The pressure waves in the endolymph
cause the basilar membrane to vibrate, which moves the hair cells
of the spiral organ against the tectorial membrane. This leads to
bending of the stereocilia and ultimately to the generation of
nerve impulses in first-order neurons in cochlear nerve fibers.
- The hair cells transduce mechanical
vibrations into electrical signals. As the basilar membrane
vibrates, the hair bundles at the apex of the hair cell bend back
and forth and slide against one another.
- A tip link protein connects the tip
of each stereocilium to a mechanically gated ion channel called the
transduction channel in its taller stereocilium neighbor.
- As the stereocilia bend in the
direction of the taller stereocilia, the tip links tug on the
transduction channels and open them.
- These channels allow cations in the
endolymph, primarily K+, to enter the hair cell
cytosol.
- As cations enter, they produce a
depolarizing receptor potential.
- Depolarization quickly spreads
along the plasma membrane and opens voltage-gated Ca2+
channels in the base of the hair cell.
- The resulting inflow of
Ca2+ triggers exocytosis of synaptic vesicles containing
a neurotransmitter, which is probably glutamate.
- As more neurotransmitter is
released, the frequency of nerve impulses in the first-order
sensory neurons that synapse with the base of the hair cell
increases.
- Bending of the stereocilia in the
opposite direction closes the transduction channels, allows
hyperpolarization to occur, and reduces neurotransmitter release
from the hair cells.
- This decreases the frequency of
nerve impulses in the sensory neurons.
Sound waves of various frequencies
cause certain regions of the basilar membrane to vibrate more
intensely than other regions. Each segment of the basilar membrane
is “tuned” for a particular pitch. Because the membrane is narrower
and stiffer at the base of the cochlea (closer to the oval window),
high-frequency (high-pitched) sounds induce maximal vibrations in
this region. Toward the apex of the cochlea, the basilar membrane
is wider and more flexible; low-frequency (low-pitched) sounds
cause maximal vibration of the basilar membrane there. Loudness is
determined by the intensity of sound waves. High-intensity sound
waves cause larger vibrations of the basilar membrane, which leads
to a higher frequency of nerve impulses reaching the brain. Louder
sounds also may stimulate a larger number of hair cells.