Sense of smell - Knowledge (XXG)

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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.

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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.

1026:, 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 1110:. 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. 612:

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.

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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 80

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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

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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.

1102:. 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 302:, 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). 1227:

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

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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

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compared to respiratory epithelium (not innervated, or supplied with nerves) gives an indication of the animal's olfactory sensitivity. Humans have about 10 cm (1.6 sq in) of olfactory epithelium, whereas some dogs have 170 cm (26 sq in). A dog's olfactory epithelium is

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million 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

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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,

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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

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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

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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

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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

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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

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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

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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

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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

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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

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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

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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

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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

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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

433:; 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. 350:, 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. 1459: 1455: 1209:

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

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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

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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

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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

622:, 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 1146:

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

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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.

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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.

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Jaeger, SR; McRae, JF; Bava, CM; Beresford, MK; Hunter, D; Jia, Y; Chheang, SL; Jin, D; Peng, M; Gamble, JC; Atkinson, KR; Axten, LG; Paisley, AG; Tooman, L; Pineau, B; Rouse, SA; Newcomb, RD (2013).

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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

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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

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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

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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.

1894: 4309:"Olfactory object recognition, segmentation, adaptation, target seeking, and discrimination by the network of the olfactory bulb and cortex: computational model and experimental data" 282:. 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 2523:

Santos, PS; Schinemann, JA; Gabardo, J; da Graça Bicalho, M (2005). "New evidence that the MHC influences odor perception in humans: a study with 58 Southern Brazilian students".

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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

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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.

5061: 5260: 979:). Granular cells also mediate inhibition and excitation of mitral cells through pathways from centrifugal fibers and the anterior olfactory nuclei. Neuromodulators like 586:

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

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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.

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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

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Weisfeld, GE; Czilli, T; Phillips, KA; Gall, JA; Lichtman, CM (2003). "Possible olfaction-based mechanisms in human kin recognition and inbreeding avoidance".

3447: 962:, 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 3204: 1329:

has applied its standard in regulating numerous industries, landfills, and sewage treatment plants. Example applications this district has engaged are the

1155:. 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 317:. However, the behavioral predictions of this theory have been called into question. There is no theory yet that explains olfactory perception completely. 41: 4382: 6498: 2831: 2458: 2862: 1326: 4344: 1114:

remains a highly controversial concept, evidence exists for perceptual information implemented in the spatial dimensions of olfactory networks.

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

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have numerical standards of acceptability for the intensity of odor that is allowed to cross into a residential property. For example, the

3108: 724:. The olfactory epithelium is made up of at least six morphologically and biochemically different cell types. The proportion of olfactory 4687: 1160: 1127:, although pheromones can also be detected by the main olfactory system. In the accessory olfactory system, stimuli are detected by the 828: 2935:"Genetic variation in the odorant receptor OR2J3 is associated with the ability to detect the "grassy" smelling odor, cis-3-hexen-1-ol" 5053: 890:

Averaged activity of the receptor neurons can be measured in several ways. In vertebrates, responses to an odor can be measured by an

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de March, CA; Ryu, SE; Sicard, G; Moon, C; Golebiowski, J (2015). "Structure–odour relationships reviewed in the postgenomic era".

228: 5725: 3584:"Trpc2-expressing sensory neurons in the mouse main olfactory epithelium of type B express the soluble guanylate cyclase Gucy1b2" 422: 290:

There are, at present, a number of competing theories regarding the mechanism of odor coding and perception. According to the

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complex also acts to inhibit the binding of cAMP to the cAMP-dependent channel, thus contributing to olfactory adaptation.

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Crocker-Henderson system, which rates smells on a 0-8 scale for each of four "primary" smells: fragrant, acid, burnt, and

1277: 3056:"Odour of King Penguin feathers analysed using direct thermal desorption discriminates between individuals but not sexes" 2983: 1454:

Viruses can also infect the olfactory epithelium leading to a loss of the sense of olfaction. About 50% of patients with

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and hippocampus, areas of the brain that have long been known to be involved in emotion and place memory, respectively.

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Masaoka, Y; Satoh, H; Akai, L; Homma, I (2010). "Expiration: The moment we experience retronasal olfaction in flavor".

776:). This mucus acts as a solvent for odor molecules, flows constantly, and is replaced approximately every ten minutes. 6440: 1741: 1587: 3173: 445:. Functional imaging shows that this olfactory kinship detection process involves the frontal-temporal junction, the 6752: 6583: 6394: 5730: 1508: 1003: 579: 6203: 518:

Different people smell different odors, and most of these differences are caused by genetic differences. Although

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results from interactions between smell and taste, a smound may result from interactions between smell and sound.

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from ORs of one specific type and connect to the principal neurons of the OB, mitral and tufted cells (MT cells).

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is associated with the ability to detect the "grassy" odor, cis-3-hexen-1-ol. The preference (or dislike) of

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that the smell's contribution to flavor occurs, in contrast to that of proper smell, which occurs during the

313:, posits that odor receptors detect the frequencies of vibrations of odor molecules in the infrared range by 6726: 6624: 6130: 5506: 999: 772:(these are highly important, as the olfactory neurons provide a direct passage for infection to pass to the 757: 426: 3862:"Activation of raphe nuclei triggers rapid and distinct effects on parallel olfactory bulb output channels" 2854: 894:

or through calcium imaging of receptor neuron terminals in the olfactory bulb. In insects, one can perform

472:(MUP) gene cluster provides a highly polymorphic scent signal of genetic identity that appears to underlie 6808: 6782: 6744: 6578: 6294: 6125: 5735: 5521: 5340: 5213: 1503: 1492: 1334: 991:

all send axons to the olfactory bulb and have been implicated in gain modulation, pattern separation, and

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There are many different things which can interfere with a normal sense of smell, including damage to the

171: 143:) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and 110: 3633:"Contribution of the receptor guanylyl cyclase GC-D to chemosensory function in the olfactory epithelium" 6666: 6654: 6644: 6332: 5322: 5302: 1330: 1027: 891: 741: 465: 4821:

Gadenne, C; Barrozo, RB; Anton, S (2016). "Plasticity in insect olfaction: to smell or not to smell?".

733:. Often, land organisms will have separate olfaction systems for smell and taste (orthonasal smell and 5208: 4308: 2693:

Sherborne, AL; Thom, MD; Paterson, S; Jury, F; Ollier, WE; Stockley, P; Beynon, RJ; Hurst, JL (2007).

6521: 6193: 6160: 5859: 5574: 5478: 4987: 4890: 4787: 4014: 3813:"Cholinergic inputs from basal forebrain add an excitatory bias to odor coding in the olfactory bulb" 3703: 3644: 3536: 3481: 3396: 3239: 2897: 2799: 2706: 2200: 2106: 1973: 1497: 1372: 1346: 1098:

After binding the odorant, the receptor is activated and will send a signal to the glomeruli in the

1070:, smells are sensed by sensilla located on the antenna and maxillary palp and first processed by the 895: 788: 721: 458: 438: 5979: 5849: 5834: 5720: 5659: 5595: 1526: 1311: 1250: 1023: 968: 761: 279: 3055: 866:

This mechanism of transduction is somewhat unusual, in that cAMP works by directly binding to the

851:, further depolarizing the cell and triggering an action potential. Ca is then extruded through a 756:

of the olfactory sensory neurons. This may occur by diffusion or by the binding of the odorant to

6803: 6561: 6538: 5933: 5885: 5839: 5778: 5763: 5317: 4856: 4751: 4679: 4617: 4500: 4336: 4284: 4229: 4127: 3983: 3505: 3296: 2823: 2591: 2548: 2505: 2344: 2259: 1886: 1849: 1769: 1264: 1007: 976: 947: 884: 875: 749: 623: 519: 442: 314: 159: 3692:"An olfactory subsystem that detects carbon disulfide and mediates food-related social learning" 3104: 1432: 5201: 1921:"A novel multigene family may encode odorant receptors: a molecular basis for odor recognition" 679:

fishes use the sense of smell to identify mating partners or to alert to the presence of food.

