Theinnerear contains parts (the nonauditory labyrinth or vestibular organ) that are sensitive to acceleration in space,rotation, andorientation in the gravitational field. Rotation is signaled by way of thesemicircular canals, three bony tubes in each ear that lie embedded in the skull roughly at right angles to each other. These canals are filled with fluid calledendolymph; in theampulla of each canal are fine hairs equipped with mechanosensing stereocilia and a kinocilium that project into the cupula, a gelatinous component of theampulla. When rotation begins, the cupula is displaced as the endolymph lags behind, causing the stereocilia to bend toward the kinocilium and therebytransmit signals to thebrain. When rotation is maintained at a steady velocity, the fluid catches up, and stimulation of thehair cells no longer occurs until rotation suddenly stops, again circulating the endolymph. Whenever the hair cells are thus stimulated, one normally experiences asensation of rotation in space. During rotation one exhibits reflexnystagmus (back-and-forth movement) of the eyes. Slow displacement of the eye occurs against the direction of rotation and serves to maintain the gaze at a fixed point in space; this is followed by a quick return to the initial eye position in the direction of the rotation. Stimulation of the hair cells in the absence of actual rotation tends to produce an apparent “swimming” of the visual field, often associated withdizziness andnausea.

Two sacs or enlargements of thevestibule (thesaccule andutricle) react to steady (static) pressures (e.g., those of gravitational forces). Hair cells within these structures, similar to those of the semicircular canal, possess stereocilia and a kinocilium. They also are covered by a gelatinous cap in which are embedded small granular particles ofcalcium carbonate, calledotoliths, that weigh against the hairs. Unusual stimulation of the vestibular receptors and semicircular canals can cause sensory distortions in visual and motor activity. The resultingdiscord between visual and motor responses and the externalspace (as aboard a ship in rough waters) often leads to nausea and disorientation (e.g.,seasickness). In space flight abnormal gravitational and acceleratory forces may contribute to nausea or disequilibrium.

In somediseases (e.g., ear infections), irritation of vestibular nerve endings may cause the affected individual to be subject to falling as well as to spells of disorientation andvertigo. Similarsymptoms may be induced by flushing hot and cold water into the outer opening of the ear, since the temperature changes produce currents in the endolymph of the semicircular canals. This effect is used in clinical tests for vestibular functions and in physiological experiments. Externally applied electrical currents may also stimulate the nerve endings of the vestibule. When a current is applied to the right mastoid bone (just behind the ear), nystagmus to the right tends to occur with a reflex right movement of the head; movement tends to the left for the opposite mastoid. Destruction of the labyrinth in only one ear causes vertigo and other vestibular symptoms, such as nystagmus, inaccurate pointing, and tendency to fall.

Taste (gustatory) sense

The sensory structures for taste are thetaste buds, clusters of cells contained in goblet-shaped structures called papillae that open by a small pore to the mouth cavity. A single taste bud contains about 50 to 75 slender taste receptor cells, all arranged in a banana-like cluster pointed toward the gustatory pore.Taste receptor cells, whichdifferentiate from the surrounding epithelium, are replaced by new cells in a turnover period as short as 7 to 10 days. The various types of cells in the taste bud appear to be different stages in this turnover process. Slender nerve fibres entwine among and make contact usually with many cells. Taste buds are located primarily infungiform (mushroom-shaped),foliate, andcircumvallate (walled-around) papillae of thetongue or inadjacent structures of thepalate andthroat. Many gustatory receptors in small papillae on thesoft palate and back roof of the mouth in adults are particularly sensitive to sour and bitter tastes, whereas the tongue receptors are relatively more sensitive to sweet and salty tastes. Some loss of taste sensitivity suffered among denture wearers may occur because of mechanical interference of the dentures with taste receptors on the roof of the mouth.

Nerve supply

There is no single sensory nerve for taste. The anterior (front) two-thirds of the tongue is supplied by one nerve (thelingual nerve), the back of the tongue by another (theglossopharyngeal nerve), and the throat and larynx by certain branches of a third (thevagus nerve), all of which subserve touch, temperature, andpain sensitivity in the tongue, as well as taste. The gustatory fibres of the anterior tongue leave the lingual nerve to form thechorda tympani, a slender nerve thattraverses the eardrum on the way to the brainstem. When the chorda tympani at one ear is cut or damaged (by injury to the eardrum), taste buds begin to disappear and gustatory sensitivity is lost on the anterior two-thirds of the tongue on the same side. The taste fibres from all the sensory nerves from the mouth come together in themedulla oblongata. Here and at all levels of the brain, gustatory fibres run in distinct and separate pathways, lying close to the pathways for othermodalities from the tongue and mouth cavity. From themedulla, the gustatory fibres ascend by a pathway to a small cluster of cells in the thalamus and then to a taste-receiving area in the anteriorcerebral cortex.