Olfaction

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Olfaction
Luna White Pyriness dog smells red flowers 03.jpg
Image of a dog smelling a poppy. Olfaction uses chemoreceptors that create signals processed in the brain that form the sense of smell.
Details
System Olfactory system
Functionsense chemicals in the environment that are used to form the sense of smell
Identifiers
MeSH D012903
Anatomical terminology

Olfaction, or the sense of smell, [1] is the special sense through which smells (or odors) are perceived. [2] It occurs when an odor binds to a receptor within the nasal cavity, transmitting a signal through the olfactory system. [3] Olfaction has many functions, including detecting hazards, and pheromones, and plays a role in taste.

Contents

Glomeruli aggregate signals from these receptors and transmit them to the olfactory bulb, where the sensory input will start to interact with parts of the brain responsible for smell identification, memory, and emotion. [4]

There are many different causes for alteration, lack, or disturbance to normal olfaction, and can include damage to the peripheral nose or smell receptors, or central problems affecting the brain. Some causes include upper respiratory infections, traumatic brain injury, and neurodegenerative disease. [5] [6]

Vertebrates

Physiology

Main olfactory system

In humans and other vertebrates, smells are sensed by olfactory sensory neurons in the olfactory epithelium. The olfactory epithelium is made up of at least six morphologically and biochemically different cell types. [7] The proportion of olfactory epithelium compared to respiratory epithelium (not innervated, or supplied with nerves) gives an indication of the animal's olfactory sensitivity. Humans have about 10 cm2 (1.6 sq in) of olfactory epithelium, whereas some dogs have 170 cm2 (26 sq in). A dog's olfactory epithelium is also considerably more densely innervated, with a hundred times more receptors per square centimeter. [8] The sensory olfactory system integrates with other senses to form the perception of flavor. [9] Often, land organisms will have separate olfaction systems for smell and taste (orthonasal smell and retronasal smell), but water-dwelling organisms usually have only one system. [10]

Molecules of odorants passing through the superior nasal concha of the nasal passages dissolve in the mucus that lines the superior portion of the cavity and are detected by olfactory receptors on the dendrites of the olfactory sensory neurons. This may occur by diffusion or by the binding of the odorant to odorant-binding proteins. The mucus overlying the epithelium contains mucopolysaccharides, salts, enzymes, and antibodies (these are highly important, as the olfactory neurons provide a direct passage for infection to pass to the brain). This mucus acts as a solvent for odor molecules, flows constantly, and is replaced approximately every ten minutes.

In insects, 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 (odor molecule or odorant) to the receptor leads to an action potential in the receptor neuron, via a second messenger pathway, depending on the organism. In mammals, the odorants stimulate adenylate cyclase to synthesize cAMP via a G protein called Golf. cAMP, which is the second messenger here, opens a cyclic nucleotide-gated ion channel (CNG), producing an influx of cations (largely Ca 2+ with some Na +) into the cell, slightly depolarising it. The Ca2+ in turn opens a Ca2+-activated chloride channel, leading to efflux of Cl −, further depolarizing the cell and triggering an action potential. Ca2+ is then extruded through a sodium-calcium exchanger. A calcium-calmodulin 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. [11] Other subpopulations, such as those that express the receptor guanylyl cyclase GC-D (Gucy2d) [12] or the soluble guanylyl cyclase Gucy1b2, [13] use a cGMP cascade to transduce their odorant ligands. [14] [15] [16] 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 rather than through activation of protein kinase A. 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.

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 or calcium imaging within the olfactory bulb.

Olfactory bulb projections

Schematic of the early olfactory system including the olfactory epithelium and bulb. Each ORN expresses one OR that responds to different odorants. Odorant molecules bind to ORs on cilia. ORs activate ORNs that transduce the input signal into action potentials. In general, glomeruli receive input from ORs of one specific type and connect to the principal neurons of the OB, mitral and tufted cells (MT cells). Early Olfactory System.svg
Schematic of the early olfactory system including the olfactory epithelium and bulb. Each ORN expresses one OR that responds to different odorants. Odorant molecules bind to ORs on cilia. ORs activate ORNs that transduce the input signal into action potentials. In general, glomeruli receive input from ORs of one specific type and connect to the principal neurons of the OB, mitral and tufted cells (MT cells).

