The sense of smell, or olfaction, is the
special sense
In medicine and anatomy, the special senses are the senses that have specialized organs devoted to them:
* vision (the eye)
* hearing and balance (the ear, which includes the auditory system and vestibular system)
* smell (the nose)
* taste (th ...
through which smells (or odors) are perceived.
The sense of smell has many functions, including detecting desirable foods, hazards, and
pheromone
A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s, and plays a role in
taste
The gustatory system or sense of taste is the sensory system that is partially responsible for the perception of taste (flavor). Taste is the perception produced or stimulated when a substance in the mouth reacts chemically with taste receptor ...
.
In humans, it occurs when an
odor
An odor (American English) or odour (English in the Commonwealth of Nations, Commonwealth English; American and British English spelling differences#-our, -or, see spelling differences) is caused by one or more volatilized chemical compounds ...
binds to a
receptor
Receptor may refer to:
* Sensory receptor, in physiology, any structure which, on receiving environmental stimuli, produces an informative nerve impulse
*Receptor (biochemistry), in biochemistry, a protein molecule that receives and responds to a ...
within the
nasal cavity
The nasal cavity is a large, air-filled space above and behind the nose in the middle of the face. The nasal septum divides the cavity into two cavities, also known as fossae. Each cavity is the continuation of one of the two nostrils. The nasal c ...
, transmitting a signal through the
olfactory system
The olfactory system, or sense of smell, is the sensory system used for smelling ( olfaction). Olfaction is one of the special senses, that have directly associated specific organs. Most mammals and reptiles have a main olfactory system and an ...
.
Glomeruli
''Glomerulus'' () is a common term used in anatomy to describe globular structures of entwined vessels, fibers, or neurons. ''Glomerulus'' is the diminutive of the Latin ''glomus'', meaning "ball of yarn".
''Glomerulus'' may refer to:
* the filter ...
aggregate signals from these receptors and transmit them to the
olfactory bulb
The olfactory bulb (Latin: ''bulbus olfactorius'') is a neural structure of the vertebrate forebrain involved in olfaction, the sense of smell. It sends olfactory information to be further processed in the amygdala, the orbitofrontal cortex ( ...
, where the sensory input will start to interact with parts of the brain responsible for smell identification,
memory
Memory is the faculty of the mind by which data or information is encoded, stored, and retrieved when needed. It is the retention of information over time for the purpose of influencing future action. If past events could not be remembered, ...
, and
emotion
Emotions are mental states brought on by neurophysiological changes, variously associated with thoughts, feelings, behavioral responses, and a degree of pleasure or displeasure. There is currently no scientific consensus on a definition. ...
.
There are many different causes for alteration,
lack
Lack may refer to:
Places
* Lack, County Fermanagh, a townland in Northern Ireland
* Lack, Poland
* Łąck, Poland
* Lack Township, Juniata County, Pennsylvania, US
Other uses
* Lack (surname)
* Lack (manque), a term in Lacan's psychoanalyti ...
, or disturbance to a normal sense of smell, and can include damage to the
nose
A nose is a protuberance in vertebrates that houses the nostrils, or nares, which receive and expel air for respiration alongside the mouth. Behind the nose are the olfactory mucosa and the sinuses. Behind the nasal cavity, air next passes th ...
or smell receptors, or central problems affecting the brain. Some causes include
upper respiratory infections
An upper respiratory tract infection (URTI) is an illness caused by an acute infection, which involves the upper respiratory tract, including the nose, sinuses, pharynx, larynx or trachea. This commonly includes nasal obstruction, sore throat, ...
,
traumatic brain injury
A traumatic brain injury (TBI), also known as an intracranial injury, is an injury to the brain caused by an external force. TBI can be classified based on severity (ranging from mild traumatic brain injury TBI/concussionto severe traumatic b ...
, and
neurodegenerative disease
A neurodegenerative disease is caused by the progressive loss of structure or function of neurons, in the process known as neurodegeneration. Such neuronal damage may ultimately involve cell death. Neurodegenerative diseases include amyotrophic ...
.
History of study
Early scientific study of the sense of smell includes the extensive doctoral dissertation of
Eleanor Gamble
Eleanor Acheson McCulloch Gamble (March 2, 1868 – August 30, 1933) was an influential American psychologist from the late 19th century through the early 20th century. Gamble published most of her work on audition and memory influenced by Georg ...
, published in 1898, which compared olfactory to other
stimulus modalities Stimulus modality, also called sensory modality, is one aspect of a stimulus or what is perceived after a stimulus. For example, the temperature modality is registered after heat or cold stimulate a receptor. Some sensory modalities include: light, ...
, and implied that smell had a lower intensity discrimination.
As the Epicurean and atomistic Roman philosopher
Lucretius
Titus Lucretius Carus ( , ; – ) was a Roman poet and philosopher. His only known work is the philosophical poem ''De rerum natura'', a didactic work about the tenets and philosophy of Epicureanism, and which usually is translated into E ...
(1stcentury BCE) speculated, different odors are attributed to different shapes and sizes of "atoms" (odor molecules in the modern understanding) that stimulate the olfactory organ.
A modern demonstration of that theory was the cloning of olfactory receptor proteins by
Linda B. Buck
Linda Brown Buck (born January 29, 1947) is an American biologist best known for her work on the olfactory system. She was awarded the 2004 Nobel Prize in Physiology or Medicine, along with Richard Axel, for their work on olfactory receptors. She ...
and
Richard Axel
Richard Axel (born July 2, 1946) is an American molecular biologist and List of university professors at Columbia University, university professor in the Department of Neuroscience at Columbia University and investigator at the Howard Hughes Medic ...
(who were awarded the
Nobel Prize
The Nobel Prizes ( ; sv, Nobelpriset ; no, Nobelprisen ) are five separate prizes that, according to Alfred Nobel's will of 1895, are awarded to "those who, during the preceding year, have conferred the greatest benefit to humankind." Alfr ...
in 2004), and subsequent pairing of odor molecules to specific receptor proteins. Each odor receptor molecule recognizes only a particular molecular feature or class of odor molecules.
Mammals
Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
have about a thousand
gene
In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
s that code for
odor reception. Of the genes that code for odor receptors, only a portion are functional. Humans have far fewer active odor receptor genes than other primates and other mammals. In mammals, each
olfactory receptor neuron
An olfactory receptor neuron (ORN), also called an olfactory sensory neuron (OSN), is a sensory neuron within the olfactory system.
Structure
Humans have between 10 and 20 million olfactory receptor neurons (ORNs). In vertebrates, ORNs are b ...
expresses only one functional odor receptor. Odor receptor nerve cells function like a key–lock system: if the airborne molecules of a certain chemical can fit into the lock, the nerve cell will respond.
There are, at present, a number of competing theories regarding the mechanism of odor coding and perception. According to the
shape theory, each receptor detects a feature of the odor
molecule
A molecule is a group of two or more atoms held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions which satisfy this criterion. In quantum physics, organic chemistry, and bioch ...
. The weak-shape theory, known as the
odotope theory, suggests that different receptors detect only small pieces of molecules, and these minimal inputs are combined to form a larger olfactory perception (similar to the way visual perception is built up of smaller, information-poor sensations, combined and refined to create a detailed overall perception).
According to a new study, researchers have found that a functional relationship exists between molecular volume of odorants and the olfactory neural response. An alternative theory, the
vibration theory proposed by
Luca Turin
Luca Turin (born 20 November 1953) is a biophysicist and writer with a long-standing interest in bioelectronics, the sense of smell, perfumery, and the fragrance industry.
Early life and education
Turin was born in Beirut, Lebanon on 20 November ...
, posits that odor receptors detect the frequencies of vibrations of odor molecules in the infrared range by
quantum tunnelling
Quantum tunnelling, also known as tunneling ( US) is a quantum mechanical phenomenon whereby a wavefunction can propagate through a potential barrier.
