vagal trigone of the fourth ventricle. It is a general visceral efferent nucleus and is the largest parasympathetic nucleus in the brainstem. Most (80%) of its neurones give rise to the preganglionic parasympathetic fibres that pass into the vagus nerve. The remainder are interneurones or neurones that project centrally. The nucleus innervates the cardiac and smooth muscle of the thoracic viscera (heart, bronchi, lungs, oesophagus) and abdomen (stomach, liver, pancreas, spleen, small intestine and proximal part of the colon), and glandular epithelium. Sparse sensory afferents from the nodose ganglion project directly to the dorsal and lateral edges of the dorsal vagal nucleus, and possibly beyond it into the nucleus of the solitary tract ( ).

Neurones within the nucleus are heterogeneous and can be classified into nine subnuclei, grouped into rostral, intermediate and caudal divisions. Topographic maps of visceral representation in animals suggest that the heart and lungs are represented in the caudal and lateral parts of the nucleus; the stomach and pancreas in intermediate regions; and the remaining abdominal organs in the rostral and medial parts of the nucleus.

Extracranial course

The vagus nerve exits the skull through the jugular foramen, accompanied by the accessory nerve, with which it shares an arachnoid and a dural sheath. Both nerves lie anterior to a fibrous septum that separates them from the glossopharyngeal nerve. The vagus descends vertically in the neck in the carotid sheath, between the internal jugular vein and the internal carotid artery, to the upper border of the thyroid cartilage, and then passes between the vein and the common carotid artery to the root of the neck. Its relationships in this part of its course are therefore similar to those described for these structures (see Figs 35.7A , 35.8 , 35.17 ). Its further course differs on the two sides. The right vagus descends posterior to the internal jugular vein to cross the first part of the subclavian artery and enter the thorax. The left vagus enters the thorax between the left common carotid and subclavian arteries, and behind the left brachiocephalic vein.

After emerging from the jugular foramen, the vagus bears two marked enlargements: a small, round, superior (jugular) ganglion and a larger inferior (nodose) ganglion.

Superior (jugular) ganglion

The superior ganglion is greyish, spherical and approximately 4 mm in diameter. It is connected to the inferior glossopharyngeal ganglion and the sympathetic trunk, the latter by a filament from the superior cervical ganglion.

Inferior (nodose) ganglion

The inferior or nodose ganglion is larger than the superior ganglion, and is elongated and cylindrical in shape with a length of 25 mm and a maximum breadth of 5 mm. It is connected with the hypoglossal nerve, the loop between the first and second cervical spinal nerves, and with the superior cervical sympathetic ganglion. Most visceral afferent axons travelling in the vagus nerve are the peripheral processes of cell bodies located in the nodose ganglion.

Both vagal ganglia are exclusively sensory and contain somatic, special visceral and general visceral afferent neurones. The superior ganglion is mainly somatic and most of its neurones enter the auricular nerve, whilst neurones in the inferior ganglion are concerned with visceral sensation from the heart, larynx, lungs and the alimentary tract from the pharynx to the transverse colon. Some fibres transmit impulses from taste endings in the vallecula and epiglottis. Large afferent fibres are derived from muscle spindles in the laryngeal muscles. Vagal sensory neurones in the nodose ganglion may show some somatotopic organization. Both ganglia are traversed by parasympathetic and perhaps some sympathetic fibres, but there is no evidence that vagal parasympathetic components relay in the inferior ganglion. Preganglionic motor fibres from the dorsal vagal nucleus and the special visceral efferents from the nucleus ambiguus, which descend to the inferior vagal ganglion, commonly form a visible band that skirts the ganglion in some mammals. These larger fibres probably provide motor innervation to the larynx in the recurrent laryngeal nerve, together with some contribution to the superior laryngeal nerve supplying cricothyroid.

Vagus nerve in the neck

The branches of the vagus in the neck are the meningeal, auricular, pharyngeal, carotid body, superior and recurrent laryngeal nerves and cardiac branches.

Meningeal branches

Meningeal branches appear to start from the superior vagal ganglion and pass through the jugular foramen to be distributed to the dura mater in the posterior cranial fossa (see Fig. 25.3B ).