Olfactory sensory neurons project axons to the brain within the olfactory nerve, (cranial nerve  I). These nerve fibers, lacking myelin sheaths, pass to the olfactory bulb of the brain through perforations in the cribriform plate, which in turn projects olfactory information to the olfactory cortex and other areas. [17] The axons from the olfactory receptors converge in the outer layer of the olfactory bulb within small (≈50 micrometers in diameter) structures called glomeruli. Mitral cells, 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 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 cells and granular cells that inhibit the mitral cells surrounding it (lateral inhibition). Granular cells also mediate inhibition and excitation of mitral cells through pathways from centrifugal fibers and the anterior olfactory nuclei. Neuromodulators like acetylcholine, serotonin and norepinephrine all send axons to the olfactory bulb and have been implicated in gain modulation, [18] pattern separation, [19] and memory functions, [20] 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 olfactory tubercle, the amygdala, the piriform cortex, and the entorhinal cortex. The anterior olfactory nucleus projects, via the anterior commissure, 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). [21] The piriform cortex projects to the medial dorsal nucleus of the thalamus, which then projects to the orbitofrontal cortex. The orbitofrontal cortex mediates conscious perception of the odor.[ citation needed ] The three-layered piriform cortex projects to a number of thalamic and hypothalamic nuclei, the hippocampus 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 and has strong connections to emotional memory. This is possibly due to the olfactory system's close anatomical ties to the limbic system 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 nostrils 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. [22]

In insects, smells are sensed by sensilla located on the antenna and maxillary palp and first processed by the antennal lobe (analogous to the olfactory bulb), and next by the mushroom bodies 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. [23] 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. [24]

After binding the odorant, the receptor is activated and will send a signal to the glomeruli. [24] 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. All of the signals from the glomeruli are then sent to the brain, where the combination of glomeruli activation encodes the different chemical features of the odorant. The brain then essentially puts the pieces of the activation pattern back together in order to identify and perceive the odorant. [24] This distributed code allows the brain to detect specific odors in mixtures of many background odors. [25]

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 olfaction with concepts such as a layout corresponding to chemical features (called chemotopy) or perceptual features. [26] While chemotopy remains a highly controversial concept, [27] evidence exists for perceptual information implemented in the spatial dimensions of olfactory networks. [26]

Although conventional wisdom and lay literature, based on impressionistic findings in the 1920s, have long presented human olfaction 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. [28] 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 80 million scents. [29] 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." [30] 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". [31] [32] The logic of his paper has in turn been criticized by the authors of the original paper. [33]

Accessory olfactory system

Many animals, including most mammals and reptiles, but not humans, [34] 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 pheromones, although pheromones can also be detected by the main olfactory system. In the accessory olfactory system, stimuli are detected by the vomeronasal organ, located in the vomer, between the nose and the mouth. 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. 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 and bed nucleus of the stria terminalis, and from there to the hypothalamus, where they may influence aggression and mating behavior.

Genetics

Different people smell different odors, and most of these differences are caused by genetic differences. [35] Although odorant receptor 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 and its genetic variants (alleles) are responsible for our ability (or failure) to smell β-ionone, a key aroma in foods and beverages. [36] Similarly, the odorant receptor OR2J3 is associated with the ability to detect the "grassy" odor, cis-3-hexen-1-ol. [37] The preference (or dislike) of cilantro (coriander) has been linked to the olfactory receptor OR6A2. [38]

Functions

Taste

Flavor perception is an aggregation of auditory, taste, haptic, and smell sensory information. [9] Retronasal smell plays the biggest role in the sensation of flavor. During the process of mastication, the tongue manipulates food to release odorants. These odorants enter the nasal cavity during exhalation. [39] The olfaction of food has the sensation of being in the mouth because of co-activation of the motor cortex and olfactory epithelium during mastication. [9]

Olfaction, taste, and trigeminal receptors (also called chemesthesis) together contribute to flavor. The human tongue 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 that the olfaction contribution to flavor occurs, in contrast to that of proper smell, which occurs during the inhalation phase of breathing. [39] The olfactory system is the only human sense that bypasses the thalamus and connects directly to the forebrain. [7]

Hearing

Olfaction and sound information has been shown to converge in the olfactory tubercles of rodents. [40] This neural convergence is proposed to give rise to a perception termed smound. [41] Whereas a flavor 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; 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. [42] [43]

Humans can detect blood relatives from olfaction. [44] 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. [45] Functional imaging shows that this olfactory kinship detection process involves the frontal-temporal junction, the insula, and the dorsomedial prefrontal cortex, but not the primary or secondary olfactory cortices, or the related piriform cortex or orbitofrontal cortex. [46]