The transmission through the barrier can be finite and depends exponentially on the barrier h ...
. However, the behavioral predictions of this theory have been called into question. There is no theory yet that explains olfactory perception completely.
Functions
Taste
Flavor perception is an aggregation of
auditory,
taste
The gustatory system or sense of taste is the sensory system that is partially responsible for the perception of taste (flavor). Taste is the perception produced or stimulated when a substance in the mouth reacts chemically with taste receptor ...
,
haptic, and smell sensory information.
Retronasal smell Retronasal smell, retronasal olfaction, is the ability to perceive flavor dimensions of foods and drinks. Retronasal smell is a sensory modality that produces flavor. It is best described as a combination of traditional smell (orthonasal smell) and ...
plays the biggest role in the sensation of flavor. During the process of
mastication
Chewing or mastication is the process by which food is crushed and ground by teeth. It is the first step of digestion, and it increases the surface area of foods to allow a more efficient break down by enzymes. During the mastication process, th ...
, the tongue manipulates food to release odorants. These odorants enter the nasal cavity during exhalation.
The smell of food has the sensation of being in the mouth because of co-activation of the motor cortex and olfactory epithelium during mastication.
Smell,
taste
The gustatory system or sense of taste is the sensory system that is partially responsible for the perception of taste (flavor). Taste is the perception produced or stimulated when a substance in the mouth reacts chemically with taste receptor ...
, and
trigeminal
In neuroanatomy, the trigeminal nerve ( lit. ''triplet'' nerve), also known as the fifth cranial nerve, cranial nerve V, or simply CN V, is a cranial nerve responsible for sensation in the face and motor functions such as biting and chewing; ...
receptors (also called
chemesthesis
Chemesthesis is the chemical sensitivity of the skin and mucous membranes. Chemesthetic sensations arise when chemical compounds activate receptors associated with other senses that mediate pain, touch, and thermal perception. These chemical-ind ...
) together contribute to
flavor
Flavor or flavour is either the sensory perception of taste or smell, or a flavoring in food that produces such perception.
Flavor or flavour may also refer to:
Science
*Flavors (programming language), an early object-oriented extension to Lis ...
. The human
tongue
The tongue is a muscular organ (anatomy), organ in the mouth of a typical tetrapod. It manipulates food for mastication and swallowing as part of the digestive system, digestive process, and is the primary organ of taste. The tongue's upper surfa ...
can distinguish only among five distinct qualities of taste, while the nose can distinguish among hundreds of substances, even in minute quantities. It is during
exhalation
Exhalation (or expiration) is the flow of the breath out of an organism. In animals, it is the movement of air from the lungs out of the airways, to the external environment during breathing.
This happens due to elastic properties of the lungs, ...
that the smell's contribution to flavor occurs, in contrast to that of proper smell, which occurs during the
inhalation
Inhalation (or Inspiration) happens when air or other gases enter the lungs.
Inhalation of air
Inhalation of air, as part of the cycle of breathing, is a vital process for all human life. The process is autonomic (though there are exceptions ...
phase of breathing.
The olfactory system is the only human sense that bypasses the
thalamus
The thalamus (from Greek θάλαμος, "chamber") is a large mass of gray matter located in the dorsal part of the diencephalon (a division of the forebrain). Nerve fibers project out of the thalamus to the cerebral cortex in all directions, ...
and connects directly to the
forebrain
In the anatomy of the brain of vertebrates, the forebrain or prosencephalon is the Anatomical terms of location#Directional terms, rostral (forward-most) portion of the brain. The forebrain (prosencephalon), the midbrain (mesencephalon), and hin ...
.
Hearing
Smell and
sound
In physics, sound is a vibration that propagates as an acoustic wave, through a transmission medium such as a gas, liquid or solid.
In human physiology and psychology, sound is the ''reception'' of such waves and their ''perception'' by the ...
information has been shown to converge in the olfactory tubercles of
rodents. This neural convergence is proposed to give rise to a perception termed
smound.
Whereas a
flavor
Flavor or flavour is either the sensory perception of taste or smell, or a flavoring in food that produces such perception.
Flavor or flavour may also refer to:
Science
*Flavors (programming language), an early object-oriented extension to Lis ...
results from interactions between smell and taste, a smound may result from interactions between smell and sound.
Inbreeding avoidance
The
MHC genes (known as
HLA in humans) are a group of genes present in many animals and important for the
immune system
The immune system is a network of biological processes that protects an organism from diseases. It detects and responds to a wide variety of pathogens, from viruses to parasitic worms, as well as cancer cells and objects such as wood splinte ...
; in general, offspring from parents with differing MHC genes have a stronger immune system. Fish, mice, and female humans are able to smell some aspect of the MHC genes of potential sex partners and prefer partners with MHC genes different from their own.
Humans can detect blood relatives from olfaction. Mothers can identify by body odor their biological children but not their stepchildren. Pre-adolescent children can olfactorily detect their full siblings but not half-siblings or step siblings, and this might explain
incest avoidance and the
Westermarck effect
The Westermarck effect, also known as reverse sexual imprinting, is a psychological hypothesis that people tend not to be attracted to peers with whom they lived like siblings before age six. This hypothesis was first proposed by Finnish anthr ...
. Functional imaging shows that this olfactory kinship detection process involves the frontal-temporal junction, the
insula, and the dorsomedial
prefrontal cortex
In mammalian brain anatomy, the prefrontal cortex (PFC) covers the front part of the frontal lobe of the cerebral cortex. The PFC contains the Brodmann areas BA8, BA9, BA10, BA11, BA12, BA13, BA14, BA24, BA25, BA32, BA44, BA45, BA46, ...
, but not the primary or secondary olfactory cortices, or the related
piriform cortex
The piriform cortex, or pyriform cortex, is a region in the brain, part of the rhinencephalon situated in the cerebrum. The function of the piriform cortex relates to the sense of smell.
Structure
The piriform cortex is part of the rhinencepha ...
or
orbitofrontal cortex
The orbitofrontal cortex (OFC) is a prefrontal cortex region in the frontal lobes of the brain which is involved in the cognitive process of decision-making. In non-human primates it consists of the association cortex areas Brodmann area 11, 12 ...
.
Since
inbreeding
Inbreeding is the production of offspring from the mating or breeding of individuals or organisms that are closely related genetically. By analogy, the term is used in human reproduction, but more commonly refers to the genetic disorders and o ...
is detrimental, it tends to be avoided. In the house mouse, the
major urinary protein (MUP) gene cluster provides a highly polymorphic scent signal of genetic identity that appears to underlie
kin recognition Kin recognition, also called kin detection, is an organism's ability to distinguish between close genetic kin and non-kin. In evolutionary biology and psychology, such an ability is presumed to have evolved for inbreeding avoidance, though animals d ...
and inbreeding avoidance. Thus, there are fewer matings between mice sharing MUP haplotypes than would be expected if there were random mating.
Guiding movement
Some animals use
scent trails to guide movement, for example social insects may lay down a trail to a food source, or a
tracking dog
Tracking refers to a dog's ability to detect, recognize and follow a specific scent. Possessing heightened olfactory abilities, dogs, especially scent hounds, are able to detect, track and locate the source of certain odours.Hepper, P. G., & Wells ...
may follow the scent of its target.
Genetics
Different people smell different odors, and most of these differences are caused by genetic differences. Although
odorant receptor
Olfactory receptors (ORs), also known as odorant receptors, are chemoreceptors expressed in the cell membranes of olfactory receptor neurons and are responsible for the detection of odorants (for example, compounds that have an odor) which give ri ...
genes make up one of the largest gene families in the human genome, only a handful of genes have been linked conclusively to particular smells. For instance, the odorant receptor
OR5A1
Olfactory receptor 5A1 is a protein that in humans is encoded by the ''OR5A1'' gene.
