DAVID COX: Now that we know a little bit about the anatomy of the cochlea,

let's discuss what happens when sound waves hit our ears.
Here's the overall picture.
When the sound waves hit the middle ear and cause the ossicles to move,
the footplate of the stapes pushes onto the oval window membrane.
This displaces the perilymph in the scala vestibuli and scala tympani,
causing the basilar membrane to vibrate.
When the basilar membrane moves in relation to the tectorial membrane,
there is a shearing force on the hair cells of the organ of Corti.
These shearing forces cause ion channels in the hair cells to open and close,
which depolarizes or hyperpolarizes the cells, respectively.
The hair cells, therefore, transduce a mechanical signal--
the force of shearing that they feel--
into an electrical signal that neurons can use,
either depolarization or hyperpolarization.
So sound waves vibrate bones, which vibrate
fluids, which fluctuate a membrane, which
physically moves cells, which opens ion channels, which causes depolarization.
Let's take a closer look at the hair cells of the organ of Corti,
because that's where the mechanical signal is actually
transduced into an electrical signal.
A hair cell is anchored in the basilar membrane
and surrounded by a non-sensory supporting cell.
At the top of a hair cell are 60 or so tiny hair-like structures
called stereocilia that, together, form a hair bundle.
The stereocilia are attached to the bottom of the tectorial membrane.
When a sound wave causes the basilar membrane
to vibrate relative to the tectorial membrane, the stereocilia tilt
either toward or away from the tall edge of the hair bundle.
When the stereocilia are tilted toward their tall edge
from the shearing force, a set of non-selective cation channels open.
Tilting them away from their tall edge closes these channels.
There are numerous experiments that suggest that these ion channels are
mechanically gated, meaning a simple push or pull will open or close them,
as opposed to being voltage- or ligand-gated, like the other ion
channels we have encountered so far in this course.
When the shearing force tilts stereocilia towards the tall edge,
their mechanically-gated ion channels open,
and we see a depolarization of the hair cells.
Normally, about 10% of these channels are open at any given moment.
When stereocilia are sheared towards their tall edge, more of these channels
open, cations rush in, and the cell is depolarized.
When stereocilia are sheared away from their tall edge,
the 10% that were open close, and the cells hyperpolarize.
So a particular hair cell will depolarize or hyperpolarize
based on how and in which direction the basilar membrane is
displaced at any given moment.
These changes in electrical potential in the hair cells
cause the hair cells, which themselves do not fire action potentials,
to release neurotransmitter, which causes
depolarization and spiking of cells in a structure called the spiral ganglion.
Spikes are carried to the brain along the auditory nerve, which
is a bundle of axons that goes to the brainstem, where signals are relayed
through several weigh stations before eventually finding
their way to auditory thalamus and then cortex.