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Keller, A; Vosshall, LB (2004). "A psychophysical test of the vibration theory of olfaction".

2216: 2173: 2132: 2075: 2024: 2001: 1942: 1878: 1761: 1722: 1669: 1607: 1513: 1304: 1233: 1224: 1182: 1128: 1060: 1019: 951: 812: 660: 559: 450: 339: 241: 206: 4648: 3015:"Highly contiguous genomes improve the understanding of avian olfactory receptor repertoires" 2811: 236:

Early scientific study of the sense of smell includes the extensive doctoral dissertation of

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Liberles, S (2006). "A second class of chemosensory receptors in the olfactory epithelium".

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Turin, L (2002). "A method for the calculation of odor character from molecular structure".

2163: 2122: 2114: 2065: 2057: 1991: 1981: 1932: 1868: 1753: 1712: 1702: 1636: 1471: 1176: 1107: 1041: 1037: 943: 939: 871: 844: 808: 804: 734: 707: 594:, which must always be aware of each other, and in those that smell for their food, such as 358: 343: 306: 167: 74: 526:

and its genetic variants (alleles) are responsible for our ability (or failure) to smell β-

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to guide movement, for example social insects may lay down a trail to a food source, or a

485: 473: 454: 5022: 2046:"The scent of life. The exquisite complexity of the sense of smell in animals and humans" 5223: 4991: 4894: 4791: 4738: 4713: 4018: 3707: 3648: 3540: 3485: 3243: 2901: 2886:"A mendelian trait for olfactory sensitivity affects odor experience and food selection" 2803: 2710: 2204: 2110: 1977: 1854:"What makes a discovery successful? The story of Linda Buck and the olfactory receptors" 258:

A modern demonstration of that theory was the cloning of olfactory receptor proteins by

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McRae, JF; Mainland, JD; Jaeger, SR; Adipietro, KA; Matsunami, H; Newcomb, RD (2012).

2622: 1996: 1961: 1757: 6797: 6430: 6425: 6379: 6317: 6302: 6198: 5920: 5864: 5679: 5646: 5630: 5549: 5493: 5453: 5402: 5393: 5350: 4860: 4774:

Carraher, C; Dalziel, J; Jordan, MD; Christie, DL; Newcomb, RD; Kralicek, AV (2015).

4621: 4264: 4162: 3559: 3524: 3418: 3134: 2827: 2579: 1937: 1920: 1890: 1362: 1322: 1071: 1045: 980: 924: 656: 595: 430: 259: 136: 4755: 4683: 4605: 4504: 4340: 4131: 3987: 3911:"Noradrenergic induction of odor-specific neural habituation and olfactory memories" 2595: 2509: 6730: 6710: 6384: 6363: 6348: 6322: 6140: 5961: 5938: 5910: 5905: 5829: 5768: 5531: 5516: 4978:

Syed, Z (2015). "Chemical ecology and olfaction in arthropod vectors of diseases".

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Porter, RH; Cernoch, JM; Balogh, RD (1985). "Odor signatures and kin recognition".

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Boehm, T; Zufall, F (2006). "MHC peptides and the sensory evaluation of genotype".

2417: 2378: 2348: 2263: 2095:"Odorant receptors of Drosophila are sensitive to the molecular volume of odorants" 1773: 1315: 1164: 972: 633: 627: 583: 489: 381:

phase of breathing. The olfactory system is the only human sense that bypasses the

362: 263: 163: 69: 3105:"Dogs' dazzling sense of smell: what lies behind their exceptional gift of sniff?" 1691:"Olfactory impairment & traumatic brain injury in blast-injured combat troops" 5153: 4324: 4102:

Leon, M; Johnson, BA (2003). "Olfactory coding in the mammalian olfactory bulb".

3764: 3525:"A receptor guanylyl cyclase expressed specifically in olfactory sensory neurons" 2332: 1707: 1656: 1441:– psychological disorder that causes the patient to imagine he or she has strong 103: 6543: 6445: 6353: 6327: 6312: 6274: 6097: 6072: 6067: 6057: 6043: 5669: 5625: 5501: 5430: 2536: 1689:

Xydakis, MS; Mulligan, LP; Smith, AB; Olsen, CH; Lyon, DM; Belluscio, L (2015).

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Proceedings of the National Academy of Sciences of the United States of America

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Proceedings of the National Academy of Sciences of the United States of America

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and low molecular weight nitrogenous compounds. Some insects, such as the moth

843:) into the cell, slightly depolarising it. The Ca in turn opens a Ca-activated 550:

The importance and sensitivity of smell varies among different organisms; most

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Frank, Marion E.; Fletcher, Dane B.; Hettinger, Thomas P. (1 September 2017).

4076: 4059: 4026: 3749:"A sensor for low environmental oxygen in the mouse main olfactory epithelium" 3715: 3599: 2910: 2885: 2764: 2718: 2168: 2151: 1463: 1447: 1424: 1394: 1238: 1044:. This is possibly due to the olfactory system's close anatomical ties to the 856: 769: 725: 713: 663:, scent glands occur atop the head. In many species, smell is highly tuned to 641: 619: 618:

as a group can smell one- to ten-million times more acutely than a human, and

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in humans) are a group of genes present in many animals and important for the

378: 374: 310: 5161: 4488: 4429: 4332: 4272: 4217: 4170: 2819: 2302: 2061: 1582: 500:, anemotaxis, klinotaxis, and tropotaxis. Their success is influenced by the 6213: 6077: 6023: 5890: 5420: 5410: 5355: 5345: 4421: 3657: 3251: 2950: 1986: 1475: 1442: 1202: 1151:, which in the mouse is located on the dorsal-posterior portion of the main 1124: 984: 820: 753: 664: 587: 567: 386: 248: 144: 85: 47: 5238: 5179: 5007: 4961: 4920: 4852: 4807: 4747: 4675: 4613: 4556: 4496: 4447: 4123: 4085: 4044: 3979: 3944: 3895: 3846: 3782: 3733: 3676: 3617: 3549: 3501: 3381: 3330: 3269: 3165: 3040: 2968: 2919: 2736: 2679: 2630: 2544: 2501: 2435: 2402:"Smelling sounds: olfactory-auditory convergence in the olfactory tubercle" 2386: 2340: 2255: 2220: 2212: 2136: 2079: 2005: 1882: 1765: 1726: 5196: 4280: 4225: 3568: 3031: 3014: 2786:

Reddy, Gautam; Murthy, Venkatesh N.; Vergassola, Massimo (10 March 2022).

2587: 2177: 1946: 405:. This neural convergence is proposed to give rise to a perception termed 6702: 6246: 6236: 6226: 6165: 6038: 5895: 5693: 1418: 1412: 1400: 1259: 1245:

Insects are capable of smelling and differentiating between thousands of

1194: 1156: 1011: 848: 784: 672: 668: 591: 571: 535: 382: 346:

plays the biggest role in the sensation of flavor. During the process of

295: 4178: 3493: 3362: 3311: 1384:

Specific terms are used to describe disorders associated with smelling:

116: 17: 6415: 6269: 6221: 6183: 6087: 6028: 5360: 4952: 4935: 4843: 4729: 4479: 4462: 4209: 3971: 1388: 1186: 1140: 1056: 836: 811:

pathway, depending on the organism. In mammals, the odorants stimulate

765: 648: 644: 402: 331: 271: 194: 183: 4936:"The insect ovipositor as a volatile sensor within a closed microcosm" 3074: 2661: 2118: 530:, a key aroma in foods and beverages. Similarly, the odorant receptor 453:, but not the primary or secondary olfactory cortices, or the related 5956: 5674: 5664: 5370: 5218: 4193: 4147:"Spatial properties of an EEG event in the olfactory bulb and cortex" 3811:

Rothermel, M; Carey, RM; Puche, A; Shipley, MT; Wachowiak, M (2014).