Olfactory receptors interact with odorant molecules in the nose, to initiate a neuronal response that triggers the perception of a smell. The olfactory receptor p ...
and its genetic variants (alleles) are responsible for our ability (or failure) to smell β-
ionone
The ionones are a series of closely related chemical substances that are part of a group of compounds known as rose ketones, which also includes damascones and damascenones. Ionones are aroma compounds found in a variety of essential oils, inclu ...
, a key aroma in foods and beverages.
Similarly, the odorant receptor
OR2J3
Olfactory receptor 2J3 is a protein that in humans is encoded by the ''OR2J3'' gene.
Olfactory receptors interact with odorant molecules in the nose, to initiate a neuronal response that triggers the perception of a smell. The olfactory receptor p ...
is associated with the ability to detect the "grassy" odor, cis-3-hexen-1-ol.
The preference (or dislike) of
(coriander) has been linked to the olfactory receptor
OR6A2
Olfactory receptor 6A2 is a protein that in humans is encoded by the ''OR6A2'' gene. It is Class II (tetrapod-specific) olfactory receptor and a rhodopsin-like receptor.
Function
Olfactory receptors interact with odorant molecules in the nose, ...
.
Variability amongst vertebrates
The importance and sensitivity of smell varies among different organisms; most
mammal
Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s have a good sense of smell, whereas most
bird
Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
s do not, except the
tubenoses (e.g.,
petrel
Petrels are tube-nosed seabirds in the bird order Procellariiformes.
Description
The common name does not indicate relationship beyond that point, as "petrels" occur in three of the four families within that group (all except the albatross f ...
s and
albatross
Albatrosses, of the biological family Diomedeidae, are large seabirds related to the procellariids, storm petrels, and diving petrels in the order Procellariiformes (the tubenoses). They range widely in the Southern Ocean and the North Pacifi ...
es), certain species of new world
vultures
A vulture is a bird of prey that scavenges on carrion. There are 23 extant species of vulture (including Condors). Old World vultures include 16 living species native to Europe, Africa, and Asia; New World vultures are restricted to North and ...
, and the
kiwi
Kiwi most commonly refers to:
* Kiwi (bird), a flightless bird native to New Zealand
* Kiwi (nickname), a nickname for New Zealanders
* Kiwifruit, an edible berry
* Kiwi dollar or New Zealand dollar, a unit of currency
Kiwi or KIWI may also ref ...
s. Also, birds have hundreds of olfactory receptors. Although, recent analysis of the chemical composition of
volatile organic compounds
Volatile organic compounds (VOCs) are organic compounds that have a high vapour pressure at room temperature
Colloquially, "room temperature" is a range of air temperatures that most people prefer for indoor settings. It feels comfortable to a ...
(VOCs) from
king penguin
The king penguin (''Aptenodytes patagonicus'') is the second largest species of penguin, smaller, but somewhat similar in appearance to the emperor penguin. There are two subspecies: ''A. p. patagonicus'' and ''A. p. halli''; ''patagonicus'' ...
feathers suggest that VOCs may provide olfactory cues, used by the penguins to locate their colony and recognize individuals. Among mammals, it is well developed in the
carnivore
A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other sof ...
s and
ungulate
Ungulates ( ) are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. These include odd-toed ungulates such as horses, rhinoceroses, and tapirs; and even-toed ungulates such as cattle, pigs, giraffes, cam ...
s, which must always be aware of each other, and in those that smell for their food, such as
moles Moles can refer to:
* Moles de Xert, a mountain range in the Baix Maestrat comarca, Valencian Community, Spain
* The Moles (Australian band)
*The Moles, alter ego of Scottish band Simon Dupree and the Big Sound
People
*Abraham Moles, French engin ...
. Having a strong sense of smell is referred to as ''macrosmatic''.
Figures suggesting greater or lesser sensitivity in various species reflect experimental findings from the reactions of animals exposed to aromas in known extreme dilutions. These are, therefore, based on perceptions by these animals, rather than mere nasal function. That is, the brain's smell-recognizing centers must react to the stimulus detected for the animal to be said to show a response to the smell in question. It is estimated that dogs, in general, have an olfactory sense approximately ten thousand to a hundred thousand times more acute than a human's. This does not mean they are overwhelmed by smells our noses can detect; rather, it means they can discern a molecular presence when it is in much greater dilution in the carrier, air.
Scenthound
Franz Rudolf Frisching in the uniform of an officer of the Bernese Huntsmen Corps with his Berner Laufhund, painted by Jean Preudhomme in 1785
Scent hounds (or scenthounds) are a Dog type, type of hound that primarily hunts by scent rather than ...
s as a group can smell one- to ten-million times more acutely than a human, and
bloodhound
The bloodhound is a large scent hound, originally bred for hunting deer, wild boar and, since the Middle Ages, for tracking people. Believed to be descended from hounds once kept at the Abbey of Saint-Hubert, Belgium, in French it is called, ...
s, which have the keenest sense of smell of any dogs,
have noses ten- to one-hundred-million times more sensitive than a human's. They were bred for the specific purpose of tracking humans, and can detect a
scent trail
An odor (American English) or odour (Commonwealth English; see spelling differences) is caused by one or more volatilized chemical compounds that are generally found in low concentrations that humans and animals can perceive via their sense ...
a few days old. The second-most-sensitive nose is possessed by the
Basset Hound
The Basset Hound is a short-legged breed of dog in the hound family. The Basset is a scent hound that was originally bred for the purpose of hunting hare. Their sense of smell and ability to ''ground-scent'' is second only to the Bloodhound.Har ...
, which was bred to track and hunt rabbits and other small animals.
Grizzly bear
The grizzly bear (''Ursus arctos horribilis''), also known as the North American brown bear or simply grizzly, is a population or subspecies of the brown bear inhabiting North America.
In addition to the mainland grizzly (''Ursus arctos horri ...
s have a sense of smell seven times stronger than that of the bloodhound, essential for locating food underground. Using their elongated claws, bears dig deep trenches in search of burrowing animals and nests as well as roots, bulbs, and insects. Bears can detect the scent of food from up to eighteen miles away; because of their immense size, they often scavenge new kills, driving away the predators (including packs of wolves and human hunters) in the process.
The sense of smell is less developed in the
catarrhine
The parvorder Catarrhini , catarrhine monkeys, Old World anthropoids, or Old World monkeys, consisting of the Cercopithecoidea and apes (Hominoidea). In 1812, Geoffroy grouped those two groups together and established the name Catarrhini, "Old ...
primate
Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...
s, and nonexistent in
cetacea
Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel them ...
ns, which compensate with a well-developed sense of
taste
The gustatory system or sense of taste is the sensory system that is partially responsible for the perception of taste (flavor). Taste is the perception produced or stimulated when a substance in the mouth reacts chemically with taste receptor ...
.
In some
strepsirrhines
Strepsirrhini or Strepsirhini (; ) is a suborder of primates that includes the lemuriform primates, which consist of the lemurs of Madagascar, galagos ("bushbabies") and pottos from Africa, and the lorises from India and southeast Asia. Colle ...
, such as the
red-bellied lemur
The red-bellied lemur (''Eulemur rubriventer'') is a medium-sized strepsirrhine primate with a luxuriant chestnut brown coat. This lemur is endemic to eastern Madagascan rainforests and is distinguished by patches of white skin below the e ...
, scent glands occur atop the head. In many species, smell is highly tuned to
pheromone
A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s; a male
silkworm
The domestic silk moth (''Bombyx mori''), is an insect from the moth family Bombycidae. It is the closest relative of ''Bombyx mandarina'', the wild silk moth. The silkworm is the larva or caterpillar of a silk moth. It is an economically imp ...
moth, for example, can sense a single molecule of
bombykol
Bombykol is a pheromone released by the female silkworm moth to attract mates. It is also the sex pheromone in the wild silk moth (''Bombyx mandarina''). Discovered by Adolf Butenandt
Adolf Friedrich Johann Butenandt (; 24 March 1903 – 18 J ...