Since inbreeding 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 and inbreeding avoidance. Thus, there are fewer matings between mice sharing MUP haplotypes than would be expected if there were random mating. [47]

Variability amongst vertebrates

The importance and sensitivity of smell varies among different organisms; most mammals have a good sense of smell, whereas most birds do not, except the tubenoses (e.g., petrels and albatrosses), certain species of vultures, and the kiwis. Although, recent analysis of the chemical composition of volatile organic compounds (VOCs) from king penguin feathers suggest that VOCs may provide olfactory cues, used by the penguins to locate their colony and recognise individuals. [48] Among mammals, it is well developed in the carnivores and ungulates, which must always be aware of each other, and in those that smell for their food, such as moles. 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. [49] 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.

Scenthounds as a group can smell one- to ten-million times more acutely than a human, and bloodhounds, which have the keenest sense of smell of any dogs, [50] 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 a few days old. The second-most-sensitive nose is possessed by the Basset Hound, which was bred to track and hunt rabbits and other small animals.

Grizzly bears 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 primates, and nonexistent in cetaceans, which compensate with a well-developed sense of taste. [50] In some strepsirrhines, such as the red-bellied lemur, scent glands occur atop the head. In many species, olfaction is highly tuned to pheromones; a male silkworm moth, for example, can sense a single molecule of bombykol.

Fish, too, have a well-developed sense of smell, even though they inhabit an aquatic environment.[ citation needed ] 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.

Insects

Insect olfaction refers to the function of chemical receptors that enable insects to detect and identify volatile compounds for foraging, predator avoidance, finding mating partners (via pheromones) and locating oviposition habitats. [51] Thus, it is the most important sensation for insects. [51] Most important insect behaviors must be timed perfectly which is dependent on what they smell and when they smell it. [52] For example, olfaction is essential for hunting in many species of wasps, including Polybia sericea .

The two organs insects primarily use for detecting odors are the antennae and specialized mouth parts called the maxillary palps. [53] However, a recent study has demonstrated the olfactory role of ovipositor in fig wasps. [54] 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 membrane. The majority of olfactory receptor neurons typically reside in the antenna. These neurons can be very abundant, for example Drosophila flies have 2,600 olfactory sensory neurons. [53]

Insects are capable of smelling and differentiating between thousands of volatile compounds both sensitively and selectively. [51] [55] 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 acids, aldehydes and low molecular weight nitrogenous compounds. [55] Some insects, such as the moth Deilephila elpenor , use olfaction as a means to find food sources.

Plants

The tendrils of plants are especially sensitive to airborne volatile organic compounds. Parasites such as dodder make use of this in locating their preferred hosts and locking on to them. [56] 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 compounds to their foliage.

Machines

Nasal Ranger, an olfactometer, in use Nasal Ranger.JPG
Nasal Ranger, an olfactometer, in use

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 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 has applied its standard in regulating numerous industries, landfills, and sewage treatment plants. Example applications this district has engaged are the San Mateo, California, wastewater treatment plant; the Shoreline Amphitheatre in Mountain View, California; and the IT Corporation waste ponds, Martinez, California.

Classification

Systems of classifying odors include:

Disorders

Specific terms are used to describe disorders associated with smelling:

Viruses can also infect the olfactory epithelium leading to a loss of the sense of olfaction. About 50% of patients with SARS-CoV-2 (causing COVID-19) experience some type of disorder associated with their sense of smell, including anosmia and parosmia. SARS-CoV-1, MERS-CoV and even the flu (influenza virus) can also disrupt olfaction. [60]

History

The Lady and the Unicorn, a Flemish tapestry depicting the sense of smell, 1484-1500. Musee national du Moyen Age, Paris. The Lady and the unicorn Smell.jpg
The Lady and the Unicorn , a Flemish tapestry depicting the sense of smell, 1484–1500. Musée national du Moyen Âge , Paris.

