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Sensory cranial nerves: want to learn more about it?

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Sensory cranial nerves

Before we continue our discussion of the cranial nerves, perhaps we should address the elephant in the room. It’s quite large and loud, and awfully difficult to miss… Can’t you see it? Smell it? How about hear it? How are you able to do this?

Actually, forget the elephant; the best way to answer these questions may be just to continue our exploration of the cranial nerves. Sight, smell and sound signals, as well as vestibular signals informing our sense of balance, are all carried by sensory cranial nerves. Although exceptionally complex, the following article attempts to delve a little deeper into the sensory cranial nerves responsible for transmitting signals to our brain for interpretation, informing us of the world that surrounds us (including, of course, the elephant).

Cranial nerves

List

The cranial nerves (CN) are twelve pairs of nerves that, with the exception of the spinal accessory nerve (CN XI), originate in the brain and contribute to the peripheral nervous system (PNS), supplying the head and neck.

Optic nerve (lateral-left view)

These 12 paired nerves, and their main branches, include:

  • The olfactory nerve (CN I)
  • The optic nerve (CN II)
  • The oculomotor nerve (CN III)
  • The trochlear nerve (CN IV)
  • The trigeminal nerve (CN V)
  • The abducens nerve (CN VI)
  • The facial nerve (CN VII)
    • Temporal branch
    • Zygomatic branch
    • Buccal branch
    • Mandibular branch
    • Cervical branch
  • The vestibulocochlear nerve (CN VIII)
  • The glossopharyngeal nerve (CN IX)
  • The vagus nerve (CN X)
  • The spinal accessory nerve (CN XI)
  • The hypoglossal nerve (CN XII)

Types

Of these nerves, some have special sensory functions, some have somatic sensory functions, some have autonomic functions, some have somatic motor functions, and some have a combination of the aforementioned functions. The olfactory nerve, the optic nerve, the facial nerve, the vestibulocochlear nerve, the glossopharyngeal nerve, and the vagus nerve each play roles in special sensory functions (i.e. olfaction, vision, gustation, audition, and balance).

The trigeminal nerve (all three branches: the ophthalmic, maxillary, and mandibular) and the glossopharyngeal nerve play roles in somatic sensory functions.

The oculomotor nerve, the facial nerve, the glossopharyngeal nerve, and the vagus nerve have important autonomic functions.

Finally, the oculomotor nerve, the trochlear nerve, the mandibular branch of the trigeminal nerve (V3), the abducens nerve, the facial nerve, the glossopharyngeal nerve, the vagus nerve, the spinal accessory nerve, and the hypoglossal nerve are responsible for motor functions.

One way to simplify efforts to understand each nerve and each of its separate functions is to separate the 12 cranial nerves into smaller groups. This can be done by first dividing the overarching group of 12 nerves into nerves which are considered to have primarily motor functions; nerves which are considered to have primarily sensory functions; and nerves which have a combination of both motor and sensory components.

To remember the names of the cranial nerves and whether they are primarily motor, sensory (or both) in nature, check out this cranial nerve mnemonics video:

This article will provide an introduction to the cranial nerves which are considered primarily sensory nerves, which includes the olfactory nerve, the optic nerve, and the vestibulocochlear nerve. The terminal nerve (CN 0, or CN N), although not conventionally included in the list of cranial nerves, will also be discussed among this group.

Olfactory nerve

Even at very low concentrations, humans can detect odors chemically. These chemicals stimulate chemoreceptors attached to cell bodies in the olfactory mucosa, a region of specialized epithelium in the nasal cavity located just below a sheet of bone in the skull called the cribriform plate. These bipolar sensory receptor neurons each give off single dendrites at one end that terminate as olfactory knobs in the olfactory mucosa, and single unmyelinated special visceral afferent axonal fibers which transmit olfactory information to the brain for interpretation at the other end.

Olfactory nerve (medial view)

Rather than coalescing into a single structure, it is small bundles of these individual axons which make up the olfactory nerve (CN I). These olfactory nerve bundles enter the brain via the cribriform plate of the ethmoid bone and transmit information to neurons in the olfactory bulb on the ipsilateral (same) side.

Olfactory bulb (medial view)

The olfactory bulb lies on the inferior (ventral) surface of the prefrontal cortex of the forebrain bilaterally. It receives signals from the olfactory nerve and is the first part of the central nervous system in which these signals are processed. The axons of the olfactory sensory neurons spread themselves over the surface of the bulbs, forming an olfactory nerve layer. The glomerular layer, located near the surface of the bulb, contains clusters of nerve terminals of the olfactory sensory neurons as well as dendrites of various other cell types. Mitral and tufted cells in particular are excited by the olfactory sensory neurons, and transmit the signals they receive from the olfactory bulbs to forebrain structures for continued processing. They do this via axonal bundles referred to as the olfactory tracts. There are two olfactory tracts or branches on each side: the medial and lateral olfactory striae.

