19.6: Hair Cells
Hair cells are the sensory receptors of the auditory system—they transduce mechanical sound waves into electrical energy that the nervous system can understand. Hair cells are located in the organ of Corti within the cochlea of the inner ear, between the basilar and tectorial membranes. The actual sensory receptors are called inner hair cells. The outer hair cells serve other functions, such as sound amplification in the cochlea, and are not discussed in detail here.
Hair cells are named after the hair-like stereocilia that protrude from their tops and touch the tectorial membrane. The stereocilia are arranged by height and are attached by thin filaments called tip links. The tip links are connected to stretch-activated cation channels on the tips of the stereocilia.
When a sound wave vibrates the basilar membrane, it creates a shearing force between the basilar and tectorial membranes that moves the hair cell stereocilia from side to side. When the cilia are displaced towards the tallest cilium, the tip links stretch, opening the cation channels. Potassium (K+) then flows into the cell, because there is a very high concentration of K+ in the fluid outside of the stereocilia. This large voltage difference creates an electrochemical gradient that causes an influx of K+ once the channels are opened.
This influx of positive charge depolarizes the cell, increasing the voltage across the membrane. This causes voltage-gated calcium (Ca2+) channels in the cell body to open, and Ca2+ flows into the cell. Ca2+ triggers a signaling cascade that causes synaptic vesicles containing excitatory neurotransmitter molecules to fuse to the cell membrane and be released, exciting the postsynaptic auditory nerve cell and increasing the transmission of action potentials to the brain. When the stereocilia are pushed in the opposite direction, towards the shortest stereocilia, the tip links relax, the cation channels close, and the cell becomes hyperpolarized (i.e., the membrane potential is more negative) compared to its resting state.
Characteristics of the sound wave, such as frequency, are encoded in the pattern of hair cell activation and, consequently, auditory nerve cell activation. This information is then sent to the brain for interpretation.