Early scientific study of olfaction includes the extensive doctoral dissertation of Eleanor Gamble, published in 1898, which compared olfactory to other stimulus modalities, and implied that smell had a lower intensity discrimination. [61]

As the Epicurean and atomistic Roman philosopher Lucretius (1st century BCE) speculated, different odors are attributed to different shapes and sizes of "atoms" (odor molecules in the modern understanding) that stimulate the olfactory organ. [62]

A modern demonstration of that theory was the cloning of olfactory receptor proteins by Linda B. Buck and Richard Axel (who were awarded the Nobel Prize in 2004), and subsequent pairing of odor molecules to specific receptor proteins. [63] Each odor receptor molecule recognizes only a particular molecular feature or class of odor molecules. Mammals have about a thousand genes that code for odor reception. [64] 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. [65] In mammals, each olfactory receptor neuron expresses only one functional odor receptor. [66] 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. 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). [67]

According to a new study, researchers have found that a functional relationship exists between molecular volume of odorants and the olfactory neural response. [68] An alternative theory, the vibration theory proposed by Luca Turin, [69] [70] posits that odor receptors detect the frequencies of vibrations of odor molecules in the infrared range by quantum tunnelling. However, the behavioral predictions of this theory have been called into question. [71] There is no theory yet that explains olfactory perception completely.

The state of the field as of 2020—its history and current laboratory routines and practice—has been surveyed and studied in a new book Smellosophy: What the Nose tells the Mind, [72] authored by historian, philosopher, and cognitive scientist Ann-Sophie Barwich.

See also

Related Research Articles

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 name is derived from the fact that it lies adjacent to the unpaired vomer bone in the nasal septum. It is present and functional in all snakes and lizards, and in many mammals, including cats, dogs, cattle, pigs, and some primates; in humans and horses it is present, but is vestigial and non-functional.

Sensory nervous system Part of the nervous system responsible for processing sensory information

The sensory nervous system is a part of the nervous system responsible for processing sensory information. A sensory system consists of sensory neurons, neural pathways, and parts of the brain involved in sensory perception. Commonly recognized sensory systems are those for vision, hearing, touch, taste, smell, and balance. In short, senses are transducers from the physical world to the realm of the mind where we interpret the information, creating our perception of the world around us.

Olfactory bulb

The olfactory bulb 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 (OFC) and the hippocampus where it plays a role in emotion, memory and learning. The bulb is divided into two distinct structures: the main olfactory bulb and the accessory olfactory bulb. The main olfactory bulb connects to the amygdala via the piriform cortex of the primary olfactory cortex and directly projects from the main olfactory bulb to specific amygdala areas. The accessory olfactory bulb resides on the dorsal-posterior region of the main olfactory bulb and forms a parallel pathway. Destruction of the olfactory bulb results in ipsilateral anosmia, while irritative lesions of the uncus can result in olfactory and gustatory hallucinations.

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, sound, temperature, taste, pressure, and smell. The type and location of the sensory receptor activated by the stimulus plays the primary role in coding the sensation. All sensory modalities work together to heighten stimuli sensation when necessary.

Olfactory system Sensory system used for smelling

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 accessory olfactory system. The main olfactory system detects airborne substances, while the accessory system senses fluid-phase stimuli.

Olfactory receptor neuron

An olfactory receptor neuron (ORN), also called an olfactory sensory neuron (OSN), is a sensory neuron within the olfactory system.

Olfactory epithelium Specialised epithelial tissue in the nasal cavity that detects odours

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 epithelium is the part of the olfactory system directly responsible for detecting odors.

Glomerulus (olfaction)

The glomerulus is a spherical structure located in the olfactory bulb of the brain where synapses form between the terminals of the olfactory nerve and the dendrites of mitral, periglomerular and tufted cells. Each glomerulus is surrounded by a heterogeneous population of juxtaglomerular neurons and glial cells.

Olfactory receptors (ORs), also known as odorant receptors, are expressed in the cell membranes of olfactory receptor neurons and are responsible for the detection of odorants which give rise to the sense of smell. Activated olfactory receptors trigger nerve impulses which transmit information about odor to the brain. These receptors are members of the class A rhodopsin-like family of G protein-coupled receptors (GPCRs). The olfactory receptors form a multigene family consisting of around 800 genes in humans and 1400 genes in mice.

The docking theory of olfaction proposes that the smell of an odorant molecule is due to a range of weak non-covalent interactions between the odorant [a ligand] and its protein odorant receptor, such as electrostatic and Van der Waals interactions as well as H-bonding, dipole attraction, pi-stacking, metal ion, Cation–pi interaction, and hydrophobic effects, in addition to odorant conformation. While this type of recognition has previously been termed the shape theory of olfaction, which primarily considers molecular shape and size, this latter model is oversimplified since two scent molecules may have similar shapes and sizes but different sets of weak intermolecular forces and therefore activate different combinations of odorant receptors. Earlier “lock and key” and "hand in glove" models of protein−ligand binding has been replaced by a more nuanced pictures which consider the distortion of flexible molecules so as to form the optimal interactions with binding partners as in molecular docking of non-olfactory G-protein coupled receptors.

