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Sense of smell

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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.
1015:, 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 1099:. 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. 601:
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.
1091:. 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 291:, 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). 1216:
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
422:; 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. 339:, 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. 1198:
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
611:, 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 1135:
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.
1883: 4298:"Olfactory object recognition, segmentation, adaptation, target seeking, and discrimination by the network of the olfactory bulb and cortex: computational model and experimental data" 271:. 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 2512:
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.
5050: 968:). Granular cells also mediate inhibition and excitation of mitral cells through pathways from centrifugal fibers and the anterior olfactory nuclei. Neuromodulators like 575:
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".
3436: 951:, 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 3193: 1318:
has applied its standard in regulating numerous industries, landfills, and sewage treatment plants. Example applications this district has engaged are the
1144:. 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 306:. However, the behavioral predictions of this theory have been called into question. There is no theory yet that explains olfactory perception completely. 4371: 2820: 2447: 5249: 2851: 4333: 1103:
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
3097: 713:. The olfactory epithelium is made up of at least six morphologically and biochemically different cell types. The proportion of olfactory 4676: 1116:, although pheromones can also be detected by the main olfactory system. In the accessory olfactory system, stimuli are detected by the 6487: 2924:"Genetic variation in the odorant receptor OR2J3 is associated with the ability to detect the "grassy" smelling odor, cis-3-hexen-1-ol" 5042: 879:
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".
1315: 3573:"Trpc2-expressing sensory neurons in the mouse main olfactory epithelium of type B express the soluble guanylate cyclase Gucy1b2" 206: 5242: 1266: 279:
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
6296: 3045:"Odour of King Penguin feathers analysed using direct thermal desorption discriminates between individuals but not sexes" 2972: 1443:
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".
765:). This mucus acts as a solvent for odor molecules, flows constantly, and is replaced approximately every ten minutes. 1730: 5235: 5081: 3162: 2281: 2017: 434:. Functional imaging shows that this olfactory kinship detection process involves the frontal-temporal junction, the 187: 5714: 411: 217: 507:
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
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or through calcium imaging of receptor neuron terminals in the olfactory bulb. In insects, one can perform
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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
132:) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and 6429: 3622:"Contribution of the receptor guanylyl cyclase GC-D to chemosensory function in the olfactory epithelium" 1576: 5968: 1137: 6741: 6572: 6383: 5719: 4810:
Gadenne, C; Barrozo, RB; Anton, S (2016). "Plasticity in insect olfaction: to smell or not to smell?".
1497: 992: 722:. Often, land organisms will have separate olfaction systems for smell and taste (orthonasal smell and 568: 5197: 4297: 2682:
Sherborne, AL; Thom, MD; Paterson, S; Jury, F; Ollier, WE; Stockley, P; Beynon, RJ; Hurst, JL (2007).
6555: 6473: 6192: 6134: 5945: 5301: 3802:"Cholinergic inputs from basal forebrain add an excitatory bias to odor coding in the olfactory bulb" 1327: 1272: 1235: 1218: 1179: 1087:
After binding the odorant, the receptor is activated and will send a signal to the glomeruli in the
1059:, smells are sensed by sensilla located on the antenna and maxillary palp and first processed by the 280: 272: 6605: 6139: 5973: 841: 706: 211: 87: 3044: 855:
This mechanism of transduction is somewhat unusual, in that cAMP works by directly binding to the
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of the olfactory sensory neurons. This may occur by diffusion or by the binding of the odorant to
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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".
6655: 6643: 6633: 6321: 5311: 5291: 4637: 3004:"Highly contiguous genomes improve the understanding of avian olfactory receptor repertoires" 2800: 1319: 1016: 880: 730: 454: 225:
Early scientific study of the sense of smell includes the extensive doctoral dissertation of
5127:"Smell and taste disorders in COVID-19: From pathogenesis to clinical features and outcomes" 4765:"Towards an understanding of the structural basis for insect olfaction by odorant receptors" 3461:
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".
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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
8: 5838: 5823: 5709: 5648: 5584: 5011: 2035:"The scent of life. The exquisite complexity of the sense of smell in animals and humans" 1515: 1300: 1239: 1012: 957: 750: 268: 5212: 4980: 4883: 4780: 4727: 4702: 4007: 3696: 3637: 3529: 3474: 3232: 2890: 2875:"A mendelian trait for olfactory sensitivity affects odor experience and food selection" 2792: 2699: 2193: 2099: 1966: 1843:"What makes a discovery successful? The story of Linda Buck and the olfactory receptors" 247:
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).