Auricular branch

The auricular branch (Arnold’s nerve) arises from the superior vagal ganglion and is joined by a branch from the inferior ganglion of the glossopharyngeal nerve. It passes behind the internal jugular vein and enters the mastoid canaliculus on the lateral wall of the jugular fossa. Traversing the temporal bone, it crosses the facial canal approximately 4 mm above the stylomastoid foramen and here supplies an ascending branch to the facial nerve. Fibres of the nervus intermedius may pass to the auricular branch at this point, which may explain the cutaneous vesiculation in the auricle that sometimes accompanies geniculate herpes. The auricular branch then traverses the tympanomastoid fissure and divides into two rami. One ramus joins the posterior auricular nerve and the other is distributed to the skin of part of the ear and to the external acoustic meatus.

Pharyngeal branch

The pharyngeal branch of the vagus is the main motor nerve of the pharynx and consists of axons arising from neuronal cell bodies in the nucleus ambiguus. It supplies all the muscles of the soft palate (excluding tensor veli palatini, which is supplied by the mandibular division of the trigeminal nerve via the nerve to medial pterygoid) and all the muscles of the pharynx (excluding stylopharyngeus, which is supplied by the glossopharyngeal nerve). It emerges from the upper part of the inferior vagal ganglion, passes between the external and internal carotid arteries to reach the upper border of the middle pharyngeal constrictor, and divides into numerous filaments that join rami of the sympathetic trunk and glossopharyngeal nerve to form the pharyngeal plexus. It also gives off a minute filament, the ramus lingualis vagi, which joins the hypoglossal nerve as it curves round the occipital artery.

Branches to the carotid body

Branches to the carotid body are variable in number. They may arise from the inferior ganglion or travel in the pharyngeal branch, or in the superior laryngeal nerve. They form a plexus with the glossopharyngeal rami and branches of the cervical sympathetic trunk.

Superior laryngeal nerve

The superior laryngeal nerve is larger than the pharyngeal branch and issues from the middle of the inferior vagal ganglion. It receives one or more communicating branches from the superior cervical sympathetic ganglion; most frequently, the connection is with the external laryngeal nerve. The sympathetic communication contributes to the innervation of the carotid body ( ) and the thyroid gland ( ). The superior laryngeal nerve descends alongside the pharynx, at first posterior, then medial, to the internal carotid artery: it may be found on the lateral side of the artery. It divides into a smaller, external, and a larger, internal, branch, approximately 1.5 cm below the ganglion: rarely, both branches may arise from the ganglion (see Figure 35.8, Figure 35.12 ).

Internal laryngeal nerve

The internal laryngeal nerve passes forwards approximately 7 mm before piercing the thyrohyoid membrane, usually at a higher level than the superior thyroid artery ( ). It splits into superior, middle and inferior branches on entering the larynx. The superior branch supplies the mucosa of the piriform fossa. The large middle branch is distributed to the mucosa of the ventricle, specifically the quadrangular membrane, and therefore probably conveys the afferent component of the cough reflex. The inferior ramus is mainly distributed to the mucosa of the ventricle and subglottic cavity. On the medial wall of the piriform fossa, descending branches give twigs to the interarytenoid muscle and share a number of communicating branches with the recurrent laryngeal nerve ( ).

The internal laryngeal nerve is sensory to the laryngeal mucosa down to the level of the vocal folds. It also carries afferent fibres from the laryngeal neuromuscular spindles and other stretch receptors. It descends to the thyrohyoid membrane, pierces it above the superior laryngeal artery and divides into an upper and lower branch. The upper branch is horizontal and supplies the mucosa of the pharynx, the epiglottis, vallecula and laryngeal vestibule. The lower branch descends in the medial wall of the piriform recess, supplies the aryepiglottic fold, the mucosa on the back of the arytenoid cartilage and one or two branches to transverse arytenoid (the latter unite with twigs from the recurrent laryngeal to supply the same muscle). The internal laryngeal nerve ends by piercing the inferior pharyngeal constrictor to unite with an ascending branch from the recurrent laryngeal nerve. As it ascends in the neck, it supplies branches, more numerous on the left, to the mucosa and tunica muscularis of the oesophagus and trachea, and to the inferior constrictor.

