Introduction

• The sense of smell, or olfaction is due to olfactory neurons (receptor cells) in the roof of the vertebrate
nasal cavity.
• These cells, which are specialized endings of the fibers that make up the olfactory nerve (first cranial
nerve), lie among supporting epithelial cells.
• They are densely packed; for example, a dog has up to 40 million olfactory receptor cells per
square centimeter.
• Each olfactory cell ends in a tuft of cilia containing receptor sites for various chemicals.
•Olfactory receptors are regularly replaced.
•This is an exception to the usual (human, mammalian) rule that neurons last a lifetime and are not replaced.
•For an odorant to be detected, molecules of th substance must first diffuse into the air and then pass into
the region of the olfactory epithelium.
• Once there, the odorant dissolves in the watery mucus and then binds to specific odorant receptors.
• Proteins in the mucus may interact with the odorant molecules, transporting them to the receptors and
facilitating their binding to the receptors.
•Although there are many thousands of receptor cells, each contains one or only a few of the 1,000 or so
different receptor types.
• Each type only responds to a specific chemically related group of odorant molecules.
• Each odorant has specific chemical groups that distinguish it from other odorants, and each of these
groups activates a different plasma membrane odorant receptor type.
• Such an interaction alters the membrane permeability and leads to a generator potential.
• In most fishes, openings (external nares) in the snout lead to the olfactory receptors.
• Recent research has revealed that some fishes rely heavily on their sense of smell.
• For example, salmon and lampreys return to spawn in the same streams in which they hatched years
earlier.
• Their migrations to these streams often involve distances of hundreds of miles and are guided by the
fishes' perception of characteristic odors of their spawning stream.
• Olfaction is an important sense for many amphibians.
• It is used in mate recognition, as well as in detecting noxious chemicals and locating food.
• Olfactory senses are better developed in reptiles than in amphibians.
• In addition to having more olfactory epithelium, most reptiles (except crocodilians) possess blind-ending
pouches that open into the mouth.
• These pouches, called Jacobson's (vomeronasal) organs, are best developed in snakes and lizards.
• The protrusible, forked tongues of snakes and lizards are accessory olfactory organs used to sample
airborne chemicals.
• A snake's tongue flicks out and then moves to the Jacobson's organs, which perceive odor molecules.
• Turtles and the tuatara use Jacobson's organs to taste objects held in the mouth.
• Olfaction apparently plays a minor role in the lives of most birds.
• External nares open near the base of the beak, but the olfactory epithelium is poorly developed.
• Vultures are exceptions, in that they locate dead and dying prey largely by smell.
• Many mammals can perceive olfactory stimuli over long distances during either the day or night.
• They use the stimuli to locate food, recognize members of the same species, and avoid predators.
• One example of such a stimulus is a par-ticular class of environmental chemicals called pheromones.
• Pheromones are chemical signals that are released by an animal that affect the behavior of another
animal of the same species.
• Pheromones play an important role in maintaining social hierarchies and stimulating
reproduction in many animals