Olfactory tract (medial view)

The lateral pathway sends olfactory information to the temporal lobe and nearby limbic structures: these axons project to the primary olfactory cortex (piriform cortex), amygdala, and entorhinal cortex located in the temporal lobe. Neurons in the piriform cortex, amygdala, and entorhinal cortex project to the prefrontal cortex. Neurons in the entorhinal cortex, via the perforant fiber pathway, also project to and modulate the functions of the hippocampus, a structure located in the medial temporal lobe associated with memory formation. Although some olfactory projections reach the prefrontal cortex without passing through and synapsing in the thalamus, the piriform cortex does send some axons to the the mediodorsal thalamic nucleus which in turn projects to a number of regions in the frontal lobe, including the prefrontal cortex.

Olfactory nerve (caudal view)

The medial pathway contains axonal fibers of some mitral and tufted cells which project to a number of medial limbic structures, including the substantia innominata, medial septal nucleus, and bed nucleus of the stria terminalis. Some mitral and tufted cell axons also project to the anterior olfactory nucleus, a region of cell bodies within the posterior part of each olfactory bulb. Neurons in the anterior olfactory nucleus subsequently gives off axons which also travel via the medial olfactory pathway; but instead of remaining on the ipsilateral side, these fibers cross to the opposite cerebral cortex via the anterior commissure (a fiber bundle rostral to the descending column of the fornix) and synapse on the contralateral olfactory bulb.

Olfactory projections to the limbic system mediate affective (emotional) aspects of olfactory sensation; whereas descending projections to the hypothalamus, midbrain periaqueductal gray, and autonomic centers of the brainstem and spinal cord mediate autonomic responses to olfactory stimuli.

Optic nerve

Pathway and parts

The optic nerve (CN II) is the cranial nerve that carries visual information, allowing us to see the world around us. It is made up of special sensory afferent fibers which transmit visual information from the retina, the innermost layer of the eye, to the brain. The signals transmitted by this nerve are first received by specialized receptors in the retina called photoreceptors, of which there are two types: rods, which respond to light, and cones, which detect color.

The retina represents a map of the visual field in which everything is reversed and upside down due to the angles at which light enters the eyes. As a result, the left visual field falls on the right side of the retina, the right visual field falls on the left side of the retina, the upper visual field falls on the lower side of the retina, and the lower visual field falls on the upper side of the retina. As such, the inferior, inner (nasal) quadrant of the retina corresponds to the upper, outer quadrant of the visual field, etc.

Optic nerve (cranial view)

The axonal processes of rods and cones synapse in the retina with other cells known as horizontal cells and bipolar cells, which process signals from the photoreceptors. Ganglion cells are the only retinal cells able to fire action potentials, and it is their axons which travel toward the posterior pole of the eye and coalesce, preparing to exit the eye at the optic disc.

Optic part of retina (axial view)

The optic disc is a pale circular region in the retina, colloquially known as the blind spot because it has no photoreceptors and thus is incapable of picking up visual stimuli. Despite this, we do not have an obvious hole in our visual fields because our brain essentially fills in the blank (although the blind spot can be demonstrated by looking straight forward with one eye closed and holding up one’s pointed index finger on the hand on the opposite side as the closed eye approximately a foot from one’s open eye. Then, moving the index finger from the midline slowly into the periphery; when the top of the finger disappears, you have found your blind spot).

Optic nerve (cranial view)

At the optic disc, the ganglionic axons become myelinated and exit each eye as the optic nerves. The optic nerves are additionally covered by extensions of the meninges which cover the brain. The optic nerves enter the cranial cavity via the optic canals, and enter the brain at the pre-optic region of the diencephalon. Upon entering the brain, the optic nerves join to form an “x”-shaped structure called the optic chiasm. It is here that the fibers extending from the inner (nasal) half of each retina cross to the contralateral side, whereas fibers extending from the outer (temporal) half of each retina remain ipsilateral and uncrossed. This ultimately means that each cerebral hemisphere receives information on the opposite visual field, and does so by receiving some information from both eyes.

Optic chiasm (cranial view)

The fibers then depart the optic chiasm as optic tracts, with the left optic tract containing axons from the temporal hemiretina of the left eye and nasal hemiretina of the right eye, and the right optic tract containing axons from the temporal hemiretina of the right eye and nasal hemiretina of the left eye.