External laryngeal nerve

The external laryngeal nerve, smaller than the internal, descends behind sternohyoid with the superior thyroid artery, but on a deeper plane. It lies first on (occasionally within) the inferior pharyngeal constrictor, to which it contributes some small branches, and then pierces it to curve round the inferior thyroid tubercle. It passes beneath the attachment of sternothyroid to the oblique line of the thyroid cartilage to reach cricothyroid, which it supplies. A communicating nerve continues from the posterior surface of cricothyroid, crosses the piriform fossa and enters thyroarytenoid, where it anastomoses with branches from the recurrent laryngeal nerve. It has been suggested that these communicating branches may provide both additional motor components to thyroarytenoid and sensory fibres to the mucosa in the region of the subglottis. An anastomosis between the external and internal laryngeal nerves has also been described ( ). The external laryngeal nerve also gives branches to the pharyngeal plexus and communicates with the superior cardiac nerve and superior cervical sympathetic ganglion.

The close relationship of the external laryngeal nerve to the superior thyroid artery puts the nerve at potential risk when the artery is clamped during thyroid lobectomy, particularly when it is close to the artery (in approximately 20% of cases), or where, instead of crossing the superior thyroid vessels approximately 1 cm or more above the superior pole of the gland, it actually passes below this point (in some 20% of cases) ( , ). The external laryngeal nerve is also at risk in parathyroidectomy, carotid endarterectomy and anterior cervical spine procedures.

Recurrent laryngeal nerve

The recurrent laryngeal nerve supplies all the laryngeal muscles except cricothyroid, communicates with the internal laryngeal nerve and carries sensory filaments from the laryngeal mucosa below the vocal folds and from laryngeal stretch receptors. The nerve differs in origin and course on the two sides. On the right, it almost always arises from the vagus anterior to the first part of the subclavian artery, and curves backwards below and then behind it to ascend obliquely to the side of the trachea behind the common carotid artery. Near the lower pole of the lateral lobe of the thyroid gland, it is closely related to the inferior thyroid artery and crosses either in front of, behind or between its branches. On the left, the recurrent laryngeal nerve arises from the vagus on the left of the aortic arch, curves below it immediately behind the attachment of the ligamentum arteriosum to the concavity of the aortic arch and ascends to the side of the trachea (see Fig. 57.61 ). As the recurrent laryngeal nerve curves round either the subclavian artery or the aortic arch, it gives cardiac filaments to the deep cardiac plexus.

On both sides, the recurrent laryngeal nerve ascends in or near the tracheo-oesophageal groove. It is closely related to the medial surface of the thyroid gland before it passes under the lower border of the inferior constrictor, and it enters the larynx behind the articulation of the inferior thyroid cornu with the cricoid cartilage, passing either deep to (usually) or between (sometimes) the fibres of cricopharyngeus at its attachment to the lateral aspect of the cricoid cartilage. The nerve supplies cricopharyngeus as it passes. At this point, the nerve is in intimate proximity to the posteromedial aspect of the thyroid gland. The main trunk divides into two or more branches, usually below the lower border of the inferior constrictor, although branching may take place higher up. The anterior branch is mainly motor and is sometimes called the inferior laryngeal nerve (a term that is best avoided to prevent any confusion with a former synonym for the recurrent laryngeal nerve). It ascends posterior to the cricothyroid joint and its ligament, usually covered by fibres of posterior cricoarytenoid, then bends over the joint and continues forwards over lateral cricoarytenoid before terminating within thyroarytenoid. It innervates posterior cricoarytenoid by one or more branches, and interarytenoid, lateral cricoarytenoid and thyroarytenoid (Maranillo et al 2005). The posterior branch of the recurrent laryngeal nerve is mainly sensory and ascends deep to posterior cricoarytenoid to join the descending branch of the internal laryngeal nerve.