1640: 1292: 1287: 1198: 1067: 992: 931: 840: 832: 800: 783:, smells are sensed by olfactory sensory neurons in the chemosensory 780: 563: 551: 527: 496:

have been studied in different species, including gradient search or

406: 401:

information has been shown to converge in the olfactory tubercles of

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Mastrangelo, Andrea; Bonato, Matteo; Cinque, Paola (23 March 2021).

4667: 4647:

Soucy, ER; Albeanu, DF; Fantana, AL; Murthy, VN; Meister, M (2009).

4538: 4194:"Modeling the olfactory bulb and its neural oscillatory processings" 3877: 3301: 950:

converge in the outer layer of the olfactory bulb within small (≈50

630:, which was bred to track and hunt rabbits and other small animals. 604:

in contrast to having a weak sense of smell which is referred to as

3401: 2247: 626:

a few days old. The second-most-sensitive nose is possessed by the

6605: 5900: 5702: 5615: 5610: 5384: 5294: 5269: 4249:"Olfactory cortex: model circuit for study of associative memory?" 2646:"The neuronal substrates of human olfactory based kin recognition" 1815: 1211: 1136: 905: 773: 748:

that lines the superior portion of the cavity and are detected by

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Magnasco, MO (2015). "On the dimensionality of olfactory space".

2451:"Making scents of sounds: noises may alter how we perceive odors" 1223:

The two organs insects primarily use for detecting odors are the

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Neurogastronomy: how the brain creates flavor and why it matters

1816:"Lucretius, the biochemistry of smell, and scientific discovery" 1132: 963: 916: 555: 468:

is detrimental, it tends to be avoided. In the house mouse, the

275: 190: 174:

aggregate signals from these receptors and transmit them to the

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Lundström, JN; Boyle, JA; Zatorre, RJ; Jones-Gotman, M (2009).

244:, and implied that smell had a lower intensity discrimination. 5415: 4649:"Precision and diversity in an odor map on the olfactory bulb" 737:), but water-dwelling organisms usually have only one system. 226:, a Flemish tapestry depicting the sense of smell, 1484–1500. 5054:"Is there a dewey decimal system for the library of smells?" 2788:"Olfactory Sensing and Navigation in Turbulent Environments" 1427:– distortion in the absence of an odor, "hallucinated smell" 1282:

The tendrils of plants are especially sensitive to airborne

1002:, which synapses on five major regions of the cerebrum: the 2152:"A spectroscopic mechanism for primary olfactory reception" 1167:, where they may influence aggression and mating behavior. 88:

in the environment that are used to form the sense of smell

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3054:

Gabirot, M; Buatois, B; Müller, TC; Bonadonna, F (2018).

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can be utilized to determine the magnitude of an odor.

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3860:

Kapoor, V; Provost, A; Agarwal, P; Murthy, VN (2016).

1409:– the natural decline in the sense of smell in old age 942:, which in turn projects olfactory information to the 538:(coriander) has been linked to the olfactory receptor 4368:

Olfactory Cortex - an overview | ScienceDirect Topics

4247:

Haberly, Lewis B.; Bower, James M. (1 January 1989).

3798:

Morris' Human Anatomy: A Complete Systematic Treatise

2984:"Soapy taste of coriander linked to genetic variants" 1450:– aversion or psychological hypersensitivity to odors 1278:

Plant perception (physiology) § Senses in plants

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Electroencephalography and Clinical Neurophysiology