.
Fish, too, have a well-developed sense of smell, even though they inhabit an aquatic environment. Salmon utilize their sense of smell to identify and return to their home stream waters. Catfish use their sense of smell to identify other individual catfish and to maintain a social hierarchy. Many fishes use the sense of smell to identify mating partners or to alert to the presence of food.
Human smell abilities
Although conventional wisdom and lay literature, based on impressionistic findings in the 1920s, have long presented human smell as capable of distinguishing between roughly 10,000 unique odors, recent research has suggested that the average individual is capable of distinguishing over one trillion unique odors. Researchers in the most recent study, which tested the psychophysical responses to combinations of over 128 unique odor molecules with combinations composed of up to 30 different component molecules, noted that this estimate is "conservative" and that some subjects of their research might be capable of deciphering between a thousand trillion odorants, adding that their worst performer could probably still distinguish between 80million scents. Authors of the study concluded, "This is far more than previous estimates of distinguishable olfactory stimuli. It demonstrates that the human olfactory system, with its hundreds of different olfactory receptors, far out performs the other senses in the number of physically different stimuli it can discriminate." However, it was also noted by the authors that the ability to distinguish between smells is not analogous to being able to consistently identify them, and that subjects were not typically capable of identifying individual odor stimulants from within the odors the researchers had prepared from multiple odor molecules. In November 2014 the study was strongly criticized by Caltech scientist Markus Meister, who wrote that the study's "extravagant claims are based on errors of mathematical logic." The logic of his paper has in turn been criticized by the authors of the original paper.
Physiological basis in vertebrates
Main olfactory system
In humans and other
vertebrate
Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
s, smells are sensed by
olfactory sensory neuron
An olfactory receptor neuron (ORN), also called an olfactory sensory neuron (OSN), is a sensory neuron within the olfactory system.
Structure
Humans have between 10 and 20 million olfactory receptor neurons (ORNs). In vertebrates, ORNs are b ...
s in the
olfactory epithelium
The olfactory epithelium is a specialized epithelial tissue inside the nasal cavity that is involved in smell. In humans, it measures
9 cm2 and lies on the roof of the nasal cavity about 7 cm above and behind the nostrils. The olfactory ...
. The olfactory epithelium is made up of at least six morphologically and biochemically different cell types.
The proportion of olfactory
epithelium
Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercellul ...
compared to respiratory epithelium (not innervated, or supplied with nerves) gives an indication of the animal's olfactory sensitivity. Humans have about of olfactory epithelium, whereas some dogs have . A dog's olfactory epithelium is also considerably more densely innervated, with a hundred times more receptors per square centimeter. The sensory olfactory system integrates with other senses to form the perception of
flavor
Flavor or flavour is either the sensory perception of taste or smell, or a flavoring in food that produces such perception.
Flavor or flavour may also refer to:
Science
*Flavors (programming language), an early object-oriented extension to Lis ...
.
Often, land organisms will have separate olfaction systems for smell and taste (orthonasal smell and
retronasal smell Retronasal smell, retronasal olfaction, is the ability to perceive flavor dimensions of foods and drinks. Retronasal smell is a sensory modality that produces flavor. It is best described as a combination of traditional smell (orthonasal smell) and ...
), but water-dwelling organisms usually have only one system.
Molecules of odorants passing through the
superior nasal concha
The superior nasal concha is a small, curved plate of bone representing a medial bony process of the labyrinth of the ethmoid bone. The superior nasal concha forms the roof of the superior nasal meatus.
Anatomy
Anatomical relations
The superi ...
of the nasal passages dissolve in the
mucus
Mucus ( ) is a slippery aqueous secretion produced by, and covering, mucous membranes. It is typically produced from cells found in mucous glands, although it may also originate from mixed glands, which contain both serous and mucous cells. It is ...
that lines the superior portion of the cavity and are detected by
olfactory receptor
Olfactory receptors (ORs), also known as odorant receptors, are chemoreceptors expressed in the cell membranes of olfactory receptor neurons and are responsible for the detection of odorants (for example, compounds that have an odor) which give r ...
s on the
dendrites
Dendrites (from Greek δένδρον ''déndron'', "tree"), also dendrons, are branched protoplasmic extensions of a nerve cell that propagate the electrochemical stimulation received from other neural cells to the cell body, or soma, of the n ...
of the olfactory sensory neurons. This may occur by diffusion or by the binding of the odorant to
odorant-binding protein Odorant-binding proteins (OBPs) are small (10 to 30 kDa) soluble proteins secreted by auxiliary cells surrounding olfactory receptor neurons, including the nasal mucus of many vertebrate species and in the sensillar lymph of chemosensory sensilla of ...
s. The mucus overlying the epithelium contains
mucopolysaccharides
Glycosaminoglycans (GAGs) or mucopolysaccharides are long, linear polysaccharides consisting of repeating disaccharide units (i.e. two-sugar units). The repeating two-sugar unit consists of a uronic sugar and an amino sugar, except in the case ...
, salts,
enzymes
Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different molecule ...
, and
antibodies
An antibody (Ab), also known as an immunoglobulin (Ig), is a large, Y-shaped protein used by the immune system to identify and neutralize foreign objects such as pathogenic bacteria and viruses. The antibody recognizes a unique molecule of the ...
(these are highly important, as the olfactory neurons provide a direct passage for infection to pass to the
brain
A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a v ...
). This mucus acts as a solvent for odor molecules, flows constantly, and is replaced approximately every ten minutes.
In
insect
Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s, smells are sensed by olfactory sensory neurons in the chemosensory
sensilla, which are present in insect antenna, palps, and tarsa, but also on other parts of the insect body. Odorants penetrate into the cuticle pores of chemosensory sensilla and get in contact with insect odorant-binding proteins (OBPs) or
Chemosensory proteins (CSPs), before activating the sensory neurons.
Receptor neuron
The binding of the
ligand
In coordination chemistry, a ligand is an ion or molecule (functional group) that binds to a central metal atom to form a coordination complex. The bonding with the metal generally involves formal donation of one or more of the ligand's electr ...
(odor molecule or odorant) to the receptor leads to an
action potential
An action potential occurs when the membrane potential of a specific cell location rapidly rises and falls. This depolarization then causes adjacent locations to similarly depolarize. Action potentials occur in several types of animal cells, ...
in the receptor neuron, via a
second messenger
Second messengers are intracellular signaling molecules released by the cell in response to exposure to extracellular signaling molecules—the first messengers. (Intercellular signals, a non-local form or cell signaling, encompassing both first me ...
pathway, depending on the organism. In mammals, the odorants stimulate
adenylate cyclase
Adenylate cyclase (EC 4.6.1.1, also commonly known as adenyl cyclase and adenylyl cyclase, abbreviated AC) is an enzyme with systematic name ATP diphosphate-lyase (cyclizing; 3′,5′-cyclic-AMP-forming). It catalyzes the following reaction:
:A ...
to synthesize
cAMP
Camp may refer to:
Outdoor accommodation and recreation
* Campsite or campground, a recreational outdoor sleeping and eating site
* a temporary settlement for nomads
* Camp, a term used in New England, Northern Ontario and New Brunswick to descri ...
via a
G protein
G proteins, also known as guanine nucleotide-binding proteins, are a family of proteins that act as molecular switches inside cells, and are involved in transmitting signals from a variety of stimuli outside a cell to its interior. Their act ...
called G
olf. cAMP, which is the second messenger here, opens a
cyclic nucleotide-gated ion channel
Cycle, cycles, or cyclic may refer to:
Anthropology and social sciences
* Cyclic history, a theory of history
* Cyclical theory, a theory of American political history associated with Arthur Schlesinger, Sr.
* Social cycle, various cycles in so ...
(CNG), producing an influx of
cation
An ion () is an atom or molecule with a net electrical charge.
