Just as we do not actually smell with the bumps on our faces called noses, neither do we perceive sound solely with the flaps we call ears. Although hearing begins with the ear flap or pinna, the receptor cells that change sound energy into the electrical currency of the nervous system lie deep inside the temporal bone of the skull. Like olfactory cells that detect odors, auditory receptor cells (hair cells) are recessed from the surface of the body. Unlike olfactory or taste receptors, however, hair cells are not renewed when they die or are damaged. Although taste and olfactory cells interact directly with molecules in the environment, auditory receptors are quite far removed from the phenomena they detect. Sound waves are converted into vibrations in a fluid in the inner ear, and these vibrations indirectly move the hair cells, which then send electrical signals to the brain.
Sound activates the external, middle, and inner ear.
External Ear
The folds and ridges of the pinna channel sound efficiently into the ear canal and to the eardrum, or tympanic membrane, at its end. The pattern of folds captures sounds in a way that helps us localize the origin of sound in space, especially on the vertical axis. The ear canal carries sound to the eardrum, and its lining produces ear wax to keep the eardrum and canal from drying out and to trap dirt before it gets to the eardrum. When sound waves vibrate the eardrum, sound energy is transferred to the middle ear.
Middle Ear
The middle ear is a small, air-filled pocket bounded by the eardrum on one side and the oval window of the inner ear on the other. This pocket is connected to the common mouth and nasal cavity, or pharynx, by the Eustachian tube. The Eustachian tube allows air pressure to equalize between the outside of the eardrum (surrounding atmosphere) and the inside of the eardrum (the middle ear). This is also the pathway that allows infections from the mouth and nose cavities to enter the middle ear, causing the common ear infections of childhood. The middle ear houses the 3 smallest bones in the body, the malleus, incus, and stapes (hammer, anvil, and stirrup), which form a chain of levers connected by joints. The malleus is attached to the eardrum by ligaments, as is the stapes to the oval window. Thus, this series of membranes and bones forms a pathway that carries vibrations from the eardrum to the inner ear. The stapes, the last bone in the chain, pulls or pushes on the membranous oval window when the eardrum and the 3 bones are vibrated by sound waves; the oval window is a closed membrane, but acts as the entrance to the inner ear for sound energy.
Inner ear
What does this pulling and pushing on the oval window do in the inner ear? A look at the structure of this area helps show how sound wave energy is transmitted to fluid in the inner ear.
The inner ear is composed of the cochlea, from the Greek word for snail, and the semicircular canals. (These latter canals are part of the vestibular system for balance and will not be considered here.) The cochlea is a membranous tube that is covered by a very thin layer of bone and wound around a tiny central bone (the modiolus) into a shape that resembles a snail; it is only about nine millimeters across - well under one-half inch. The cochlea is filled with a special fluid, and the pushing and pulling of the stapes on the oval window moves the fluid in this coiled tube.
If we stretch out the cochlear tube, we see that inside are actually 3 tubes, two larger and one smaller, with the small tube, (the scala media) lying between the two larger ones. All three tubes are filled with fluids, which vary somewhat in composition.
The cochlea is a tube made of three inner tubes: the scala vestibuli, scala media, and scala tympani.
The fluid in the top-most tube is set in motion by the piston-like movements of the stapes on the oval window. The vibrations travel into the fluid of the upper tube of the cochlea and around the tip of the organ into the fluid of the lower tube. The pushing or pulling of the oval window on this fluid must have a release or dampening mechanism: this is provided by the round window, a membrane located at the end of the lower of the large tubes.
Forming the lengthwise partition between the lower large tube and the small tube is the basilar membrane. On this membrane sit the stars of the show in the auditory system, the auditory receptor cells, or hair cells. When the basilar membrane moves, it stimulates the hair cells, which then send signals about sounds to the brain.
We can summarize the workings of the ear as follows:
The pinna captures sound waves and channels them through the ear canal to the eardrum.
Vibrations of the eardrum pass along the three bones of the middle ear, with the base of the stapes then rocking the oval window in and out.
The membranous oval window acts something like a piston in a hydraulic system: it pushes and pulls on the enclosed fluid of the cochlea.
The fluid vibrations move the basilar membrane, and this motion activates auditory receptor cells (hair cells) sitting on the membrane, which send signals to the brain.
Hair cells sit on the basilar membrane and are innervated by fibers from the auditory nerve, one of the cranial nerves.
4. The basilar membrane distributes vibrations to hair cells
The motion of the fluid in the cochlear tubes sets the basilar membrane in motion, generating traveling waves along its length. These are somewhat like the waves produced in a long rope that is grasped at one end and flicked. The basilar membrane is much more complicated, though. To begin with, it is not uniform throughout its length, but rather is relatively wide and thin at the apex (top) of the cochlea, and narrow but thick at the base. Because of these properties, a sound wave in the cochlear fluid produces a peak amplitude or height of displacement of the membrane at a certain point along its length. This point is determined by the frequency (number of waves per unit time) of the sound that originally produced the fluid motion. High frequencies cause a peak wave near the base (narrow part of the membrane), and low frequencies produce their peaks toward the apex (broad part of the membrane). Thus, the basilar membrane is sometimes called a frequency analyzer. In addition, the hair cells on the membrane are also tuned to particular frequencies, so that each hair cell responds best to sound of a given frequency.