The recurrent laryngeal nerve does not always lie in a protected position in the tracheo-oesophageal groove but may be slightly anterior to it (more often on the right), and it may be markedly lateral to the trachea at the level of the lower part of the thyroid gland. The upper part of the nerve has a close but variable relationship to the inferior thyroid artery. On the right, it is as often anterior to, or posterior to, or intermingled with, the terminal branches of the artery; on the left, the nerve is usually posterior to the artery, though occasionally lies anterior to it. The stated incidences of these relationships vary between different authors. The recurrent laryngeal nerve has a highly variable branching pattern (see Fig. 41.13 ) ( ).

A very rare anomaly of relevance to laryngeal pathology and surgery is the ‘non-recurrent’ laryngeal nerve, where the right recurrent laryngeal nerve arises directly from the vagus nerve trunk high up in the neck and enters the larynx close to the inferior pole of the thyroid gland. Only the right side is affected, and it is always associated with an abnormal origin of the right subclavian artery from the aortic arch on the left side. If unrecognized, a non-recurrent laryngeal nerve may be susceptible to injury during surgery. It may also potentially be compressed by small tumours of the thyroid gland ( ).

Pharyngeal plexus

Almost all of the nerve supply to the pharynx, whether motor or sensory, is derived from the pharyngeal plexus, which is formed by the pharyngeal branches of the glossopharyngeal and vagus nerves with contributions from the superior cervical sympathetic ganglion. The plexus lies on the external surface of the pharynx, especially on the middle constrictor. Filaments from the plexus ascend or descend external to the superior and inferior constrictors before branching within the muscular layer and mucosa of the pharynx. Whether a ‘cranial root’ of the accessory nerve serves as the conduit for the major motor drive to the pharyngeal plexus, albeit via the vagus nerve, is a matter of debate. The plexiform connections that may be demonstrated between the accessory nerve and the vagus nerve within the posterior cranial fossa are too variable and insubstantial to support this function.

Laryngeal nerve anastomoses

Anastomotic communications between the internal, external and recurrent laryngeal nerves probably explain why normal or near normal voice function can be present following neck surgery when one or more branches of these nerves have been divided ( , , ). Numerous communications have been described ( , ). The ansa Galeni, lying on the interarytenoid muscles, forms a direct connection between the recurrent and internal laryngeal nerves: it may be a single or double trunk or a plexus. The cricoid communication connects branches that originate bilaterally from the recurrent laryngeal nerve with the superior branch of the deep portion of the arytenoid plexus. The thyroarytenoid communication links the ascending branch of the recurrent laryngeal nerve and the descending branch from the anterior branch of the internal laryngeal nerve. There are also communications between the external laryngeal and recurrent laryngeal nerves, the internal and external laryngeal nerves and the recurrent laryngeal and superior laryngeal nerves. The anastomosis between the recurrent laryngeal nerve and the sympathetic trunk probably explains why ptosis is a rare complication following thyroidectomy.

Lesions of the vagus nerve and recurrent laryngeal nerve paralysis

The consequences of vagal nerve lesions are complex, reflecting the long course of the nerve and the possible involvement of three of its branches, namely the pharyngeal, superior laryngeal and recurrent laryngeal nerves. Collectively these nerves innervate the muscles of the larynx, soft palate and pharynx, and injury may therefore have deleterious effects on phonation and/or soft palate movements and/or swallowing. A lesion of the vagus nerve above the level at which the pharyngeal branch is given off will affect both the superior and recurrent laryngeal nerves. This causes immobility of the vocal fold on the affected side and imparts a breathy voice with lack of pitch and limited loudness. The affected fold is paralysed and lies in the so-called ‘cadaveric’ position halfway between abduction and adduction. If the lesion is unilateral, the voice is weak and hoarse, but if it is bilateral, phonation is almost absent, the vocal pitch cannot be altered, and the cough is weak and ineffective. There will also be a degree of hypernasality because of the effects on movements of the soft palate caused by paralysis of levator veli palatini. Unilateral palsies impact very significantly on the quality of life of the patient because of impaired vocalization and a tendency to aspirate. Bilateral palsies are extremely serious and a tracheostomy may be required to protect the airway.