The charge of an electron is considered to be negative by convention and this charge is equal and opposite to the charge of a proton, which is considered to be positive by convent ...
s (largely
Ca2+ with some
Na+) into the cell, slightly depolarising it. The Ca
2+ in turn opens a Ca
2+-activated
chloride channel
Chloride channels are a superfamily of poorly understood ion channels specific for chloride. These channels may conduct many different ions, but are named for chloride because its concentration ''in vivo'' is much higher than other anions. Severa ...
, leading to efflux of
Cl−, further depolarizing the cell and triggering an action potential. Ca
2+ is then extruded through a
sodium-calcium exchanger
The sodium-calcium exchanger (often denoted Na+/Ca2+ exchanger, exchange protein, or NCX) is an antiporter membrane protein that removes calcium from cells. It uses the energy that is stored in the electrochemical gradient of sodium (Na+) by al ...
. A calcium-
calmodulin
Calmodulin (CaM) (an abbreviation for calcium-modulated protein) is a multifunctional intermediate calcium-binding messenger protein expressed in all eukaryotic cells. It is an intracellular target of the secondary messenger Ca2+, and the bind ...
complex also acts to inhibit the binding of cAMP to the cAMP-dependent channel, thus contributing to olfactory adaptation.
The main olfactory system of some mammals also contains small subpopulations of olfactory sensory neurons that detect and transduce odors somewhat differently. Olfactory sensory neurons that use trace amine-associated receptors (TAARs) to detect odors use the same second messenger signaling cascade as do the canonical olfactory sensory neurons. Other subpopulations, such as those that express the receptor guanylyl cyclase GC-D (Gucy2d) or the soluble guanylyl cyclase Gucy1b2, use a cGMP cascade to transduce their odorant ligands. These distinct subpopulations (olfactory subsystems) appear specialized for the detection of small groups of chemical stimuli.
This mechanism of transduction is somewhat unusual, in that cAMP works by directly binding to the
ion channel
Ion channels are pore-forming membrane proteins that allow ions to pass through the channel pore. Their functions include establishing a resting membrane potential, shaping action potentials and other electrical signals by gating the flow of io ...
rather than through activation of
protein kinase A
In cell biology, protein kinase A (PKA) is a family of enzymes whose activity is dependent on cellular levels of cyclic AMP (cAMP). PKA is also known as cAMP-dependent protein kinase (). PKA has several functions in the cell, including regulatio ...
. It is similar to the transduction mechanism for
photoreceptors, in which the second messenger
cGMP works by directly binding to ion channels, suggesting that maybe one of these receptors was evolutionarily adapted into the other. There are also considerable similarities in the immediate processing of stimuli by
lateral inhibition
In neurobiology, lateral inhibition is the capacity of an excited neuron to reduce the activity of its neighbors. Lateral inhibition disables the spreading of action potentials from excited neurons to neighboring neurons in the lateral direction ...
.
Averaged activity of the receptor neurons can be measured in several ways. In vertebrates, responses to an odor can be measured by an
electro-olfactogram or through calcium imaging of receptor neuron terminals in the olfactory bulb. In insects, one can perform
electroantennography Electroantennography or EAG is a technique for measuring the average output of an insect antenna to its brain for a given odor. It is commonly used in electrophysiology while studying the function of the olfactory pathway in insects. The techniqu ...
or calcium imaging within the olfactory bulb.
Olfactory bulb projections
![Early Olfactory System](https://upload.wikimedia.org/wikipedia/commons/0/07/Early_Olfactory_System.svg)
Olfactory sensory neurons project
axon
An axon (from Greek ἄξων ''áxōn'', axis), or nerve fiber (or nerve fibre: see spelling differences), is a long, slender projection of a nerve cell, or neuron, in vertebrates, that typically conducts electrical impulses known as action po ...
s to the brain within the
olfactory nerve
The olfactory nerve, also known as the first cranial nerve, cranial nerve I, or simply CN I, is a cranial nerve that contains sensory nerve fibers relating to the sense of smell.
The afferent nerve fibers of the olfactory receptor neurons tr ...
, (
cranial nerve
Cranial nerves are the nerves that emerge directly from the brain (including the brainstem), of which there are conventionally considered twelve pairs. Cranial nerves relay information between the brain and parts of the body, primarily to and ...
I). These nerve fibers, lacking
myelin
Myelin is a lipid-rich material that surrounds nerve cell axons (the nervous system's "wires") to insulate them and increase the rate at which electrical impulses (called action potentials) are passed along the axon. The myelinated axon can be ...
sheaths, pass to the
olfactory bulb
The olfactory bulb (Latin: ''bulbus olfactorius'') is a neural structure of the vertebrate forebrain involved in olfaction, the sense of smell. It sends olfactory information to be further processed in the amygdala, the orbitofrontal cortex ( ...
of the brain through perforations in the
cribriform plate
In mammalian anatomy, the cribriform plate (Latin for lit. ''sieve-shaped''), horizontal lamina or lamina cribrosa is part of the ethmoid bone. It is received into the ethmoidal notch of the frontal bone and roofs in the nasal cavities. It supp ...
, which in turn projects olfactory information to the
olfactory cortex and other areas. The axons from the
olfactory receptor
Olfactory receptors (ORs), also known as odorant receptors, are chemoreceptors expressed in the cell membranes of olfactory receptor neurons and are responsible for the detection of odorants (for example, compounds that have an odor) which give r ...
s converge in the outer layer of the olfactory bulb within small (≈50
micrometers
The micrometre (American and British English spelling differences#-re, -er, international spelling as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American and British English spelling differences# ...
in diameter) structures called
glomeruli
''Glomerulus'' () is a common term used in anatomy to describe globular structures of entwined vessels, fibers, or neurons. ''Glomerulus'' is the diminutive of the Latin ''glomus'', meaning "ball of yarn".
''Glomerulus'' may refer to:
* the filter ...
.
Mitral cell
Mitral cells are neurons that are part of the olfactory system. They are located in the olfactory bulb in the mammalian central nervous system. They receive information from the axons of olfactory receptor neurons, forming synapses in neuropils ca ...
s, located in the inner layer of the olfactory bulb, form synapses with the axons of the sensory neurons within glomeruli and send the information about the
odor
An odor (American English) or odour (English in the Commonwealth of Nations, Commonwealth English; American and British English spelling differences#-our, -or, see spelling differences) is caused by one or more volatilized chemical compounds ...
to other parts of the olfactory system, where multiple signals may be processed to form a synthesized olfactory perception. A large degree of convergence occurs, with 25,000 axons synapsing on 25 or so mitral cells, and with each of these mitral cells projecting to multiple glomeruli. Mitral cells also project to
periglomerular cell
Periglomerular cells mediate lateral inhibition in the olfactory system together with granule cells. They have inhibitory synapses on mitral cells and tufted cell Tufted cells are found within the olfactory glomeruli. They receive input from the ...
s and
granular cells that inhibit the mitral cells surrounding it (
lateral inhibition
In neurobiology, lateral inhibition is the capacity of an excited neuron to reduce the activity of its neighbors. Lateral inhibition disables the spreading of action potentials from excited neurons to neighboring neurons in the lateral direction ...
). Granular cells also mediate inhibition and excitation of mitral cells through pathways from centrifugal fibers and the anterior olfactory nuclei. Neuromodulators like
acetylcholine
Acetylcholine (ACh) is an organic chemical that functions in the brain and body of many types of animals (including humans) as a neurotransmitter. Its name is derived from its chemical structure: it is an ester of acetic acid and choline. Part ...
,
serotonin
Serotonin () or 5-hydroxytryptamine (5-HT) is a monoamine neurotransmitter. Its biological function is complex and multifaceted, modulating mood, cognition, reward, learning, memory, and numerous physiological processes such as vomiting and vas ...
and
norepinephrine
Norepinephrine (NE), also called noradrenaline (NA) or noradrenalin, is an organic chemical in the catecholamine family that functions in the brain and body as both a hormone and neurotransmitter. The name "noradrenaline" (from Latin '' ad'', ...
all send axons to the olfactory bulb and have been implicated in gain modulation, pattern separation, and
memory
Memory is the faculty of the mind by which data or information is encoded, stored, and retrieved when needed. It is the retention of information over time for the purpose of influencing future action. If past events could not be remembered, ...
functions, respectively.