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Erlbaum Associates. 1232:typically reside in the 1163:, and from there to the 1149:accessory olfactory bulb 919:to the brain within the 853:sodium-calcium exchanger 758:odorant-binding proteins 223:The Lady and the Unicorn 6727:Medial forebrain bundle 6625:Lateral olfactory stria 6608:areas involved in smell 6131:Auditory verbal agnosia 5985:Juxtacapillary receptor 5224:Olfaction and Gustation 4253:Trends in Neurosciences 4077:10.1093/chemse/26.5.577 3915:Journal of Neuroscience 3817:Journal of Neuroscience 3658:10.1073/pnas.0704965104 3252:10.1126/science.1249168 2482:Trends in Neurosciences 2406:Journal of Neuroscience 2169:10.1093/chemse/21.6.773 1987:10.1073/pnas.0535697100 1189:to detect and identify 1000:lateral olfactory tract 847:, leading to efflux of 716:, smells are sensed by 361:receptors (also called 6783:Evolution of olfaction 6745:Medial olfactory stria 6126:Auditory hallucination 5736:Surface wave detection 5341:Multimodal integration 4198:Biological Cybernetics 4104:Brain Research Reviews 3550:10.1073/pnas.92.8.3571 3439:Boroditsky, L (1999). 2213:10.1006/jtbi.2001.2504 1561:"to emit a smell" and 1550: 1504:Olfactic Communication 1493:Evolution of olfaction 1335:Shoreline Amphitheatre 1299:Machine-based smelling 912: 514:Evolution of olfaction 449:, and the dorsomedial 274:have about a thousand 266:(who were awarded the 233: 203:traumatic brain injury 147:, and plays a role in 111:Anatomical terminology 6667:Periamygdaloid cortex 6655:Hippocampal formation 6645:EC-hippocampus system 6385:Phantom limb syndrome 6333:Tactile hallucination 5522:Glossopharyngeal (IX) 5323:Active sensory system 5209:Olfaction at cf.ac.uk 5204:at Wikimedia Commons 4422:10.1093/chemse/bjx031 3201:Science Now/AAAS News 2951:10.1093/chemse/bjs049 2525:Hormones and Behavior 2283:Shepherd, GM (2013). 1814:Holtsmark, E (1978). 1553:, past participle of 1331:San Mateo, California 1090:Coding and perception 1028:medial dorsal nucleus 934:sheaths, pass to the 909: 789:Chemosensory proteins 742:superior nasal concha 702:Main olfactory system 683:Human smell abilities 647:, and nonexistent in 470:major urinary protein 220: 6194:Labyrinthine fistula 6161:Spatial hearing loss 5860:Campaniform sensilla 5575:Somatosensory cortex 5142:Neuroscience Letters 4712:Brennan, PA (2001). 4588:Auffarth, B (2013). 3747:Bleymehl, K (2016). 2321:Neuroscience Letters 1746:The Lancet Neurology 1498:Nasal administration 1391:– inability to smell 1373:Hendrik Zwaardemaker 1347:Martinez, California 1286:. Parasites such as 1055:Inputs from the two 969:periglomerular cells 896:electroantennography 722:olfactory epithelium 712:In humans and other 459:orbitofrontal cortex 417:Inbreeding avoidance 193:or smell receptors, 6814:Otorhinolaryngology 5980:Nociceptin receptor 5850:Merkel nerve ending 5835:Mechanotransduction 5228:Neuroscience Online 5111:Hawkes, CH (2002). 5083:Henning, H (1915). 4992:2015COIS...10...83S 4895:2015CBio...25.R120L 4792:2015IBMB...66...31C 4656:Nature Neuroscience 4527:Nature Neuroscience 4058:Wilson, DA (2001). 4019:2009CBio...19.1561Z 3960:Nature Neuroscience 3921:(42): 10711–10719. 3866:Nature Neuroscience 3708:2010CBio...20.1438M 3649:2007PNAS..10414507L 3643:(36): 14507–14512. 3541:1995PNAS...92.3571F 3494:10.1038/nature05066 3486:2006Natur.442..645L 3363:10.7554/eLife.07865 3344:Meister, M (2015). 3312:10.7554/eLife.07865 3244:2014Sci...343.1370B 3238:(6177): 1370–1372. 3032:10.1093/icb/icab150 2902:2013CBio...23.1601J 2804:2022ARCMP..13..191R 2711:2007CBio...17.2061S 2650:Human Brain Mapping 2455:Scientific American 2236:Nature Neuroscience 2205:2002JThBi.216..367T 2111:2016NatSR...625103S 2044:Rinaldi, A (2007). 2019:Pinel, JPJ (2006). 1978:2003PNAS..100.3324G 1527:Scent transfer unit 1474:) can also disrupt 1078:), and next by the 1024:anterior commissure 948:olfactory receptors 892:electro-olfactogram 799:The binding of the 762:mucopolysaccharides 750:olfactory receptors 558:do not, except the 242:stimulus modalities 6562:Olfactory receptor 6539:Sustentacular cell 6342:Nociception (pain) 5934:Olfactory receptor 5886:Photoreceptor cell 5840:Lamellar corpuscle 5764:Photomorphogenesis 5626:nociception (pain) 5318:Sensory processing 5027:The New York Times 4953:10.1242/jeb.152777 4730:10.1007/pl00000880 4480:10.3758/BF03196369 4210:10.1007/BF00200803 3972:10.1038/nn0709-813 3690:Munger, S (2010). 3523:Fulle, HJ (1995). 2758:. Annual Reviews. 2099:Scientific Reports 1850:Ann-Sophie Barwich 1500:olfactory transfer 1470:and even the flu ( 1265:Deilephila elpenor 1191:volatile compounds 1183:chemical receptors 1074:(analogous to the 1008:olfactory tubercle 977:lateral inhibition 913: 885:lateral inhibition 443:Westermarck effect 315:quantum tunnelling 234: 6791: 6790: 6761: 6760: 6739: 6738: 6650:Entorhinal cortex 6596:Olfactory trigone 6474: 6473: 6459:Biases and errors 6454: 6453: 6390:Somatoparaphrenia 6359:Pain dissociation 6204:Ménière's disease 6136:Cortical deafness 6014:Visual impairment 5993: 5992: 5855:Bulbous corpuscle 5845:Tactile corpuscle 5813:sensory receptors 5805: 5804: 5688: 5687: 5641: 5640: 5606:Olfaction (smell) 5560:Vestibular cortex 5542:Cerebral cortices 5379: 5378: 5366:Motion perception 5200:Media related to 5122:978-0-7506-7287-0 4378:978-0-08-045046-9 4013:(18): 1561–1565. 3823:(13): 4654–4664. 3702:(16): 1438–1444. 3582:Omura, M (2015). 3480:(7103): 645–650. 3075:10.1111/ibi.12544 2896:(16): 1601–1605. 2756:Knowable Magazine 2705:(23): 2061–2066. 2662:10.1002/hbm.20686 2150:Turin, L (1996). 2119:10.1038/srep25103 1675:978-1-4292-3719-2 1613:978-0-87893-572-7 1581:Harper, Douglas. 1514:Olfactory fatigue 1305:Machine olfaction 1129:vomeronasal organ 1061:binocular rivalry 1020:entorhinal cortex 813:adenylate cyclase 661:red-bellied lemur 484:Some animals use 451:prefrontal cortex 125: 124: 120: 50:. Olfaction uses 16:(Redirected from 6821: 6778:Olfactory system 6707:Stria terminalis 6687:Habenular nuclei 6683:Stria medullaris 6662:Prepyriform area 6630: 6612: 6501: 6494: 6487: 6478: 6121:Auditory agnosia 6053:Optic neuropathy 6002: 5870:Stretch receptor 5818: 5716:Magnetoreception 5711:Electroreception 5699: 5621:mechanoreception 5570:Gustatory cortex 5565:Olfactory cortex 5399: 5390: 5308:Sensory receptor 5291: 5263: 5256: 5249: 5240: 5199: 5184: 5183: 5173: 5133: 5127: 5126: 5108: 5099: 5098: 5080: 5074: 5073: 5071: 5069: 5058:Nadia Berenstein 5049: 5043: 5042: 5040: 5038: 5018: 5012: 5011: 4975: 4966: 4965: 4955: 4946:(9): 1554–1557. 