A lesion affecting the superior laryngeal nerve may be unilateral or bilateral. Complete section is most likely during the ligation of the vessels forming the vascular pedicle of the superior pole of the thyroid gland during thyroid lobectomy. Unilateral lesions may result in the vocal folds appearing relatively normal and the effect on voice is barely noticeable and is often overlooked. A more detailed examination may detect some shortening of the vocal folds on the affected side with asymmetric tilt of the epiglottis and the anterior larynx canted towards the unaffected side. The result is a mildly hoarse voice with loss of pitch control. Bilateral superior laryngeal nerve lesions result in shortening of both vocal folds, the overhanging of the epiglottis over the folds and reduced tilt between the thyroid and cricoid cartilage. Effects on the voice are correspondingly greater, with reduced loudness and pitch but variable breathiness.

Damage to the internal laryngeal nerve causes loss of mechanoreceptive and proprioceptive sensation from the larynx. Unilateral loss produces a feeling that something is stuck in the throat, whereas bilateral loss will result in aspiration and can cause dysphagia, with a risk of choking.

Unilateral complete palsy of the recurrent laryngeal nerve is more common on the left side, presumably because the nerve is longer on this side. There is isolated paralysis of all the laryngeal muscles on the affected side except cricothyroid, which is innervated by the external laryngeal nerve. The patient may be asymptomatic or have a hoarse, breathy voice and there will be diminished ability to manipulate pitch. The hoarseness may be permanent or may become less severe with time as the contralateral fold develops the ability to hyperadduct and oppose the paralysed fold, thereby closing the glottis during phonation and coughing, although this is seldom enough to restore full voice quality. There may be aspiration of food and drink. Bilateral lesions of the recurrent laryngeal nerve result in both vocal folds being paralysed and taking up a paramedian position. Phonation can be nearly normal because the vocal folds lie so close to the midline, but there will be audible stridor and a compromised airway.

Clinically the position of the vocal fold in the acute phase after section of the recurrent laryngeal nerve is very variable. Though stridor is more common after bilateral lesions and sometimes only audible to the educated ear, the folds may be sufficiently abducted so that there is little problem with airway obstruction at rest, although the voice is always weaker in this situation. The folds are slightly more widely separated in chronic lesions, which renders the voice weaker but with a less precarious airway. Atrophy and fibrosis of paraglottic muscles probably affect the position of paralysed vocal folds in chronic lesions to a greater degree than variations in the strength of the apposing adductor and abductor muscle groups.

For many years, conventional wisdom held that movements of abduction were affected to a greater degree than those of adduction when the recurrent laryngeal nerve was partially damaged (Semon’s law). This effect was thought to reflect the internal segregation within the recurrent laryngeal nerve of axons supplying the laryngeal abductor muscles. The hypothesis was later undermined by the demonstration that axons destined for particular laryngeal muscles are randomly distributed within the nerve. The weak abductive action of cricothyroid that is said to produce lengthening of the vocal folds has been suggested as the explanation for the relative sparing of abduction in these lesions, but is difficult to reconcile with the alternative view that cricothyroid may be a slight adductor of the vocal folds (Mu and Sanders 2009, ). It is likely that predicting the effect of partial lesions of the recurrent laryngeal nerve is complicated by the variable patterns of anastomoses between the laryngeal nerves. For further reading, see and .

Vagus nerve in the thorax

Preganglionic parasympathetic axons arise from neuronal cell bodies in the dorsal motor nucleus of the vagus in the medulla oblongata. Those destined for the thoracic viscera travel in the pulmonary, cardiac and oesophageal branches of the vagus nerve and synapse in minute ganglia in the walls of the target viscera. Axons travelling in cardiac branches join the cardiac plexuses and synapse in ganglia distributed freely over both atria, the interatrial septum and peri-nodal subepicardial tissue. Postganglionic fibres are distributed to the atria and the atrioventricular bundle and are concentrated around the sinuatrial and, to a lesser extent, the atrioventricular nodes. Cardiac branches slow the heart rate and diminish the force of contraction and may indirectly influence ventricular muscle via their effect on the atrioventricular node. The direct postganglionic parasympathetic innervation of the ventricles is sparse. The smaller branches of the coronary arteries are innervated mainly by parasympathetic vagal neurones, whereas the larger arteries are innervated mainly by sympathetic neurones. All cardiac branches of the vagus nerves contain both afferent and efferent fibres. Pulmonary branches contain axons that synapse in ganglia of the pulmonary plexuses: they are motor (bronchoconstrictor) to the circular non-striated muscle fibres of the bronchi and bronchioles (and therefore bring about bronchoconstriction), and secretomotor to the mucous glands of the respiratory epithelium.