In medicine and anatomy, the special senses are the senses that have specialized organs devoted to them:

Olfactory tubercle Area at the bottom of the forebrain

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 play a role in locomotor and attentional behaviors, particularly in relation to social and sensory responsiveness, and it may be necessary for behavioral flexibility. The OT is interconnected with numerous brain regions, especially the sensory, arousal, and reward centers, thus making it a potentially critical interface between processing of sensory information and the subsequent behavioral responses.

A topographic map is the ordered projection of a sensory surface, like the retina or the skin, or an effector system, like the musculature, to one or more structures of the central nervous system. Topographic maps can be found in all sensory systems and in many motor systems.

Hyperosmia is an increased olfactory acuity, 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 the olfactory cortex. The causes of hyperosmia may be genetic, hormonal, environmental or the result of benzodiazepine withdrawal syndrome.

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 parosmia or phantosmia. Parosmia is a distortion in the perception of an odorant. Odorants smell different from what one remembers. Phantosmia is the perception of an odor when no odorant is present. The cause of dysosmia still remains a theory. It is typically considered a neurological disorder and clinical associations with the disorder have been made. Most cases are described as idiopathic and the main antecedents related to parosmia are URTIs, head trauma, and nasal and paranasal sinus disease. Dysosmia tends to go away on its own but there are options for treatment for patients that want immediate relief.

Olfactory memory refers to the recollection of odors. Studies have found various characteristics of common memories of odor memory including persistence and high resistance to interference. Explicit memory is typically the form focused on in the studies of olfactory memory, though implicit forms of memory certainly supply distinct contributions to the understanding of odors and memories of them. Research has demonstrated that the changes to the olfactory bulb and main olfactory system following birth are extremely important and influential for maternal behavior. Mammalian olfactory cues play an important role in the coordination of the mother infant bond, and the following normal development of the offspring. Maternal breast odors are individually distinctive, and provide a basis for recognition of the mother by her offspring.

Sniffing is a perceptually-relevant behavior, defined as the active sampling of odors through the nasal cavity for the purpose of information acquisition. This behavior, displayed by all terrestrial vertebrates, is typically identified based upon changes in respiratory frequency and/or amplitude, and is often studied in the context of odor guided behaviors and olfactory perceptual tasks. Sniffing is quantified by measuring intra-nasal pressure or flow or air or, while less accurate, through a strain gauge on the chest to measure total respiratory volume. Strategies for sniffing behavior vary depending upon the animal, with small animals displaying sniffing frequencies ranging from 4 to 12 Hz but larger animals (humans) sniffing at much lower frequencies, usually less than 2 Hz. Subserving sniffing behaviors, evidence for an "olfactomotor" circuit in the brain exists, wherein perception or expectation of an odor can trigger brain respiratory center to allow for the modulation of sniffing frequency and amplitude and thus acquisition of odor information. Sniffing is analogous to other stimulus sampling behaviors, including visual saccades, active touch, and whisker movements in small animals. Atypical sniffing has been reported in cases of neurological disorders, especially those disorders characterized by impaired motor function and olfactory perception.

Insect olfaction

Insect olfaction refers to the function of chemical receptors that enable insects to detect and identify volatile compounds for foraging, predator avoidance, finding mating partners and locating oviposition 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, olfaction is essential for hunting in many species of wasps, including Polybia sericea.

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 and taste modalities. Retronasal smell creates flavor from smell molecules in foods or drinks shunting up through the nasal passages as one is chewing. When people use the term "smell", they are usually referring to "orthonasal smell", or the perception of smell molecules that enter directly through the nose and up the nasal passages. Retronasal smell is critical for experiencing the flavor of foods and drinks. Flavor should be contrasted with taste, which refers to five specific dimensions: (1) sweet, (2) salty, (3) bitter, (4) sour, and (5) umami. Perceiving anything beyond these five dimensions, such as distinguishing the flavor of an apple from a pear for example, requires the sense of retronasal smell.

Neurogastronomy is the study of flavor perception and the ways it affects cognition and memory. This interdisciplinary field is influenced by the psychology and neuroscience of sensation, learning, satiety, and decision making. Areas of interest include how olfaction contributes to flavor, food addiction and obesity, taste preferences, and the linguistics of communicating and identifying flavor. The term neurogastronomy was coined by neuroscientist Gordon M. Shepherd.

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