DR.

DAVID COX: After the mechanical energy of sound

has been transduced into neuronal activity in the inner ear,
it begins its journey to the brain.
Signals from the hair cells in each ear's cochlea
are sent through the spiral ganglion cells and auditory
nerves to a sequence of nuclei in the brainstem
before eventually reaching the midbrain, thalamus, and the auditory cortex.
A complex of nuclei in the brainstem known as the cochlear nucleus
complex, cochlear nucleus, or simply CN, is
located in the dorsolateral side of the brainstem,
spanning the junction of the pons and the medulla.
The CN consists of two heterogeneous collections of neurons
whose inputs come from the cochlear nerve
and whose outputs go towards higher regions of the auditory brainstem.
Auditory nerves enervate the cochlear nucleus
in a spatial pattern that matches their spatial organization in the basilar
membrane of the cochlea, forming a highly organized network based
on the frequencies to which each nerve is tuned.
Auditory nerves carrying signals from hair cells near the apical basilar
membrane have low characteristic frequencies.
Those from hair cells near the basal basilar
membrane near the round and oval windows have a high characteristic frequency,
and so on.
When these auditory axons enervate the cochlear nucleus,
they do so in a highly organized pattern based
on their characteristic frequencies.
This systematic relationship between the position in the cochlear nucleus
and characteristic frequency is another example
of tonotopy, a topographic organization of neural responses
according to frequency.
Tonotopic maps are found throughout the auditory system,
beginning at the basilar membrane and carried forward
in each subsequent processing stage, just the same way
that many structures in the visual system carry retinotopic organization
all the way to cortex.
After the cochlear nucleus, auditory signals
are carried deeper into the brain via two distinct pathways
before ultimately reaching the auditory cortex.
One set of signals is sent from the cochlear nucleus to the superior olive.
The superior olive marks the first place where
auditory signals from both ears come together, are paired, and integrated.
Two specialized groups of cells in distinct regions
of the superior olive encode differences in sound intensity or loudness
as well as timing differences between the sound perceived
by the left ear versus the right ear.
From the superior olive, auditory information
is sent to the inferior colliculus, which
is the principal nucleus of the auditory pathway in the midbrain.
Anatomically, it sits below the superior colliculus,
which is also known as the optic tectum, which we learned about
in the last lesson.
The superior colliculus plays an important role
in directing and orienting head and eye movements
in response to visual stimuli.
Since these two structures are so close together,
it's perhaps not surprising that the inferior colliculus
serves in an analogous coordinating function,
allowing us to orient ourselves in response to auditory stimuli.
The second set of signals leaving the cochlear nucleus
bypasses the superior olive and goes straight to the inferior colliculus.
Once these two sets of signals are processed in the inferior colliculus,
they are sent on to the medial geniculate nucleus.
The medial geniculate nucleus, also known as the MGN,
is the auditory subdivision of the thalamus,
and it basically is to the auditory system
what the lateral geniculate nucleus, or LGN, is to the visual system.
It's made up of a number of distinct cell types, forming distinct layers
and carrying out specific functions to help us hear.
It's also thought that the MGN plays a role
in allowing us to direct and maintain our attention to a particular sound.