The first-order axons entering the brainstem from the cochlea terminate ipsilaterally on the dorsal and ventral cochlear nuclei. From here, second-order axons have several synaptic targets. Second-order axons forming the ascending auditory pathway ascend to the pons and project bilaterally on the superior olivary nucleus (eFig. 9.82). These bilateral projections are important for auditory acuity and localizing the origin of a sound. From here, fibers continue to ascend as the lateral lemniscus and terminate in the inferior colliculus of the midbrain. Axons leaving the inferior colliculus then have a synaptic relay in the medial geniculate nucleus of the thalamus before reaching the final synaptic target: the primary auditory cortex (superior temporal gyrus) of the temporal lobe (eFig. 9.82). Throughout this pathway, from the basilar membrane to the primary auditory cortex, auditory stimuli are tonotopically represented. This is analogous to the somatotopic map previously discussed in the section on somatosensory systems pathways.
In addition to the ascending auditory pathway, the superior olivary nucleus receives descending input from the primary auditory cortex as a form of feedback (eFig. 9.82). The superior olivary nucleus then sends descending olivocochlear fibers to the organ of Corti, which has an inhibitory function on the hair cells to prevent damage from harmfully loud sounds. It is also believed that the superior olivary nucleus has connections with the motor nuclei of the trigeminal and facial nerve to mediate reflexive contraction of the tensor tympani and stapedius muscles in response to loud sounds.
The vestibular nerve transmits afferent or sensory information regarding movement and position of the head from the vestibular organs, which include semicircular ducts, utricle, and saccule. Like the cochlear apparatus, each of these vestibular organs is located within a membranous (ductal) portion of the vestibular apparatus, which is surrounded by a bony (canal) portion (eFig. 9.83). The cell bodies for these sensory receptors are located in the vestibular (Scarpa’s) ganglion within the internal acoustic meatus (eFig. 9.83). The vestibular nerve then passes through the internal acoustic meatus and subarachnoid space to enter the pontomedullary junction at the cerebellopontine angle.
The central processes of the vestibular axons predominantly terminate in the four vestibular nuclei (superior, inferior, medial, and lateral), which are located in the rostral medulla and caudal pons (eFig. 9.40). Axons leaving the vestibular nuclei have several ascending and descending synaptic targets. These pathways connect with visual motor, descending motor, and cerebellar pathways to coordinate movement and maintain posture and balance (eFig. 9.84).
Axons ascending after leaving the vestibular nuclei form the medial longitudinal fasciculus to reach the oculomotor, trochlear, and abducent nuclei (eFig. 9.85). These connections coordinate movement of the head and eyes so that visual fixation on an object can be maintained. Other axons ascending from the vestibular nuclei project to the cerebral cortex after a synaptic relay in the ventral posterior nucleus of the thalamus (eFig. 9.84B). This connection assists in consciously orienting oneself in space. In addition to these connections, axons leaving the vestibular nuclei pass through the inferior cerebellar peduncle to reach the cerebellum and modulate equilibrium (eFig. 9.84B).
Axons descending after leaving the vestibular nuclei form the medial and lateral vestibulospinal tracts ipsilaterally (eFig. 9.84B). As mentioned in the previous section on medial motor systems in the spinal cord, the axons in these pathways synapse on interneurons within the anterior horn gray matter to influence spinal motor neuron activity and maintain posture and balance.
Part X: Hypothalamus