The mitral cells leave the olfactory bulb in the
lateral olfactory tract, which synapses on five major regions of the cerebrum: the
anterior olfactory nucleus
The anterior olfactory nucleus (AON; also called the anterior olfactory cortex) is a portion of the forebrain of vertebrates.
It is involved in olfaction
and has supposedly strong influence on other olfactory areas like the olfactory bulb and the ...
, the
olfactory tubercle
The olfactory tubercle (OT), also known as the tuberculum olfactorium, is a multi-sensory processing center that is contained within the olfactory cortex and ventral striatum and plays a role in reward cognition. The OT has also been shown to ...
, the
amygdala
The amygdala (; plural: amygdalae or amygdalas; also '; Latin from Greek, , ', 'almond', 'tonsil') is one of two almond-shaped clusters of nuclei located deep and medially within the temporal lobes of the brain's cerebrum in complex verteb ...
, the
piriform cortex
The piriform cortex, or pyriform cortex, is a region in the brain, part of the rhinencephalon situated in the cerebrum. The function of the piriform cortex relates to the sense of smell.
Structure
The piriform cortex is part of the rhinencepha ...
, and the
entorhinal cortex
The entorhinal cortex (EC) is an area of the brain's allocortex, located in the medial temporal lobe, whose functions include being a widespread network hub for memory, navigation, and the perception of time.Integrating time from experience in the ...
. The anterior olfactory nucleus projects, via the
anterior commissure
The anterior commissure (also known as the precommissure) is a white matter tract (a bundle of axons) connecting the two temporal lobes of the cerebral hemispheres across the midline, and placed in front of the columns of the fornix. In most exis ...
, to the contralateral olfactory bulb, inhibiting it. The piriform cortex has two major divisions with anatomically distinct organizations and functions. The anterior piriform cortex (APC) appears to be better at determining the chemical structure of the odorant molecules, and the posterior piriform cortex (PPC) has a strong role in categorizing odors and assessing similarities between odors (e.g. minty, woody, and citrus are odors that can, despite being highly variant chemicals, be distinguished via the PPC in a concentration-independent manner). The piriform cortex projects to the
medial dorsal nucleus
The medial dorsal nucleus (or dorsomedial nucleus of thalamus) is a large nucleus in the thalamus.
It is believed to play a role in memory.
Structure
It relays inputs from the amygdala and olfactory cortex and projects to the prefrontal cortex a ...
of the thalamus, which then projects to the orbitofrontal cortex. The orbitofrontal cortex mediates conscious perception of the odor. The three-layered piriform cortex projects to a number of thalamic and hypothalamic nuclei, the
hippocampus
The hippocampus (via Latin from Greek , 'seahorse') is a major component of the brain of humans and other vertebrates. Humans and other mammals have two hippocampi, one in each side of the brain. The hippocampus is part of the limbic system, a ...
and amygdala and the orbitofrontal cortex, but its function is largely unknown. The entorhinal cortex projects to the amygdala and is involved in emotional and autonomic responses to odor. It also projects to the hippocampus and is involved in motivation and memory. Odor information is stored in
long-term memory
Long-term memory (LTM) is the stage of the Atkinson–Shiffrin memory model in which informative knowledge is held indefinitely. It is defined in contrast to short-term and working memory, which persist for only about 18 to 30 seconds. Long-t ...
and has strong connections to
emotional memory
Emotion can have a powerful effect on humans and animals. Numerous studies have shown that the most vivid autobiographical memories tend to be of emotional events, which are likely to be recalled more often and with more clarity and detail than ...
. This is possibly due to the olfactory system's close anatomical ties to the
limbic system
The limbic system, also known as the paleomammalian cortex, is a set of brain structures located on both sides of the thalamus, immediately beneath the medial temporal lobe of the cerebrum primarily in the forebrain.Schacter, Daniel L. 2012. ''Ps ...
and hippocampus, areas of the brain that have long been known to be involved in emotion and place memory, respectively.
Since any one receptor is responsive to various odorants, and there is a great deal of convergence at the level of the olfactory bulb, it may seem strange that human beings are able to distinguish so many different odors. It seems that a highly complex form of processing must be occurring; however, as it can be shown that, while many neurons in the olfactory bulb (and even the pyriform cortex and amygdala) are responsive to many different odors, half the neurons in the orbitofrontal cortex are responsive to only one odor, and the rest to only a few. It has been shown through microelectrode studies that each individual odor gives a particular spatial map of excitation in the olfactory bulb. It is possible that the brain is able to distinguish specific odors through spatial encoding, but temporal coding must also be taken into account. Over time, the spatial maps change, even for one particular odor, and the brain must be able to process these details as well.
Inputs from the two
nostril
A nostril (or naris , plural ''nares'' ) is either of the two orifices of the nose. They enable the entry and exit of air and other gasses through the nasal cavities. In birds and mammals, they contain branched bones or cartilages called turbi ...
s have separate inputs to the brain, with the result that, when each nostril takes up a different odorant, a person may experience perceptual rivalry in the olfactory sense akin to that of
binocular rivalry Binocular rivalry is a phenomenon of visual perception in which perception alternates between different images presented to each eye.
When one image is presented to one eye and a very different image is presented to the other (also known as dic ...
.
In
insect
Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s, smells are sensed by sensilla located on the antenna and maxillary palp and first processed by the
antennal lobe The antennal lobe is the primary (first order) olfactory brain area in insects. The antennal lobe is a sphere-shaped deutocerebral neuropil in the brain that receives input from the olfactory sensory neurons in the antennae and mouthparts. Function ...
(analogous to the
olfactory bulb
The olfactory bulb (Latin: ''bulbus olfactorius'') is a neural structure of the vertebrate forebrain involved in olfaction, the sense of smell. It sends olfactory information to be further processed in the amygdala, the orbitofrontal cortex ( ...
), and next by the
mushroom bodies
The mushroom bodies or ''corpora pedunculata'' are a pair of structures in the brain of insects, other arthropods, and some annelids (notably the ragworm ''Platynereis dumerilii''). They are known to play a role in olfactory learning and memory ...
and
lateral horn.
Coding and perception
The process by which olfactory information is coded in the brain to allow for proper perception is still being researched, and is not completely understood. When an odorant is detected by receptors, they in a sense break the odorant down, and then the brain puts the odorant back together for identification and perception. The odorant binds to receptors that recognize only a specific functional group, or feature, of the odorant, which is why the chemical nature of the odorant is important.
After binding the odorant, the receptor is activated and will send a signal to the glomeruli
in the
olfactory bulb
The olfactory bulb (Latin: ''bulbus olfactorius'') is a neural structure of the vertebrate forebrain involved in olfaction, the sense of smell. It sends olfactory information to be further processed in the amygdala, the orbitofrontal cortex ( ...
. Each glomerulus receives signals from multiple receptors that detect similar odorant features. Because several receptor types are activated due to the different chemical features of the odorant, several glomeruli are activated as well. The signals from the glomeruli are transformed to a pattern of oscillations of neural activities of the
mitral cells, the output neurons from the olfactory bulb. Olfactory bulb sends this pattern to the
olfactory cortex. Olfactory cortex is thought to have associative memories, so that it resonates to this bulbar pattern when the odor object is recognized.
The cortex sends centrifugal feedback to the bulb. This feedback could suppress bulbar responses to the recognized odor objects, causing olfactory adaptation to background odors, so that the newly arrived foreground odor objects could be singled out for better recognition.
During odor search, feedback could also be used to enhance odor detection.