4931: 4925: 4924: 4914: 4889:(3): R120–R129. 4874: 4865: 4864: 4846: 4818: 4812: 4811: 4771: 4760: 4759: 4741: 4709: 4703: 4702: 4700: 4698: 4692: 4653: 4644: 4638: 4637: 4635: 4633: 4600:(8): 1667–1679. 4585: 4576: 4575: 4573: 4571: 4565: 4550: 4533:(9): 1225–1232. 4524: 4515: 4509: 4508: 4482: 4458: 4452: 4451: 4441: 4401: 4395: 4394: 4392: 4390: 4363: 4357: 4356: 4354: 4352: 4304: 4293: 4292: 4244: 4238: 4237: 4189: 4183: 4182: 4142: 4136: 4135: 4099: 4090: 4089: 4079: 4055: 4049: 4048: 4038: 3998: 3992: 3991: 3955: 3949: 3948: 3938: 3906: 3900: 3899: 3889: 3857: 3851: 3850: 3840: 3808: 3802: 3801: 3793: 3787: 3786: 3776: 3759:(6): 1196–1203. 3744: 3738: 3737: 3727: 3687: 3681: 3680: 3670: 3660: 3628: 3622: 3621: 3611: 3579: 3573: 3572: 3562: 3552: 3535:(8): 3571–3575. 3520: 3514: 3513: 3469: 3463: 3462: 3460: 3458: 3452: 3445: 3436: 3430: 3429: 3423: 3413: 3407: 3406: 3404: 3392: 3386: 3385: 3375: 3365: 3341: 3335: 3334: 3324: 3314: 3304: 3280: 3274: 3273: 3263: 3223: 3217: 3216: 3214: 3212: 3192: 3186: 3185: 3183: 3181: 3161: 3155: 3154: 3152: 3150: 3130: 3121: 3120: 3118: 3116: 3100: 3094: 3093: 3091: 3089: 3083: 3060: 3051: 3045: 3044: 3034: 3025:(4): 1281–1290. 3010: 3004: 3003: 3001: 2999: 2979: 2973: 2972: 2962: 2930: 2924: 2923: 2913: 2881: 2875: 2874: 2872: 2870: 2850: 2844: 2843: 2841: 2839: 2783: 2777: 2776: 2774: 2772: 2767: 2747: 2741: 2740: 2730: 2690: 2684: 2683: 2673: 2656:(8): 2571–2580. 2641: 2635: 2634: 2606: 2600: 2599: 2563: 2557: 2556: 2520: 2514: 2513: 2477: 2471: 2470: 2468: 2466: 2446: 2440: 2439: 2429: 2412:(8): 3013–3021. 2397: 2391: 2390: 2362: 2353: 2352: 2316: 2307: 2306: 2280: 2269: 2267: 2231: 2225: 2224: 2188: 2182: 2181: 2171: 2147: 2141: 2140: 2130: 2090: 2084: 2083: 2073: 2041: 2035: 2034: 2016: 2010: 2009: 1999: 1989: 1972:(6): 3324–3327. 1957: 1951: 1950: 1940: 1916: 1910: 1909: 1907: 1905: 1899: 1876: 1858: 1846: 1840: 1839: 1837: 1835: 1811: 1805: 1804: 1796: 1790: 1789: 1787: 1785: 1737: 1731: 1730: 1720: 1710: 1686: 1680: 1679: 1659: 1651: 1645: 1644: 1641:10.1002/ffj.3249 1624: 1618: 1617: 1599: 1593: 1592: 1578: 1566: 1543: 1256:carboxylic acids 1177:Insect olfaction 1108:olfactory cortex 1042:emotional memory 1038:long-term memory 967:also project to 944:olfactory cortex 940:cribriform plate 929: 872:protein kinase A 845:chloride channel 809:second messenger 805:action potential 735:retronasal smell 708:Olfactory system 691: 520:odorant receptor 480:Guiding movement 439:incest avoidance 425:genes (known as 344:Retronasal smell 307:vibration theory 254: 213:History of study 168:olfactory system 117:edit on Wikidata 75:Olfactory system 44: 32: 21: 6829: 6828: 6824: 6823: 6822: 6820: 6819: 6818: 6794: 6793: 6792: 6787: 6757: 6735: 6715: 6691: 6671: 6640:Piriform cortex 6619: 6600: 6591:Olfactory tract 6554:Olfactory nerve 6548: 6510: 6505: 6475: 6470: 6450: 6399: 6368: 6337: 6289: 6256: 6208: 6170: 6102: 6093:Stereoblindness 6034:Color blindness 5989: 5966: 5943: 5915: 5874: 5822:Mechanoreceptor 5811: 5801: 5797:Machine hearing 5792:Computer vision 5787:Robotic sensing 5773: 5750: 5684: 5637: 5579: 5555:Auditory cortex 5536: 5483: 5446:Sensory systems 5440: 5375: 5327: 5285: 5283: 5276: 5267: 5219:Smells Database 5193: 5188: 5187: 5135: 5134: 5130: 5123: 5110: 5109: 5102: 5095: 5082: 5081: 5077: 5067: 5065: 5051: 5050: 5046: 5036: 5034: 5020: 5019: 5015: 4977: 4976: 4969: 4933: 4932: 4928: 4883:Current Biology 4876: 4875: 4868: 4820: 4819: 4815: 4773: 4772: 4763: 4711: 4710: 4706: 4696: 4694: 4690: 4668:10.1038/nn.2262 4651: 4646: 4645: 4641: 4631: 4629: 4587: 4586: 4579: 4569: 4567: 4563: 4539:10.1038/nn.3775 4522: 4517: 4516: 4512: 4460: 4459: 4455: 4410:Chemical Senses 4403: 4402: 4398: 4388: 4386: 4379: 4365: 4364: 4360: 4350: 4348: 4306: 4305: 4296: 4246: 4245: 4241: 4191: 4190: 4186: 4144: 4143: 4139: 4101: 4100: 4093: 4064:Chemical Senses 4057: 4056: 4052: 4007:Current Biology 4000: 3999: 3995: 3957: 3956: 3952: 3908: 3907: 3903: 3878:10.1038/nn.4219 3859: 3858: 3854: 3810: 3809: 3805: 3795: 3794: 3790: 3746: 3745: 3741: 3696:Current Biology 3689: 3688: 3684: 3630: 3629: 3625: 3581: 3580: 3576: 3522: 3521: 3517: 3471: 3470: 3466: 3456: 3454: 3450: 3443: 3438: 3437: 3433: 3415: 3414: 3410: 3394: 3393: 3389: 3343: 3342: 3338: 3282: 3281: 3277: 3225: 3224: 3220: 3210: 3208: 3194: 3193: 3189: 3179: 3177: 3163: 3162: 3158: 3148: 3146: 3132: 3131: 3124: 3114: 3112: 3102: 3101: 3097: 3087: 3085: 3081: 3058: 3053: 3052: 3048: 3012: 3011: 3007: 2997: 2995: 2981: 2980: 2976: 2939:Chemical Senses 2932: 2931: 2927: 2890:Current Biology 2883: 2882: 2878: 2868: 2866: 2852: 2851: 2847: 2837: 2835: 2785: 2784: 2780: 2770: 2768: 2749: 2748: 2744: 2699:Current Biology 2692: 2691: 2687: 2643: 2642: 2638: 2608: 2607: 2603: 2565: 2564: 2560: 2522: 2521: 2517: 2479: 2478: 2474: 2464: 2462: 2448: 2447: 2443: 2399: 2398: 2394: 2364: 2363: 2356: 2318: 2317: 2310: 2295: 2282: 2281: 2272: 2233: 2232: 2228: 2190: 2189: 2185: 2156:Chemical Senses 2149: 2148: 2144: 2092: 2091: 2087: 2043: 2042: 2038: 2031: 2018: 2017: 2013: 1959: 1958: 1954: 1918: 1917: 1913: 1903: 1901: 1897: 1856: 1848: 1847: 1843: 1833: 1831: 1813: 1812: 1808: 1798: 1797: 1793: 1783: 1781: 1739: 1738: 1734: 1701:(15): 1559–67. 