Optic tract (cranial view)

These tracts project to the lateral geniculate nuclei (LGN) of the thalamus on their corresponding sides, where they synapse on layers of cell types finely attuned to detect different types of visual stimuli. The ventral magnocellular layers contains large cells focused on movement and location, while the dorsal parvocellular layers contains smaller cells focused on detail, color, etc. Axons of cells in the magnocellular layers project to more dorsal (superficial) regions of the primary visual cortex (the “where” pathway), whereas axons of cells in the parvocellular layers project to more ventral (deep) regions of the primary visual cortex (the “what” pathway).

Optic radiation (cranial view)

The primary visual cortex (Brodmann’s area 17) is the cortical region ultimately dedicated to the perception of the visual stimuli transmitted by the optic nerve.

Optic Nerve: Pupillary light reflex

The pupillary light reflex is the reflexive constriction of the pupils in response to light. There are two main components of this reflex: an afferent (sensory) component involving the optic nerve and an efferent component involving the oculomotor nerve. The reflex is triggered when cells in the retina are activated by light, sending signals down the optic nerve (the afferent component of the reflex). Optic nerve fibers project and synapse in the pretectal region of the midbrain, which contributes to the circuit for the pupillary light reflex.

Neurons from the pretectal region then project and synapse in the oculomotor nucleus. From here, preganglionic parasympathetic neurons from the Edinger-Westphal nucleus send axons via the oculomotor nerve to synapse with postganglionic neurons in the ciliary ganglion, which sends signals to the sphincter pupillae muscle to contract and leads to constriction of the pupil (the efferent component of the reflex).

Ciliary ganglion (lateral-left view)

Vestibulocochlear nerve

The vestibulocochlear nerve (CN VIII) transmits information regarding two very important senses: audition and balance. Auditory information is transmitted by the cochlear part of the vestibulocochlear nerve, and information on balance is transmitted by the vestibular part of the vestibulocochlear nerve.

Auditory Signals and the cochlear component

Our ability to perceive sound begins with specialized receptors called hair cells, located in a structure called the Organ of Corti in the inner ear. These cells are triggered when air pressure waves incite vibrations in the tympanic membrane, which subsequently oscillate the stapes bone against the oval window of a structure called the cochlea.

Cochlea (anterior view)

The cochlea is filled with fluid called perilymph that is non-compressible, so movement of the stapes against the oval window by air pressure waves  transmits those pressure waves into the cochlea, leading to oscillatory movement of the perilymph. This leads to vibration of a membrane within the cochlea called the basilar membrane, which contains the Organ of Corti.

Cochlear nerve (anterior view)

There are two types of hair cells in the Organ of Corti: outer hair cells, and inner hair cells. The outer hair cells are called stereocilia and lie between two membranes: their tips are situated within a membrane called the tectorial membrane, and their bodies lie on the basilar membrane. When the basilar membrane is displaced upward by pressure waves, the stereocilia are displaced.

Stereocilia (histological slide)

Lateral displacement depolarizes the cell, triggering the generation of a signal; whereas medial displacement hyperpolarizes the cell, preventing the generation of a signal. The frequency triggering signal generation by a hair cell depends on the hair cell’s location in the basilar membrane. This is referred to as a tonotopic distribution: hair cells located in the basal portion respond to high frequencies, whereas those located in the apical portion respond to low frequencies.

Cochlear nerve (anterior view)

Once signals are generated by stereocilia in the Organ of Corti, they are transmitted to the peripheral processes of bipolar neurons that form a structure called the spiral ganglion. Projections from these bipolar neurons form the cochlear nerve.

Balance/movement and the vestibular component

The anatomical components of the vestibular system are the otolith organs (the saccule and utricle) which detect linear acceleration, and three semicircular canals, which detect angular acceleration.

Vestibular nerve (anterior view)

Like the cochlea of the auditory system, these organs are filled with endolymph; and like the receptors of the auditory system, the receptors of the vestibular system are hair cells. These hair cells also come in two types: type I hair cells, which are goblet shaped; and type II hair cells, which are cylindrical; but in the vestibular system, stereocilia project from the apical part of all types of hair cells, with small kinocilium extending slightly above each stereocilium. The bodies of these hair cells are surrounded by supporting cells connected by tight junctions, separating the hair cell from the endolymph surrounding the cilia while the apical stereocilia project into a gelatinous matrix.

Kinocilia (histological slide)

Displacement of the cilia occurs when the endolymph moves in a particular direction, i.e. when there is a change in position or motion of the head. Displacement of the stereocilia toward the kinocilia results in excitation of the hair cells; whereas displacement of the stereocilia away from the kinocilia results in inhibition of the hair cells.

Vestibular nerve (anterior view)

The hair cells are innervated by peripheral processes of vestibular (Scarpa’s) ganglia within the vestibular labyrinth (the maculae of the utricle and saccule, and the ampullae of the semicircular canals). The central processes of these bipolar neurons in the vestibular ganglia make up the vestibular part of the vestibulocochlear nerve.