The distributed code allows the brain to detect specific odors in mixtures of many background odors.
It is a general idea that the layout of brain structures corresponds to physical features of stimuli (called topographic coding), and similar analogies have been made in smell with concepts such as a layout corresponding to chemical features (called chemotopy) or perceptual features.
While chemotopy remains a highly controversial concept, evidence exists for perceptual information implemented in the spatial dimensions of olfactory networks.
Accessory olfactory system
Many animals, including most mammals and reptiles, but not humans, have two distinct and segregated olfactory systems: a main olfactory system, which detects volatile stimuli, and an accessory olfactory system, which detects fluid-phase stimuli. Behavioral evidence suggests that these fluid-phase stimuli often function as
pheromone
A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s, although pheromones can also be detected by the main olfactory system. In the accessory olfactory system, stimuli are detected by the
vomeronasal organ
The vomeronasal organ (VNO), or Jacobson's organ, is the paired auxiliary olfactory (smell) sense organ located in the soft tissue of the nasal septum, in the nasal cavity just above the roof of the mouth (the hard palate) in various tetrapods. T ...
, located in the vomer, between the
nose
A nose is a protuberance in vertebrates that houses the nostrils, or nares, which receive and expel air for respiration alongside the mouth. Behind the nose are the olfactory mucosa and the sinuses. Behind the nasal cavity, air next passes th ...
and the
mouth
In animal anatomy, the mouth, also known as the oral cavity, or in Latin cavum oris, is the opening through which many animals take in food and issue vocal sounds. It is also the cavity lying at the upper end of the alimentary canal, bounded on ...
. Snakes use it to smell prey, sticking their tongue out and touching it to the organ. Some mammals make a facial expression called
flehmen to direct stimuli to this organ.
The sensory receptors of the accessory olfactory system are located in the vomeronasal organ. As in the main olfactory system, the axons of these sensory neurons project from the vomeronasal organ to the
accessory olfactory bulb, which in the mouse is located on the dorsal-posterior portion of the main
olfactory bulb
The olfactory bulb (Latin: ''bulbus olfactorius'') is a neural structure of the vertebrate forebrain involved in olfaction, the sense of smell. It sends olfactory information to be further processed in the amygdala, the orbitofrontal cortex ( ...
. Unlike in the main olfactory system, the axons that leave the accessory olfactory bulb do not project to the brain's cortex but rather to targets in the
amygdala
The amygdala (; plural: amygdalae or amygdalas; also '; Latin from Greek, , ', 'almond', 'tonsil') is one of two almond-shaped clusters of nuclei located deep and medially within the temporal lobes of the brain's cerebrum in complex verteb ...
and
bed nucleus of the stria terminalis
The stria terminalis (or terminal stria) is a structure in the brain consisting of a band of fibers running along the lateral margin of the ventricular surface of the thalamus. Serving as a major output pathway of the amygdala, the stria termina ...
, and from there to the
hypothalamus
The hypothalamus () is a part of the brain that contains a number of small nuclei with a variety of functions. One of the most important functions is to link the nervous system to the endocrine system via the pituitary gland. The hypothalamu ...
, where they may influence aggression and mating behavior.
In insects
Insect olfaction refers to the function of
chemical receptor
A chemoreceptor, also known as chemosensor, is a specialized sensory receptor which transduces a chemical substance (endogenous or induced) to generate a biological signal. This signal may be in the form of an action potential, if the chemorecept ...
s that enable
insects
Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs of j ...
to detect and identify
volatile compounds for
foraging
Foraging is searching for wild food resources. It affects an animal's Fitness (biology), fitness because it plays an important role in an animal's ability to survive and reproduce. Optimal foraging theory, Foraging theory is a branch of behaviora ...
, predator avoidance, finding
mating
In biology, mating is the pairing of either opposite-sex or hermaphroditic organisms for the purposes of sexual reproduction. ''Fertilization'' is the fusion of two gametes. ''Copulation'' is the union of the sex organs of two sexually reproduc ...
partners (via
pheromone
A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s) and locating
oviposition
The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
habitats.
Thus, it is the most important sensation for insects.
Most important insect behaviors must be timed perfectly which is dependent on what they smell and when they smell it.
For example, smell is essential for hunting in many species of
wasps
A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. ...
, including ''
Polybia sericea''.
The two organs insects primarily use for detecting odors are the
antennae and specialized mouth parts called the maxillary palps.
However, a recent study has demonstrated the olfactory role of ovipositor in fig wasps. Inside of these olfactory organs there are neurons called olfactory receptor neurons which, as the name implies, house receptors for scent molecules in their cell membranes. The majority of
olfactory receptor neuron
An olfactory receptor neuron (ORN), also called an olfactory sensory neuron (OSN), is a sensory neuron within the olfactory system.
Structure
Humans have between 10 and 20 million olfactory receptor neurons (ORNs). In vertebrates, ORNs are b ...
s typically reside in the
antenna
Antenna ( antennas or antennae) may refer to:
Science and engineering
* Antenna (radio), also known as an aerial, a transducer designed to transmit or receive electromagnetic (e.g., TV or radio) waves
* Antennae Galaxies, the name of two collid ...
. These neurons can be very abundant, for example ''
Drosophila
''Drosophila'' () is a genus of flies, belonging to the family Drosophilidae, whose members are often called "small fruit flies" or (less frequently) pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many species ...
'' flies have 2,600 olfactory sensory neurons.
Insects are capable of smelling and differentiating between thousands of
volatile compounds both
sensitively and selectively.
Sensitivity is how attuned the insect is to very small amounts of an odorant or small changes in the concentration of an odorant. Selectivity refers to the insects' ability to tell one odorant apart from another. These compounds are commonly broken into three classes: short chain
carboxylic acid
In organic chemistry, a carboxylic acid is an organic acid that contains a carboxyl group () attached to an R-group. The general formula of a carboxylic acid is or , with R referring to the alkyl, alkenyl, aryl, or other group. Carboxylic ...
s,
aldehyde
In organic chemistry, an aldehyde () is an organic compound containing a functional group with the structure . The functional group itself (without the "R" side chain) can be referred to as an aldehyde but can also be classified as a formyl grou ...
s and low molecular weight nitrogenous compounds.
Some insects, such as the moth ''
Deilephila elpenor
Deilephila elpenor, the elephant hawk moth or large elephant hawk moth, is a moth in the family Sphingidae. Its common name is derived from the caterpillar's resemblance to an elephant's trunk. It is most common in central Europe and is distribut ...
'', use smell as a means to find food sources.
In plants
The tendrils of plants are especially sensitive to airborne
volatile organic compound
Volatile organic compounds (VOCs) are organic compounds that have a high vapour pressure at room temperature. High vapor pressure correlates with a low boiling point, which relates to the number of the sample's molecules in the surrounding air, a ...
s. Parasites such as
dodder
''Cuscuta'' (), commonly known as dodder or amarbel, is a genus of over 201 species of yellow, orange, or red (rarely green) parasitic plants. Formerly treated as the only genus in the family Cuscutaceae, it now is accepted as belonging in the ...
make use of this in locating their preferred hosts and locking on to them. The emission of volatile compounds is detected when foliage is browsed by animals. Threatened plants are then able to take defensive chemical measures, such as moving
tannin
Tannins (or tannoids) are a class of astringent, polyphenolic biomolecules that bind to and precipitate proteins and various other organic compounds including amino acids and alkaloids.
The term ''tannin'' (from Anglo-Norman ''tanner'', ...
compounds to their foliage.