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Carraher, C; Dalziel, J; Jordan, MD; Christie, DL; Newcomb, RD; Kralicek, AV (2015).
4732: 4660: 4610: 4598: 4541: 4481: 4473: 4432: 4414: 4361: 4317: 4265: 4257: 4253: 4210: 4202: 4163: 4155: 4151: 4108: 4070: 4029: 3964: 3929: 3880: 3831: 3767: 3718: 3661: 3602: 3553: 3548: 3513: 3486: 3407: 3366: 3315: 3254: 3123: 3025: 2953: 2904: 2816: 2804: 2721: 2664: 2615: 2572: 2568: 2529: 2511: 2486: 2420: 2371: 2325: 2287: 2277: 2240: 2205: 2162: 2121: 2064: 2013: 1990: 1931: 1926: 1909: 1879: 1867: 1750: 1711: 1658: 1596: 1502: 1293: 1222: 1213: 1171: 1117: 1049: 1008: 940: 801: 649: 548: 439: 328: 230: 195: 4744: 4672: 4594: 4493: 4329: 4120: 3976: 3900:"Noradrenergic induction of odor-specific neural habituation and olfactory memories" 2584: 2498: 6766: 6695: 6675: 6650: 6520: 6515: 6164: 6109: 6041: 5858: 5801: 5704: 5699: 5609: 5558: 5553: 5515: 5462: 5457: 5154: 5138: 4984: 4967:
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".
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phase of breathing. The olfactory system is the only human sense that bypasses the
347: 332: 295: 156: 63: 3094:"Dogs' dazzling sense of smell: what lies behind their exceptional gift of sniff?" 1680:"Olfactory impairment & traumatic brain injury in blast-injured combat troops" 6628: 6579: 6542: 6081: 6022: 5994: 5810: 5785: 5780: 5775: 5734: 5543: 5500: 5447: 5142: 4313: 4091:
Leon, M; Johnson, BA (2003). "Olfactory coding in the mammalian olfactory bulb".
3753: 3514:"A receptor guanylyl cyclase expressed specifically in olfactory sensory neurons" 2321: 1696: 1645: 1507: 1476: 1430:– psychological disorder that causes the patient to imagine he or she has strong 1244: 1205: 1068: 1004: 909: 474: 462: 443: 92: 2632: 2525: 1678:
Xydakis, MS; Mulligan, LP; Smith, AB; Olsen, CH; Lyon, DM; Belluscio, L (2015).
6562: 6399: 6250: 6202: 5937: 5917: 5643: 5530: 5478: 5434: 5296: 4988: 4788: 4509:"An olfactory cocktail party: figure-ground segregation of odorants in rodents" 3626:
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
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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).
4065: 4048: 4015: 3738:"A sensor for low environmental oxygen in the mouse main olfactory epithelium" 3704: 3588: 2899: 2874: 2753: 2707: 2157: 2140: 1033:. This is possibly due to the olfactory system's close anatomical ties to the 652:, scent glands occur atop the head. In many species, smell is highly tuned to 607:
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
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Reddy, Gautam; Murthy, Venkatesh N.; Vergassola, Massimo (10 March 2022).
2576: 2166: 1935: 394:. This neural convergence is proposed to give rise to a perception termed 6532: 6434: 6342: 6316: 6301: 6263: 6086: 6061: 6056: 6046: 6032: 5658: 5614: 5490: 5419: 4635: 2597: 1677: 1395: 1234:
Insects are capable of smelling and differentiating between thousands of
1195: 1092: 1021: 948: 895: 856: 564: 336: 335:
plays the biggest role in the sensation of flavor. During the process of
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Specific terms are used to describe disorders associated with smelling:
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pathway, depending on the organism. In mammals, the odorants stimulate
758: 714: 702: 630: 608: 604: 490: 486: 367: 363: 299: 4925:"The insect ovipositor as a volatile sensor within a closed microcosm" 3063: 2650: 2107: 519:, a key aroma in foods and beverages. Similarly, the odorant receptor 442:, but not the primary or secondary olfactory cortices, or the related 6066: 6012: 5879: 5409: 5399: 5344: 5334: 5207: 4182: 4136:"Spatial properties of an EEG event in the olfactory bulb and cortex" 3800:
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1629: 1464: 1431: 1191: 1113: 973: 809: 772:, smells are sensed by olfactory sensory neurons in the chemosensory 742: 653: 576: 556: 485:
have been studied in different species, including gradient search or
390:
information has been shown to converge in the olfactory tubercles of
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Mastrangelo, Andrea; Bonato, Matteo; Cinque, Paola (23 March 2021).