1688: 1687: 1683: 1676: 1653: 1652: 1648: 1626: 1625: 1621: 1614: 1601: 1600: 1596: 1580: 1579: 1575: 1570: 1569: 1544: 1540: 1535: 1488:Electronic nose 1484: 1472:influenza virus 1382: 1368:Henning's prism 1355: 1307: 1301: 1280: 1274: 1249:compounds both 1217:Polybia sericea 1205:) and locating 1179: 1173: 1120: 1092: 1080:mushroom bodies 1016:piriform cortex 927: 921:olfactory nerve 904: 826: 797: 795:Receptor neuron 710: 704: 699: 689: 685: 548: 516: 510: 482: 474:kin recognition 455:piriform cortex 419: 395: 328: 323: 252: 215: 121: 59: 28: 23: 22: 15: 12: 11: 5: 6827: 6825: 6817: 6816: 6811: 6806: 6796: 6795: 6789: 6788: 6786: 6785: 6780: 6775: 6769: 6767: 6763: 6762: 6759: 6758: 6756: 6755: 6749: 6747: 6741: 6740: 6737: 6736: 6734: 6733: 6723: 6721: 6717: 6716: 6714: 6713: 6699: 6697: 6693: 6692: 6690: 6689: 6679: 6677: 6673: 6672: 6670: 6669: 6664: 6659: 6658: 6657: 6652: 6642: 6636: 6634: 6627: 6621: 6620: 6615: 6609: 6602: 6601: 6599: 6598: 6593: 6588: 6587: 6586: 6581: 6574:Olfactory bulb 6571: 6570: 6569: 6558: 6556: 6550: 6549: 6547: 6546: 6541: 6536: 6535: 6534: 6529: 6518: 6516: 6512: 6511: 6506: 6504: 6503: 6496: 6489: 6481: 6472: 6471: 6469: 6468: 6462: 6460: 6456: 6455: 6452: 6451: 6449: 6448: 6443: 6438: 6433: 6428: 6423: 6418: 6413: 6407: 6405: 6401: 6400: 6398: 6397: 6392: 6387: 6382: 6376: 6374: 6373:Proprioception 6370: 6369: 6367: 6366: 6361: 6356: 6351: 6345: 6343: 6339: 6338: 6336: 6335: 6330: 6325: 6320: 6315: 6310: 6305: 6299: 6297: 6291: 6290: 6288: 6287: 6282: 6277: 6272: 6266: 6264: 6258: 6257: 6255: 6254: 6249: 6244: 6239: 6234: 6229: 6224: 6218: 6216: 6210: 6209: 6207: 6206: 6201: 6196: 6191: 6186: 6180: 6178: 6172: 6171: 6169: 6168: 6163: 6158: 6153: 6148: 6143: 6138: 6133: 6128: 6123: 6118: 6112: 6110: 6104: 6103: 6101: 6100: 6095: 6090: 6085: 6080: 6075: 6070: 6065: 6060: 6055: 6050: 6041: 6036: 6031: 6026: 6021: 6016: 6010: 6008: 5999: 5995: 5994: 5991: 5990: 5988: 5987: 5982: 5976: 5974: 5968: 5967: 5965: 5964: 5959: 5953: 5951: 5949:Thermoreceptor 5945: 5944: 5942: 5941: 5936: 5931: 5929:Taste receptor 5925: 5923: 5917: 5916: 5914: 5913: 5908: 5903: 5898: 5893: 5888: 5882: 5880: 5876: 5875: 5873: 5872: 5867: 5862: 5857: 5852: 5847: 5842: 5837: 5832: 5826: 5824: 5815: 5807: 5806: 5803: 5802: 5800: 5799: 5794: 5789: 5783: 5781: 5775: 5774: 5772: 5771: 5766: 5760: 5758: 5752: 5751: 5749: 5748: 5743: 5738: 5733: 5728: 5723: 5718: 5713: 5707: 5705: 5696: 5690: 5689: 5686: 5685: 5683: 5682: 5677: 5672: 5667: 5662: 5657: 5655:Proprioception 5651: 5649: 5643: 5642: 5639: 5638: 5636: 5635: 5634: 5633: 5628: 5623: 5613: 5608: 5603: 5598: 5593: 5587: 5585: 5581: 5580: 5578: 5577: 5572: 5567: 5562: 5557: 5552: 5546: 5544: 5538: 5537: 5535: 5534: 5529: 5527:Trigeminal (V) 5524: 5519: 5514: 5509: 5504: 5498: 5496: 5485: 5484: 5482: 5481: 5476: 5471: 5466: 5461: 5456: 5450: 5448: 5442: 5441: 5439: 5438: 5433: 5428: 5423: 5418: 5413: 5407: 5405: 5403:Sensory organs 5396: 5387: 5381: 5380: 5377: 5376: 5374: 5373: 5368: 5363: 5358: 5353: 5348: 5343: 5337: 5335: 5329: 5328: 5326: 5325: 5320: 5315: 5310: 5305: 5299: 5297: 5288: 5278: 5277: 5268: 5266: 5265: 5258: 5251: 5243: 5237: 5236: 5231: 5221: 5216: 5211: 5192: 5191:External links 5189: 5186: 5185: 5128: 5121: 5100: 5093: 5075: 5044: 5013: 4967: 4926: 4866: 4813: 4761: 4724:(4): 546–555. 4704: 4662:(2): 210–220. 4639: 4577: 4510: 4473:(2): 244–264. 4453: 4416:(7): 537–546. 4396: 4377: 4358: 4294: 4259:(7): 258–264. 4239: 4204:(5): 379–392. 4184: 4157:(5): 586–605. 4137: 4091: 4070:(5): 577–584. 4050: 3993: 3966:(7): 813–814. 3950: 3901: 3872:(2): 813–814. 3852: 3803: 3788: 3739: 3682: 3623: 3574: 3515: 3464: 3431: 3408: 3402:10.1101/022103 3387: 3336: 3275: 3218: 3187: 3156: 3122: 3095: 3069:(2): 379–389. 3046: 3005: 2974: 2945:(7): 585–593. 2925: 2876: 2845: 2798:(1): 191–213. 2778: 2742: 2685: 2636: 2617:(3): 279–295. 2601: 2574:(3): 445–448. 2558: 2531:(4): 384–388. 2515: 2488:(2): 100–107. 2472: 2441: 2392: 2373:(1): 423–452. 2354: 2308: 2293: 2270: 2248:10.1038/nn1215 2242:(4): 337–338. 2226: 2199:(3): 367–385. 2183: 2162:(6): 773–791. 2142: 2085: 2036: 2029: 2011: 1952: 1931:(1): 175–187. 1911: 1867:(4): 749–753. 1841: 1806: 1791: 1752:(6): 415–416. 1732: 1681: 1674: 1646: 1635:(5): 342–361. 1619: 1612: 1594: 1572: 1571: 1568: 1567: 1537: 1536: 1534: 1531: 1530: 1529: 1524: 1516: 1511: 1506: 1501: 1495: 1490: 1483: 1480: 1452: 1451: 1445: 1436: 1430: 1429: 1428: 1422: 1410: 1404: 1398: 1392: 1381: 1378: 1377: 1376: 1369: 1366: 1354: 1353:Classification 1351: 1343:IT Corporation 1303:Main article: 1300: 1297: 1273: 1270: 1201:partners (via 1175:Main article: 1172: 1169: 1153:olfactory bulb 1119: 1116: 1100:olfactory bulb 1091: 1088: 1076:olfactory bulb 989:norepinephrine 973:granular cells 936:olfactory bulb 903: 900: 876:photoreceptors 824: 815:to synthesize 796: 793: 706:Main article: 703: 700: 698: 695: 684: 681: 659:, such as the 657:strepsirrhines 547: 544: 512:Main article: 509: 506: 481: 478: 418: 415: 394: 391: 327: 324: 322: 319: 300:odotope theory 280:odor reception 278:that code for 238:Eleanor Gamble 214: 211: 176:olfactory bulb 129:sense of smell 123: 122: 113: 107: 106: 101: 95: 94: 90: 89: 82: 78: 77: 72: 66: 65: 61: 60: 52:chemoreceptors 45: 37: 36: 26: 24: 14: 13: 10: 9: 6: 4: 3: 2: 6826: 6815: 6812: 6810: 6809:Limbic system 6807: 6805: 6802: 6801: 6799: 6784: 6781: 6779: 6776: 6774: 6771: 6770: 6768: 6764: 6754: 6751: 6750: 6748: 6746: 6742: 6732: 6728: 6725: 6724: 6722: 6718: 6712: 6708: 6704: 6701: 6700: 6698: 6694: 6688: 6684: 6681: 6680: 6678: 6674: 6668: 6665: 6663: 6660: 6656: 6653: 6651: 6648: 6647: 6646: 6643: 6641: 6638: 6637: 6635: 6631: 6628: 6626: 6622: 6618: 6613: 6610: 6607: 6603: 6597: 6594: 6592: 6589: 6585: 6582: 6580: 6577: 6576: 6575: 6572: 6568: 6565: 6564: 6563: 6560: 6559: 6557: 6555: 6551: 6545: 6542: 6540: 6537: 6533: 6530: 6528: 6525: 6524: 6523: 6520: 6519: 6517: 6513: 6509: 6502: 6497: 6495: 6490: 6488: 6483: 6482: 6479: 6467: 6464: 6463: 6461: 6457: 6447: 6444: 6442: 6439: 6437: 6434: 6432: 6431:Hallucination 6429: 6427: 6426:Derealization 6424: 6422: 6419: 6417: 6414: 6412: 6409: 6408: 6406: 6402: 6396: 6393: 6391: 6388: 6386: 6383: 6381: 6380:Asomatognosia 6378: 6377: 6375: 6371: 6365: 6362: 6360: 6357: 6355: 6352: 6350: 6347: 6346: 6344: 6340: 6334: 6331: 6329: 6326: 6324: 6321: 6319: 6318:Hyperesthesia 6316: 6314: 6311: 6309: 6306: 6304: 6303:Astereognosis 6301: 6300: 6298: 6296: 6292: 6286: 6283: 6281: 6278: 6276: 6273: 6271: 6268: 6267: 6265: 6263: 6259: 6253: 6250: 6248: 6245: 6243: 6240: 6238: 6235: 6233: 6230: 6228: 6225: 6223: 6220: 6219: 6217: 6215: 6211: 6205: 6202: 6200: 6199:Labyrinthitis 6197: 6195: 