This anatomy or "geography" of the basilar membrane and hair cells produces a tonotopic map along the membrane. This means that, as with geographic maps, once you know some landmarks and the scale of the map, you can calculate the point where sound of a particular frequency will have its peak, because the system is ordered and predictable. Further, groups of responding neurons in the brain auditory areas also contain tonotopic maps.
5. Hair cells encode sounds and transmit this information to neurons
The hair cells sit on an epithelial ridge called the organ of Corti on the basilar membrane; the ridge contains several other types of cells that support the hair cells. The receptor cells are called hair cells not because they sprout hairs, but because their apical or top ends are covered with cilia, which under the microscope look a bit like hairs. Over the top of the cilia lies a gelatinous membrane, sandwiching the hair cells between itself and the basilar membrane. The complex, relative movements of these two membranes activate the cilia of hair cells, causing the cells to undergo a change in the electrical potential across their cell membranes. When specific changes occur in this electrical state, neurotransmitter molecules are released from the bottom or basal parts of the hair cells. Thus, the cilia are essential in transducing, or changing, the mechanical energy of the basilar membrane into electrical changes in the hair cells. As mentioned above, hair cells are tuned to the particular frequencies that activate the portion of the basilar membrane where they reside.
Hair cells are modified epithelial cells and do not have dendrites and axons as neurons do, but they communicate, as many neurons do, by releasing neurotransmitter. They release the neurotransmitter at junctions or synapses that they form on branches from neurons whose cell bodies are in a ganglion (group of neurons) just outside the cochlea. The axons from the ganglion neurons form the auditory nerve, which carries signals into the first stop in the brain, the cochlear nucleus.
6. Sound information from each ear is distributed to both sides of the brain
Once information from one ear goes to the cochlear nucleus on that side of the head, the neurons in this nucleus send information to identical higher centers on both sides of the brain. Some of the processing stations are the superior olivary nucleus, inferior colliculus (in the midbrain), medial geniculate nucleus (in the thalamus), and the auditory cortex. From the auditory cortex, messages go to other areas of the cerebral cortex for interpretation of the meaning of sounds.
Signals from neurons that get information directly from hair cells travel in the auditory nerve to the brainstem. Here the signals activate more neurons, which send the auditory messages on to the thalamus, then to the auditory cortex in the temporal lobe of the cerebrum.
In the cochlear nucleus, the first brain relay station for sound, signals encoding sounds are not just passed on, but rather are "dissected" and sorted first. This means that different features of a sound, such as frequency, intensity, or onset and offset (beginning and ending of a sound) are carried to higher brain centers separately. This sorting out of the features of stimuli and sending messages forward in parallel nerve pathways is a common and important attribute of brain sensory systems. One of the big tasks of researchers is to find out how areas in the cerebral cortex use input from these parallel pathways to interpret the original sensations - in this case, the original sounds.
7. We need two ears to locate a sound source
When a sound occurs at the extreme left of a subject, the arrival of the sound at the left ear is about 600 to 700 microseconds (millionths of a second) earlier than at the right ear. Further, the head acts as a sound barrier, so the sound is a little louder in the left ear. How does the timing and intensity of sound in the two ears tell the brain where the sound source is?
This processing is carried out in the superior olivary nucleus (SON) of the brainstem. Axons coming into the SON from the cochlear nuclei form synapses successively across a linear series of SON neurons. Each neuron here gets messages from cochlear neurons in both ears, and in order to fire a signal to higher brain centers, each must receive simultaneous messages from the two cochlear nuclei. Because a sound from, for example, the extreme left side of a person arrives later in the right ear than the left, hair cells and neurons from the right ear send their signals slightly later than those from the left ear. Each SON neuron is activated only by simultaneous input from the two ears, so that when, for example, signals from axon d1 from the right and d2 from the left ear coincide on SON neuron d, it fires. Through experience, we learn that when neuron d fires, the sound is, for example, at 50o to the left of straight ahead. If neuron b fires, this could mean that the sound originates at 20o to the left.
Neurons in another part of the SON employ intensity cues rather than arrival-time cues. Again, the neurons need simultaneous input from the two ears to fire, but in addition, they respond best when the sound intensity on one side of the head exceeds that on the other by a certain amount.
Note that it is hard to differentiate sounds coming from directly in front of you from those originating directly behind you. Both sounds are equal distance from the two ears, so there is no difference in timing and intensity, information that our brains need to localize sound in the horizontal dimension.
Sound is processed in the superior olivary nucleus (SON). A sound arriving earlier at the left ear elicits signals more quickly in the SON than those from the right ear. At some point, as the signals from the two ears travel across the linear array of neurons in the SON, they converge on one neuron and activate it. This processing is carried out simultaneously in the left and right SONs.