Machine-based smelling
Scientists have devised methods for quantifying the intensity of odors, in particular for the purpose of analyzing unpleasant or objectionable odors released by an industrial source into a community. Since the 1800s industrial countries have encountered incidents where proximity of an industrial source or landfill produced adverse reactions among nearby residents regarding airborne odor. The basic theory of odor analysis is to measure what extent of dilution with "pure" air is required before the sample in question is rendered indistinguishable from the "pure" or reference standard. Since each person perceives odor differently, an "odor panel" composed of several different people is assembled, each
sniffing the same sample of diluted specimen air. A field
olfactometer
An olfactometer is an instrument used to detect and measure odor dilution. Olfactometers are used in conjunction with human subjects in laboratory settings, most often in market research, to quantify and qualify human olfaction. Olfactometers are ...
can be utilized to determine the magnitude of an odor.
Many air management districts in the
US have numerical standards of acceptability for the intensity of odor that is allowed to cross into a residential property. For example, the
Bay Area Air Quality Management District
The Bay Area Air Quality Management District (BAAQMD) is a public agency that regulates the stationary sources of air pollution in the nine counties of California's San Francisco Bay Area: Alameda, Contra Costa, Marin, Napa, San Francisco, Sa ...
has applied its standard in regulating numerous industries, landfills, and sewage treatment plants. Example applications this district has engaged are the
San Mateo, California
San Mateo ( ; ) is a city in San Mateo County, California, on the San Francisco Peninsula. About 20 miles (32 km) south of San Francisco, the city borders Burlingame to the north, Hillsborough to the west, San Francisco Bay and Foster C ...
, wastewater treatment plant; the
Shoreline Amphitheatre
Shoreline Amphitheatre is an outdoor amphitheater located in Mountain View, California, in the San Francisco Bay Area. The venue has a capacity of 22,500, with 6,500 reserved seats and 16,000 general admission on the lawn. When the parking lot ...
in
Mountain View, California
Mountain View is a city in Santa Clara County, California, United States. Named for its views of the Santa Cruz Mountains, it has a population of 82,376.
Mountain View was integral to the early history and growth of Silicon Valley, and is the ...
; and the
IT Corporation
IT Corporation was a United States industrial company whose principal business was the disposal of industrial hazardous waste. At least as early as the 1970s the company was one of the largest market share holders of the liquid hazardous waste d ...
waste ponds,
Martinez, California
Martinez (Spanish language, Spanish: ''Martínez'') is a city and the county seat of Contra Costa County, California, United States, in the East Bay region of the San Francisco Bay Area. Located on the southern shore of the Carquinez Strait, t ...
.
Classification
Systems of classifying odors include:
* Crocker-Henderson system, which rates smells on a 0-8 scale for each of four "primary" smells: fragrant, acid, burnt, and
caprylic.
* Henning's prism
* Zwaardemaker smell system (invented by
Hendrik Zwaardemaker)
Disorders
Specific terms are used to describe disorders associated with smelling:
*
Anosmia
Anosmia, also known as smell blindness, is the loss of the ability to detect one or more smells. Anosmia may be temporary or permanent. It differs from hyposmia, which is a decreased sensitivity to some or all smells.
Anosmia can be due to a nu ...
– inability to smell
*
Hyperosmia
Hyperosmia is an increased olfactory acuity (heightened sense of smell), usually caused by a lower threshold for odor. This perceptual disorder arises when there is an abnormally increased signal at any point between the olfactory receptors and th ...
– an abnormally acute sense of smell
*
Hyposmia
Hyposmia, or microsmia, is a reduced ability to smell and to detect odors. A related condition is anosmia, in which no odors can be detected. Some of the causes of olfaction problems are allergies, nasal polyps, viral infections and head trauma. ...
– decreased ability to smell
*
Presbyosmia – the natural decline in the sense of smell in old age
*
Dysosmia
Dysosmia is a disorder described as any qualitative alteration or distortion of the perception of smell. Qualitative alterations differ from quantitative alterations, which include anosmia and hyposmia. Dysosmia can be classified as either paros ...
– distortion in the sense of smell
**
Parosmia
Parosmia (from the Greek παρά ''pará'' and ὀσμή ''osmḗ'' "smell") is a dysfunctional smell detection characterized by the inability of the brain to correctly identify an odor's "natural" smell. Instead, the natural odor is usually tr ...
– distortion in the perception of an odor
**
Phantosmia
Phantosmia (phantom smell), also called an olfactory hallucination or a phantom odor, is smelling an odor that is not actually there. It can occur in one nostril or both. Unpleasant phantosmia, cacosmia, is more common and is often described as s ...
– distortion in the absence of an odor, "hallucinated smell"
*
Heterosmia – inability to distinguish odors
*
Olfactory reference syndrome
Olfactory reference syndrome (ORS) is a psychiatric condition in which there is a persistent false belief and preoccupation with the idea of emitting abnormal body odors which the patient thinks are foul and offensive to other individuals. People ...
– psychological disorder that causes the patient to imagine he or she has strong
body odor
Body odor or body odour (BO) is present in all animals and its intensity can be influenced by many factors (behavioral patterns, survival strategies). Body odor has a strong genetic basis, but can also be strongly influenced by various diseases ...
*
Osmophobia
Osmophobia or olfactophobia refers to a fear, aversion, or psychological hypersensitivity to odors. The phobia generally occurs in chronic migraine sufferers who may have ''odor triggered'' migraines. Such migraines are most frequently triggered ...
– aversion or psychological hypersensitivity to odors
Viruses can also infect the olfactory epithelium leading to a loss of the sense of olfaction. About 50% of patients with
SARS-CoV-2
Severe acute respiratory syndrome coronavirus 2 (SARS‑CoV‑2) is a strain of coronavirus that causes COVID-19 (coronavirus disease 2019), the respiratory illness responsible for the ongoing COVID-19 pandemic. The virus previously had a ...
(causing COVID-19) experience some type of
disorder associated with their sense of smell, including anosmia and parosmia.
SARS-CoV-1
Severe acute respiratory syndrome coronavirus 1 (SARS-CoV-1; or Severe acute respiratory syndrome coronavirus, SARS-CoV) is a strain of coronavirus that causes severe acute respiratory syndrome (SARS), the respiratory illness responsible for ...
,
MERS-CoV
''Middle East respiratory syndrome–related coronavirus'' (''MERS-CoV''), or EMC/2012 ( HCoV-EMC/2012), is the virus that causes Middle East respiratory syndrome (MERS). It is a species of coronavirus which infects humans, bats, and camels. Th ...
and even the flu (
influenza virus
''Orthomyxoviridae'' (from Greek ὀρθός, ''orthós'' 'straight' + μύξα, ''mýxa'' 'mucus') is a family of negative-sense RNA viruses. It includes seven genera: ''Alphainfluenzavirus'', ''Betainfluenzavirus'', '' Gammainfluenzavirus'', ' ...
) can also disrupt
olfaction.
See also
*
Electronic nose
An electronic nose is an electronic sensing device intended to detect odors or flavors. The expression "electronic sensing" refers to the capability of reproducing human senses using sensor arrays and pattern recognition systems.
Since 1982, res ...
*
Evolution of olfaction
*
Nasal administration
Nasal administration, popularly known as snorting, is a route of administration in which drugs are insufflated through the nose. It can be a form of either topical administration or systemic administration, as the drugs thus locally delivered c ...
olfactory transfer
*
Olfactic Communication
*
Olfactory ensheathing cell
Olfactory ensheathing cells (OECs), also known as olfactory ensheathing glia or olfactory ensheathing glial cells, are a type of macroglia (radial glia) found in the nervous system. They are also known as olfactory Schwann cells, because they e ...
*
Olfactory fatigue
Olfactory fatigue, also known as odor fatigue, olfactory adaptation, and noseblindness, is the temporary, normal inability to distinguish a particular odor after a prolonged exposure to that airborne compound. For example, when entering a restaur ...
*
''Perfume'' (novel)
*
Scent transfer unit
References
External links
Olfaction at cf.ac.ukOlfactory Systems Laboratory at Boston UniversitySmells Database ''Neuroscience Online'' (electronic neuroscience textbook by UT Houston Medical School)
Digital Olfaction Society
{{Authority control
Limbic system