6192: 6190: 6187: 6185: 6182: 6181: 6179: 6177: 6173: 6167: 6164: 6162: 6159: 6157: 6154: 6152: 6149: 6147: 6144: 6142: 6139: 6137: 6134: 6132: 6129: 6127: 6124: 6122: 6119: 6117: 6114: 6113: 6111: 6109: 6105: 6099: 6096: 6094: 6091: 6089: 6086: 6084: 6081: 6079: 6076: 6074: 6071: 6069: 6066: 6064: 6061: 6059: 6056: 6054: 6051: 6049: 6045: 6042: 6040: 6037: 6035: 6032: 6030: 6027: 6025: 6022: 6020: 6017: 6015: 6012: 6011: 6009: 6007: 6003: 6000: 5996: 5986: 5983: 5981: 5978: 5977: 5975: 5973: 5969: 5963: 5960: 5958: 5955: 5954: 5952: 5950: 5946: 5940: 5937: 5935: 5932: 5930: 5927: 5926: 5924: 5922: 5921:Chemoreceptor 5918: 5912: 5909: 5907: 5904: 5902: 5899: 5897: 5894: 5892: 5889: 5887: 5884: 5883: 5881: 5879:Photoreceptor 5877: 5871: 5868: 5866: 5865:Slit sensilla 5863: 5861: 5858: 5856: 5853: 5851: 5848: 5846: 5843: 5841: 5838: 5836: 5833: 5831: 5828: 5827: 5825: 5823: 5819: 5816: 5814: 5808: 5798: 5795: 5793: 5790: 5788: 5785: 5784: 5782: 5780: 5776: 5770: 5767: 5765: 5762: 5761: 5759: 5757: 5753: 5747: 5744: 5742: 5739: 5737: 5734: 5732: 5729: 5727: 5724: 5722: 5719: 5717: 5714: 5712: 5709: 5708: 5706: 5704: 5700: 5697: 5695: 5691: 5681: 5680:Visceral pain 5678: 5676: 5673: 5671: 5668: 5666: 5663: 5661: 5658: 5656: 5653: 5652: 5650: 5648: 5644: 5632: 5631:thermoception 5629: 5627: 5624: 5622: 5619: 5618: 5617: 5614: 5612: 5609: 5607: 5604: 5602: 5599: 5597: 5594: 5592: 5589: 5588: 5586: 5582: 5576: 5573: 5571: 5568: 5566: 5563: 5561: 5558: 5556: 5553: 5551: 5550:Visual cortex 5548: 5547: 5545: 5543: 5539: 5533: 5530: 5528: 5525: 5523: 5520: 5518: 5515: 5513: 5512:Olfactory (I) 5510: 5508: 5505: 5503: 5500: 5499: 5497: 5495: 5494:spinal nerves 5491: 5486: 5480: 5477: 5475: 5472: 5470: 5467: 5465: 5462: 5460: 5457: 5455: 5452: 5451: 5449: 5447: 5443: 5437: 5434: 5432: 5429: 5427: 5424: 5422: 5419: 5417: 5414: 5412: 5409: 5408: 5406: 5404: 5400: 5397: 5395: 5391: 5388: 5386: 5382: 5372: 5369: 5367: 5364: 5362: 5359: 5357: 5354: 5352: 5351:Consciousness 5349: 5347: 5344: 5342: 5339: 5338: 5336: 5334: 5330: 5324: 5321: 5319: 5316: 5314: 5311: 5309: 5306: 5304: 5301: 5300: 5298: 5296: 5292: 5289: 5287: 5279: 5275: 5271: 5264: 5259: 5257: 5252: 5250: 5245: 5244: 5241: 5235: 5232: 5229: 5225: 5222: 5220: 5217: 5215: 5212: 5210: 5207: 5206: 5205: 5203: 5198: 5190: 5181: 5177: 5172: 5167: 5163: 5159: 5155: 5151: 5147: 5143: 5139: 5132: 5129: 5124: 5118: 5114: 5107: 5105: 5101: 5096: 5094:9783750176669 5090: 5086: 5079: 5076: 5063: 5059: 5055: 5048: 5045: 5032: 5028: 5024: 5017: 5014: 5009: 5005: 5001: 4997: 4993: 4989: 4985: 4981: 4974: 4972: 4968: 4963: 4959: 4954: 4949: 4945: 4941: 4937: 4930: 4927: 4922: 4918: 4913: 4908: 4904: 4900: 4896: 4892: 4888: 4884: 4880: 4873: 4871: 4867: 4862: 4858: 4854: 4850: 4845: 4840: 4836: 4832: 4828: 4824: 4817: 4814: 4809: 4805: 4801: 4797: 4793: 4789: 4785: 4781: 4777: 4770: 4768: 4766: 4762: 4757: 4753: 4749: 4745: 4740: 4735: 4731: 4727: 4723: 4719: 4715: 4708: 4705: 4689: 4685: 4681: 4677: 4673: 4669: 4665: 4661: 4657: 4650: 4643: 4640: 4627: 4623: 4619: 4615: 4611: 4607: 4603: 4599: 4595: 4591: 4584: 4582: 4578: 4562: 4558: 4554: 4549: 4544: 4540: 4536: 4532: 4528: 4521: 4514: 4511: 4506: 4502: 4498: 4494: 4490: 4486: 4481: 4476: 4472: 4468: 4464: 4457: 4454: 4449: 4445: 4440: 4435: 4431: 4427: 4423: 4419: 4415: 4411: 4407: 4400: 4397: 4384: 4380: 4374: 4370: 4369: 4362: 4359: 4346: 4342: 4338: 4334: 4330: 4326: 4322: 4318: 4314: 4310: 4303: 4301: 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2032: 2030:0-205-42651-4 2026: 2022: 2021:Biopsychology 2015: 2012: 2007: 2003: 1998: 1993: 1988: 1983: 1979: 1975: 1971: 1967: 1963: 1956: 1953: 1948: 1944: 1939: 1934: 1930: 1926: 1922: 1915: 1912: 1896: 1892: 1888: 1884: 1880: 1875: 1870: 1866: 1862: 1855: 1851: 1845: 1842: 1829: 1825: 1821: 1817: 1810: 1807: 1802: 1795: 1792: 1779: 1775: 1771: 1767: 1763: 1759: 1755: 1751: 1747: 1743: 1736: 1733: 1728: 1724: 1719: 1714: 1709: 1704: 1700: 1696: 1692: 1685: 1682: 1677: 1671: 1667: 1663: 1658: 1650: 1647: 1642: 1638: 1634: 1630: 1623: 1620: 1615: 1609: 1605: 1598: 1595: 1590: 1589: 1584: 1577: 1574: 1564: 1560: 1556: 1552: 1548: 1542: 1539: 1532: 1528: 1525: 1523: 1521: 1517: 1515: 1512: 1510: 1507: 1505: 1502: 1499: 1496: 1494: 1491: 1489: 1486: 1485: 1481: 1479: 1477: 1473: 1469: 1465: 1461: 1457: 1449: 1446: 1444: 1440: 1437: 1434: 1431: 1426: 1423: 1420: 1417: 1416: 1414: 1411: 1408: 1405: 1402: 1399: 1396: 1393: 1390: 1387: 1386: 1385: 1379: 1374: 1370: 1367: 1364: 1360: 1359: 1358: 1352: 1350: 1348: 1345:waste ponds, 1344: 1340: 1336: 1332: 1328: 1324: 1319: 1317: 1313: 1306: 1298: 1296: 1294: 1289: 1285: 1279: 1271: 1269: 1267: 1266: 1261: 1257: 1252: 1248: 1243: 1241: 1240: 1235: 1231: 1226: 1221: 1219: 1218: 1213: 1208: 1204: 1200: 1196: 1192: 1188: 1184: 1178: 1170: 1168: 1166: 1162: 1158: 1154: 1150: 1144: 1142: 1138: 1134: 1130: 1126: 1117: 1115: 1111: 1109: 1105: 1101: 1096: 1089: 1087: 1085: 1081: 1077: 1073: 1072:antennal lobe 1069: 1064: 1062: 1058: 1053: 1049: 1047: 1046:limbic system 1043: 1039: 1034: 1029: 1025: 1021: 1017: 1013: 1009: 1005: 1001: 996: 994: 990: 986: 982: 981:acetylcholine 978: 974: 970: 965: 961: 957: 953: 949: 945: 941: 937: 933: 926: 925:cranial nerve 922: 918: 908: 901: 899: 897: 893: 888: 886: 881: 877: 873: 869: 864: 860: 858: 854: 850: 846: 842: 838: 834: 830: 822: 818: 814: 810: 806: 802: 794: 792: 790: 786: 782: 777: 775: 771: 767: 763: 759: 755: 751: 747: 743: 738: 736: 732: 727: 723: 719: 715: 709: 701: 696: 694: 682: 680: 676: 674: 670: 666: 662: 658: 654: 650: 646: 643: 638: 635: 634:Grizzly bears 631: 629: 625: 621: 617: 613: 609: 607: 603: 602: 597: 593: 589: 585: 581: 577: 573: 569: 565: 561: 557: 553: 545: 543: 541: 537: 533: 529: 525: 521: 515: 507: 505: 503: 499: 495: 491: 487: 479: 477: 475: 471: 467: 462: 460: 456: 452: 448: 444: 440: 434: 432: 431:immune system 428: 424: 416: 414: 412: 408: 404: 400: 392: 390: 388: 384: 380: 376: 372: 368: 364: 360: 356: 351: 349: 345: 341: 337: 333: 325: 320: 318: 316: 312: 308: 303: 301: 297: 293: 288: 285: 281: 277: 273: 269: 265: 261: 260:Linda B. Buck 256: 250: 245: 243: 239: 231: 230: 225: 224: 219: 212: 210: 208: 204: 200: 196: 192: 187: 185: 181: 177: 173: 169: 165: 161: 157: 152: 150: 146: 142: 138: 137:special sense 134: 130: 118: 112: 108: 105: 102: 100: 96: 91: 87: 83: 79: 76: 73: 71: 67: 62: 57: 53: 49: 43: 38: 33: 30: 19: 6731:Hypothalamus 6711:Hypothalamus 6584:mitral cells 6515:Microanatomy 6507: 6364:Phantom pain 6349:Hyperalgesia 6323:Hypoesthesia 6141:Hearing loss 5962:TRP channels 5939:Osmoreceptor 5906:Photopigment 5830:Baroreceptor 5769:Gravitropism 5741:Frog hearing 5721:Echolocation 5605: 5517:Facial (VII) 5227: 5194: 5145: 5141: 5131: 5112: 5084: 5078: 5066:. Retrieved 5057: 5047: 5035:. Retrieved 5026: 5016: 4983: 4979: 4943: 4939: 4929: 4886: 4882: 4826: 4822: 4816: 4783: 4779: 4721: 4717: 4707: 4695:. Retrieved 4659: 4655: 4642: 4630:. Retrieved 4597: 4593: 4568:. Retrieved 4530: 4526: 4513: 4470: 4466: 4456: 4413: 4409: 4399: 4387:. Retrieved 4367: 4361: 4349:. 