Pathway of the vestibulocochlear nerve

The vestibular and cochlear components of the vestibulocochlear nerve travel as separate nerves through the internal auditory meatus, along with the facial nerve and labyrinthine artery.

Vestibulocochlear nerve (anterior view)

They then cross the posterior cranial fossa. It is in the posterior cranial fossa, within the petrous part of the temporal bone, that the cochlear nerve and the vestibular nerve join to form a unified vestibulocochlear nerve.

Vestibulocochlear nerve (posterior view)

This unified nerve reaches the lateral surface of the brainstem at the medullopontine angle. At this point, the vestibulocochlear nerve re-divides into vestibular and cochlear branches where they  enter the brainstem at the rostral medulla.

Vestibulocochlear nerve (inferior view)

Cochlear branches

The axons making up the cochlear nerve project to cochlear nuclei in the rostral medulla in a tonotopic manner similar to that observed in the basilar membrane. Axons carrying high-frequency sound signals from the basal part of the basilar membrane project deep into the nuclei, while axons carrying low-frequency sound signals from the apical part of the basilar membrane project into more superficial regions of the nuclei.

Anterior cochlear nucleus (anterior view)

From here, signals are transmitted through various pathways to a number of brain areas, including:

  • the superior olivary nucleus
  • the inferior colliculi in the midbrain
  • the medial geniculate nucleus of the thalamus
  • the primary auditory cortex (in the transverse temporal gyri of Heschl in the medial superior temporal gyrus)
Posterior cochlear nucleus (anterior view)

Vestibular branches

The axons extending from afferent neurons in the vestibular ganglia in the labyrinth project to various vestibular nuclei in the rostral medulla and caudal pons in the brainstem. The nuclei to which they project depends on the vestibular structures from whence they originate.

Medial vestibular nucleus (anterior view)

Axons of afferent neurons from the ampullae of the semicircular canals project to the superior vestibular nucleus (SVN) and rostral portion of the medial vestibular nucleus (MVN).

Superior vestibular nucleus (anterior view)

Many axons of afferent neurons from the maculae of the utricle and saccule project to the lateral vestibular nucleus (LVN), but some axons from the macula of the saccule project to the inferior vestibular nucleus (IVN)  as well. Axons originating in these nuclei provide input to various regions of the brain, including:

  • the trochlear nuclei
  • oculomotor nuclei
  • abducens nuclei
  • cerebellum
  • parts of the cerebral cortex
Lateral vestibular nucleus (anterior view)

Vestibulocochlear nerve and the vestibulo-oculomotor reflex

Signals received and transmitted by the vestibular branches of the vestibulocochlear nerve influence eye movements by way of the vestibular nuclei, which in turn project axons to the oculomotor, trochlear, and abducens nuclei which influence movement of the extraocular muscles. This works such that when the head moves in one direction, the eyes move in the opposite direction, staying fixed on an object until the object leaves the visual field. The eyes then move rapidly with the movement until fixing on another object until that new object is out of vision. This rapid eye movement between fixations is called a saccade.

This sequence of slower eye movement in the opposite direction to head movement with fixation followed by rapid eye movement in the same direction as head movement until fixation is again achieved is called nystagmus or the vestibulo-ocular reflex. The slower eye movement in the direction opposite to head movement is controlled by vestibular input from the vestibular nuclei, and involves the medial longitudinal fasciculus (MLF). The rapid saccadic eye movement is controlled by the paramedian pontine reticular formation (PPRF).

Medial longitudinal fasciculus (axial view)

Terminal nerve

Although not typically taught in most medical programs, some consider the terminal nerve (CN 0, or CN N) as a thirteenth cranial nerve. Its sensory endings are located in the nasal mucosa. It has three main filaments in the nasal cavity, located just posterior to the antero-superior border of the nasal septum, and its branches have a similar distribution as the olfactory nerves. Also like the branches of the olfactory nerve, branches of the terminal nerve pass through the foramina of the cribriform plate as well. The nerve fibers course posteriorly along the medial surface of the olfactory tract to the surface of the brain near the olfactory trigone, between the olfactory peduncles and rostral to the optic chiasm.

There are a number of theories regarding the function of the terminal nerve, but it is believed that it plays a role in reproductive behavior. It has been speculated that it is used to detect pheromones, contributing to mate selection, and that it enhances olfaction to influence reproductive behavior (i.e. based on findings such as a woman’s sense of smell is most acute when she is ovulating, women rating smell as a man’s most attractive feature, and traumatic loss of olfactory nerves is often accompanied by a reduction in libido).

Sensory cranial nerves: want to learn more about it?

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