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4527: 4183:"Modeling the olfactory bulb and its neural oscillatory processings" 3866: 3290: 939:
converge in the outer layer of the olfactory bulb within small (≈50
619:, which was bred to track and hunt rabbits and other small animals. 593:
in contrast to having a weak sense of smell which is referred to as
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a few days old. The second-most-sensitive nose is possessed by the
580: 560: 524: 371: 284: 4238:"Olfactory cortex: model circuit for study of associative memory?" 2635:"The neuronal substrates of human olfactory based kin recognition" 1804: 737:
that lines the superior portion of the cavity and are detected by
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2440:"Making scents of sounds: noises may alter how we perceive odors" 2307: 1377: 1212:
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
1805:"Lucretius, the biochemistry of smell, and scientific discovery" 457:
is detrimental, it tends to be avoided. In the house mouse, the
163:
aggregate signals from these receptors and transmit them to the
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The tendrils of plants are especially sensitive to airborne
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in the environment that are used to form the sense of smell
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can be utilized to determine the magnitude of an odor.
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Olfactory Cortex - an overview | ScienceDirect Topics
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Morris' Human Anatomy: A Complete Systematic Treatise
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Plant perception (physiology) § Senses in plants
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Electroencephalography and Clinical Neurophysiology
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The axons from the 155:, transmitting a signal through the 4374:from the original on 14 August 2022 4336:from the original on 14 August 2022 3577:Molecular and Cellular Neuroscience 3272:Meister, Markus (1 November 2014). 3008:Integrative and Comparative Biology 3002:Driver, R; Balakrishnan, C (2021). 2854:from the original on 8 October 2017 2082:Saberi, M; Seyed-allaei, H (2016). 1788:Kimble, GA; Schlesinger, K (1985). 1404:– distortion in the sense of smell 1150:bed nucleus of the stria terminalis 818:cyclic nucleotide-gated ion channel 468: 201: 13: 5448:Auditory system (sense of hearing) 5053:from the original on 17 April 2021 4957: 4856: 4824:10.1146/annurev-ento-010715-023523 4284: 3073:from the original on 26 April 2019 2983:from the original on 29 March 2021 2450:from the original on 19 March 2011 2298: 2260: 1729:Xydakis, MS; Belluscio, L (2017). 1609: 1584: 1569: 1451:, including anosmia and parosmia. 1322:, wastewater treatment plant; the 904:Olfactory sensory neurons project 859:rather than through activation of 783: 686:Physiological basis in vertebrates 14: 6814: 5463:Gustatory system (sense of taste) 5458:Olfactory system (sense of smell) 5179: 5076:(in German). 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These nerve fibers, lacking 867:, in which the second messenger 559:), certain species of new world 5443:Visual system (sense of vision) 5118: 5065: 5034: 5003: 4929:Journal of Experimental Biology 4916: 4694: 4629: 4595:10.1016/j.neubiorev.2013.06.009 4500: 4443: 4386: 4348: 4296:Zhaoping, Li (1 October 2016). 4229: 4174: 4127: 4040: 3983: 3940: 3891: 3842: 3793: 3778: 3729: 3672: 3613: 3564: 3505: 3454: 3421: 3398: 3377: 3326: 3265: 3208: 3184:Williams, SCP (20 March 2014). 3177: 3146: 3122:Padodara, RJ; Ninan, J (2014). 3085: 3036: 2995: 2964: 2835: 2741:"How animals follow their nose" 2626: 2591: 2548: 2505: 2462: 2438:Peeples, L (23 February 2010). 2389:Wesson, DW; Wilson, DA (2010). 2382: 2216: 2173: 2132: 2075: 2026: 2001: 1942: 1901: 1831: 1819:from the original on 8 May 2022 943:in diameter) structures called 535:Variability amongst vertebrates 35:Painting of a woman smelling a 5010:Fountain, H (3 October 2006). 3916:10.1523/JNEUROSCI.3853-08.2008 3818:10.1523/JNEUROSCI.5026-13.2014 2407:10.1523/JNEUROSCI.6003-09.2010 2368:10.1146/annurev.psych.52.1.423 2354:Doty, RL (2001). 