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The human 363:chemesthesis 352: 329: 309:proposed by 304: 292:shape theory 289: 264:Richard Axel 257: 246: 235: 227: 221: 188: 164:nasal cavity 153: 132: 128: 126: 29: 6544:Tufted cell 6446:Synesthesia 6354:Hypoalgesia 6328:Paresthesia 6313:Formication 6308:CMT disease 6275:Hypergeusia 6098:Visual snow 6073:Photophobia 6068:Papilledema 6058:Oscillopsia 6044:Hemeralopia 5911:Aureochrome 5746:Toad vision 5670:Suffocation 5584:Perceptions 5037:22 February 4844:11336/19586 4829:: 317–333. 4632:19 February 3594:: 114–124. 3302:1411.0165v2 2998:7 September 2988:Nature News 2869:7 September 2859:Nature News 2465:30 December 1904:17 February 1583:"olfaction" 1407:Presbyosmia 1251:sensitively 1207:oviposition 1033:hippocampus 952:micrometers 868:ion channel 714:vertebrates 624:scent trail 620:bloodhounds 616:Scenthounds 606:microsmotic 601:macrosmatic 568:albatrosses 348:mastication 268:Nobel Prize 162:within the 158:binds to a 93:Identifiers 6798:Categories 6522:Epithelium 6466:Pareidolia 6421:Allochiria 6404:Multimodal 6285:Parageusia 6280:Hypogeusia 6252:Phantosmia 6232:Hyperosmia 6176:Vestibular 6156:Palinopsia 6116:Amblyaudia 6063:Palinopsia 6048:Nyctalopia 5972:Nociceptor 5779:Artificial 5502:Optic (II) 5333:Perception 5282:Processes 5274:perception 5148:: 135694. 5085:Der Geruch 4697:1 December 4570:4 November 3356:: e07865. 3295:: e07865. 3115:2 November 1784:10 October 1662:Psychology 1565:"to make". 1533:References 1464:SARS-CoV-1 1456:SARS-CoV-2 1448:Osmophobia 1433:Heterosmia 1425:Phantosmia 1395:Hyperosmia 1341:; and the 1276:See also: 1239:Drosophila 1203:pheromones 1171:In insects 1125:pheromones 1018:, and the 857:calmodulin 839:with some 770:antibodies 726:epithelium 665:pheromones 655:. In some 642:catarrhine 588:carnivores 574:, and the 502:turbulence 498:chemotaxis 466:inbreeding 397:Smell and 379:inhalation 375:exhalation 359:trigeminal 311:Luca Turin 145:pheromones 6804:Olfaction 6579:glomeruli 6262:Gustatory 6214:Olfactory 6078:Photopsia 6024:Amaurosis 5998:Disorders 5891:Cone cell 5810:Types of 5421:Inner ear 5356:Cognition 5346:Awareness 5295:Sensation 5270:Sensation 5162:1872-7972 5068:15 August 4986:: 83–89. 4861:207568844 4786:: 31–41. 4622:207090474 4489:1531-5320 4430:1464-3553 4389:14 August 4351:14 August 4333:2352-1546 4319:: 30–39. 4273:0166-2236 4218:1432-0770 4171:0013-4694 3149:14 August 2828:243966350 2820:1947-5454 2303:882238865 2105:: 25103. 1891:218627484 1834:14 August 1695:Neurology 1555:olfactere 1476:olfaction 1443:body odor 1380:Disorders 1272:In plants 1260:aldehydes 985:serotonin 956:glomeruli 835:(largely 821:G protein 764:, salts, 754:dendrites 667:; a male 649:cetaceans 592:ungulates 560:tubenoses 387:forebrain 249:Lucretius 172:Glomeruli 135:, is the 133:olfaction 86:chemicals 48:carnation 6703:Amygdala 6247:Parosmia 6237:Hyposmia 6227:Dysosmia 6166:Tinnitus 6108:Auditory 6083:Polyopia 6039:Diplopia 5896:Rod cell 5694:Nonhuman 5647:Internal 5488:Sensory 5394:External 5303:Stimulus 5286:concepts 5180:33600902 5062:Archived 5031:Archived 5008:29588018 4962:28468812 4921:25649823 4853:26982441 4808:26416146 4756:19372867 4748:11361090 4739:11146473 4688:Archived 4684:30978362 4676:19151709 4626:Archived 4614:23806440 4561:Archived 4557:25086608 4505:43129982 4497:16082803 4448:28641388 4383:Archived 4345:Archived 4341:27989941 4132:22505357 4124:12668289 4086:11418503 4045:19699095 3988:28094328 3980:19554043 3945:18923046 3896:26752161 3847:24672011 3783:27916458 3734:20637621 3677:17724338 3618:25701815 3502:16878137 3448:Archived 3382:26151672 3331:26151672 3270:24653035 3211:23 March 3205:Archived 3174:Archived 3170:BBC News 3109:Archived 3079:Archived 3041:34180521 2992:Archived 2969:22714804 2920:23910657 2863:Archived 2838:13 March 2832:Archived 2771:13 March 2737:17997307 2680:19067327 2631:12810039 2596:42316168 2545:15777804 2510:15621496 2502:16337283 2459:Archived 2436:20181598 2387:11148312 2341:20171264 2256:15034588 2221:12183125 2137:27112241 2080:17603536 2006:12612342 1895:Archived 1883:32413294 1852:(2020). 1828:Archived 1826:: 7–18. 1778:Archived 1766:28504103 1727:25788559 1551:olfactus 1482:See also 1468:MERS-CoV 1419:Parosmia 1413:Dysosmia 1401:Hyposmia 1363:caprylic 1312:sniffing 1247:volatile 1225:antennae 1195:foraging 1157:amygdala 1135:and the 1057:nostrils 1012:amygdala 823:called G 785:sensilla 673:bombykol 669:silkworm 645:primates 572:vultures 536:cilantro 508:Genetics 441:and the 383:thalamus 332:auditory 321:Function 296:molecule 232:, Paris. 160:receptor 81:Function 18:Smelling 6766:General 6416:Agnosia 6295:Tactile 6270:Ageusia 6222:Anosmia 6184:Vertigo 6088:Scotoma 6029:Anopsia 5490:cranial 5361:Feeling 5171:7883672 4988:Bibcode 4912:4317791 4891:Bibcode 4788:Bibcode 4548:4146660 4439:5863551 4289:3947494 4281:2475938 4234:7932310 4226:2551392 4036:2901510 4015:Bibcode 3936:2588668 3887:4948943 3838:3965788 3774:5196021 3725:2929674 3704:Bibcode 3668:1964822 3645:Bibcode 3609:4396857 3569:7724600 3537:Bibcode 3510:2864195 3482:Bibcode 3426:265–275 3397:bioRxiv 3373:4491593 3322:4491593 3261:4483192 3240:Bibcode 3232:Science 3180:20 June 3107:. 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