49 Smell
Olfaction (Smell)
Like taste, the sense of smell, or olfaction, is also responsive to chemical stimuli. The olfactory receptor neurons are located in a small region within the superior nasal cavity (Figure 8.4). This region is referred to as the olfactory epithelium and contains bipolar sensory neurons. Each olfactory sensory neuron has dendrites that extend from the apical surface of the epithelium into the mucus lining the cavity. As airborne molecules are inhaled through the nose, they pass over the olfactory epithelial region and dissolve into the mucus. These odorant molecules bind to proteins that keep them dissolved in the mucus and help transport them to the olfactory dendrites. The odorant–protein complex binds to a receptor protein within the cell membrane of an olfactory dendrite. These receptors are G protein–coupled, and will produce a graded membrane potential in the olfactory neurons. The axon of an olfactory neuron extends from the basal surface of the epithelium, through an olfactory foramen in the cribriform plate of the ethmoid bone, and into the brain. The group of axons called the olfactory tract connect to the olfactory bulb on the ventral surface of the frontal lobe. From there, the axons split to travel to several brain regions. Some travel to the cerebrum, specifically to the primary olfactory cortex that is located in the inferior and medial areas of the temporal lobe. Others project to structures within the limbic system and hypothalamus, where smells become associated with long-term memory and emotional responses. This is how certain smells trigger emotional memories, such as the smell of food associated with one’s birthplace. Smell is the one sensory modality that does not synapse in the thalamus before connecting to the cerebral cortex. This intimate connection between the olfactory system and the cerebral cortex is one reason why smell can be a potent trigger of memories and emotion. The nasal epithelium, including the olfactory cells, can be harmed by airborne toxic chemicals. Therefore, the olfactory neurons are regularly replaced within the nasal epithelium, after which the axons of the new neurons must find their appropriate connections in the olfactory bulb. These new axons grow along the axons that are already in place in the cranial nerve.
Disorders of the Olfactory System: Anosmia
Blunt force trauma to the face, such as that common in many car accidents, can lead to the loss of the olfactory nerve, and subsequently, loss of the sense of smell. This condition is known as anosmia. When the frontal lobe of the brain moves relative to the ethmoid bone, the olfactory tract axons may be sheared apart. Professional fighters often experience anosmia because of repeated trauma to face and head. In addition, certain pharmaceuticals, such as antibiotics, can cause anosmia by killing all the olfactory neurons at once. If no axons are in place within the olfactory nerve, then the axons from newly formed olfactory neurons have no guide to lead them to their connections within the olfactory bulb. There are temporary causes of anosmia, as well, such as those caused by inflammatory responses related to respiratory infections or allergies. Loss of the sense of smell can result in food tasting bland. A person with an impaired sense of smell may require additional spice and seasoning levels for food to be tasted. Anosmia may also be related to some presentations of mild depression, because the loss of enjoyment of food may lead to a general sense of despair. The ability of olfactory neurons to replace themselves decreases with age, leading to age-related anosmia. This explains why some elderly people salt their food more than younger people do. However, this increased sodium intake can increase blood volume and blood pressure, increasing the risk of cardiovascular diseases in the elderly.
Aging and the Nervous System
Anosmia is the loss of the sense of smell. It is often the result of the olfactory nerve being severed, usually because of blunt force trauma to the head. The sensory neurons of the olfactory epithelium have a limited lifespan of approximately one to four months, and new ones are made on a regular basis. The new neurons extend their axons into the CNS by growing along the existing fibers of the olfactory nerve. The ability of these neurons to be replaced is lost with age. Age-related anosmia is not the result of impact trauma to the head, but rather a slow loss of the sensory neurons with no new neurons born to replace them. Smell is an important sense, especially for the enjoyment of food. There are only five tastes sensed by the tongue, and two of them are generally thought of as unpleasant tastes (sour and bitter). The rich sensory experience of food is the result of odor molecules associated with the food, both as food is moved into the mouth, and therefore passes under the nose, and when it is chewed and molecules are released to move up the pharynx into the posterior nasal cavity. Anosmia results in a loss of the enjoyment of food. As the replacement of olfactory neurons declines with age, anosmia can set in. Without the sense of smell, many sufferers complain of food tasting bland. Often, the only way to enjoy food is to add seasoning that can be sensed on the tongue, which usually means adding table salt. The problem with this solution, however, is that this increases sodium intake, which can lead to cardiovascular problems through water retention and the associated increase in blood pressure.
Age Related Changes to Smell
The sense of smell begins to decline in most people by middle age and continues to decline gradually into old age. This is most likely a result of a decrease in olfactory sensory cells and a loss of neurons in the olfactory bulb.