Right vagus nerve

The right vagus nerve descends posterior to the right internal jugular vein in the neck and crosses the first part of the right subclavian artery to enter the thorax; the ansa subclavia is lateral, and the phrenic nerve is further lateral. After giving off the right recurrent laryngeal nerve, it descends through the superior mediastinum, at first posterior to the right brachiocephalic vein, and then lateral to the trachea and posteromedial to the superior vena cava. Superiorly, the right mediastinal part of the parietal pleura and superior lobe of the lung are lateral; inferiorly, the azygos vein and its arching terminal portion separate the right vagus nerve from the parietal pleura (see Fig. 56.7A ). The nerve next passes posterior to the right main bronchus and lies on the posterior aspect of the right lung hilum. Here, it gives off posterior bronchial branches that unite with rami from the second to fifth/sixth thoracic sympathetic ganglia to form the right posterior pulmonary plexus. Two or three branches descend from the inferior part of this plexus on the posterior aspect of the oesophagus to join a left vagal branch and form the posterior oesophageal plexus. The anterior vagal trunk, containing nerve fibres from both vagus nerves, leaves the plexus and passes inferiorly on the posterior surface of the oesophagus, entering the abdomen by passing through the oesophageal hiatus in the respiratory diaphragm.

Left vagus nerve

The left vagus nerve enters the thorax posterior to the left brachiocephalic vein, between the left common carotid artery (anterior) and left subclavian artery (posterior). It descends through the superior mediastinum, crosses the left side of the aortic arch, gives off the left recurrent laryngeal nerve and then passes posterior to the left lung hilum (see Fig. 56.7B ). Superior to the aortic arch, it is crossed anterolaterally by the left phrenic nerve, and on the arch, it is crossed laterally by the left superior intercostal vein and sometimes by a continuation of the accessory hemiazygos vein. Posterior to the hilum of the left lung, the left vagus nerve gives off posterior bronchial branches that unite with rami of the second to fourth thoracic sympathetic ganglia to form the left posterior pulmonary plexus. Two or three branches descend anteriorly on the oesophagus and join with a ramus from the right posterior pulmonary plexus to form the anterior oesophageal plexus. The posterior vagal trunk, containing nerve fibres from both vagus nerves, passes inferiorly on the anterior surface of the oesophagus and enters the abdomen through the oesophageal hiatus. Numerous branches from the anterior vagal and left sympathetic trunks are distributed to the descending thoracic aorta. This pathway transmits the typical intense, excruciating pain of an intramural aortic haematoma from aortic dissection or laceration.

Vagus nerve in the abdomen

The vagus nerves supply parasympathetic innervation to the abdominal viscera as far as the distal transverse colon, i.e. they supply the derivatives of the foregut and midgut, whereas the derivatives of the hindgut are supplied by parasympathetic fibres travelling via the pelvic splanchnic nerves; the overlap between these two supplies is variable. The vagal trunks are derived from the oesophageal plexus and enter the abdomen via the oesophageal hiatus, closely related to the anterior and posterior walls of the abdominal part of the oesophagus (see Fig. 56.6 ). The anterior vagal trunk is mostly derived from the left vagus and the posterior vagal trunk from the right vagus. The nerves supply the abdominal part of the oesophagus and stomach directly. The anterior vagal trunk gives off a hepatic branch, which innervates the liver and its vasculature, including the gallbladder and bile ducts, and the other structures in the free edge of the lesser omentum (hepatoduodenal ligament). The posterior vagal trunk supplies branches to the coeliac plexus: these fibres frequently constitute the largest portion of the fibres contributing to the plexus and arise directly from the posterior vagal trunk and from its gastric branch, and run beneath the peritoneum, deep to the posterior wall of the superior part of the omental bursa, to reach the coeliac plexus. Their synaptic relays with postganglionic neurones are situated in the myenteric (Auerbach’s) and submucosal (Meissner’s) plexuses in the wall of the hollow viscera.

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