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Erlbaum Associates. 1221:typically reside in the 1152:, and from there to the 1138:accessory olfactory bulb 908:to the brain within the 842:sodium-calcium exchanger 747:odorant-binding proteins 314: 212:The Lady and the Unicorn 6716:Medial forebrain bundle 6614:Lateral olfactory stria 6597:areas involved in smell 6120:Auditory verbal agnosia 5974:Juxtacapillary receptor 5213:Olfaction and Gustation 4242:Trends in Neurosciences 4066:10.1093/chemse/26.5.577 3904:Journal of Neuroscience 3806:Journal of Neuroscience 3647:10.1073/pnas.0704965104 3241:10.1126/science.1249168 2471:Trends in Neurosciences 2395:Journal of Neuroscience 2158:10.1093/chemse/21.6.773 1976:10.1073/pnas.0535697100 1178:to detect and identify 989:lateral olfactory tract 836:, leading to efflux of 705:, smells are sensed by 350:receptors (also called 6772:Evolution of olfaction 6734:Medial olfactory stria 6115:Auditory hallucination 5725:Surface wave detection 5330:Multimodal integration 4187:Biological Cybernetics 4093:Brain Research Reviews 3539:10.1073/pnas.92.8.3571 3428:Boroditsky, L (1999). 2202:10.1006/jtbi.2001.2504 1550:"to emit a smell" and 1539: 1493:Olfactic Communication 1482:Evolution of olfaction 1324:Shoreline Amphitheatre 1288:Machine-based smelling 901: 503:Evolution of olfaction 438:, and the dorsomedial 263:have about a thousand 255:(who were awarded the 222: 192:traumatic brain injury 136:, and plays a role in 100:Anatomical terminology 6656:Periamygdaloid cortex 6644:Hippocampal formation 6634:EC-hippocampus system 6374:Phantom limb syndrome 6322:Tactile hallucination 5511:Glossopharyngeal (IX) 5312:Active sensory system 5198:Olfaction at cf.ac.uk 5193:at Wikimedia Commons 4411:10.1093/chemse/bjx031 3190:Science Now/AAAS News 2940:10.1093/chemse/bjs049 2514:Hormones and Behavior 2272:Shepherd, GM (2013). 1803:Holtsmark, E (1978). 1542:, past participle of 1320:San Mateo, California 1079:Coding and perception 1017:medial dorsal nucleus 923:sheaths, pass to the 898: 778:Chemosensory proteins 731:superior nasal concha 691:Main olfactory system 672:Human smell abilities 636:, and nonexistent in 459:major urinary protein 209: 6183:Labyrinthine fistula 6150:Spatial hearing loss 5849:Campaniform sensilla 5564:Somatosensory cortex 5131:Neuroscience Letters 4701:Brennan, PA (2001). 4577:Auffarth, B (2013). 3736:Bleymehl, K (2016). 2310:Neuroscience Letters 1735:The Lancet Neurology 1487:Nasal administration 1380:– inability to smell 1362:Hendrik Zwaardemaker 1336:Martinez, California 1275:. Parasites such as 1044:Inputs from the two 958:periglomerular cells 885:electroantennography 711:olfactory epithelium 701:In humans and other 448:orbitofrontal cortex 406:Inbreeding avoidance 182:or smell receptors, 6803:Otorhinolaryngology 5969:Nociceptin receptor 5839:Merkel nerve ending 5824:Mechanotransduction 5217:Neuroscience Online 5100:Hawkes, CH (2002). 5072:Henning, H (1915). 4981:2015COIS...10...83S 4884:2015CBio...25.R120L 4781:2015IBMB...66...31C 4645:Nature Neuroscience 4516:Nature Neuroscience 4047:Wilson, DA (2001). 4008:2009CBio...19.1561Z 3949:Nature Neuroscience 3910:(42): 10711–10719. 3855:Nature Neuroscience 3697:2010CBio...20.1438M 3638:2007PNAS..10414507L 3632:(36): 14507–14512. 3530:1995PNAS...92.3571F 3483:10.1038/nature05066 3475:2006Natur.442..645L 3352:10.7554/eLife.07865 3333:Meister, M (2015). 3301:10.7554/eLife.07865 3233:2014Sci...343.1370B 3227:(6177): 1370–1372. 3021:10.1093/icb/icab150 2891:2013CBio...23.1601J 2793:2022ARCMP..13..191R 2700:2007CBio...17.2061S 2639:Human Brain Mapping 2444:Scientific American 2225:Nature Neuroscience 2194:2002JThBi.216..367T 2100:2016NatSR...625103S 2033:Rinaldi, A (2007). 2008:Pinel, JPJ (2006). 1967:2003PNAS..100.3324G 1516:Scent transfer unit 1463:) can also disrupt 1067:), and next by the 1013:anterior commissure 937:olfactory receptors 881:electro-olfactogram 788:The binding of the 751:mucopolysaccharides 739:olfactory receptors 547:do not, except the 231:stimulus modalities 6551:Olfactory receptor 6528:Sustentacular cell 6331:Nociception (pain) 5923:Olfactory receptor 5875:Photoreceptor cell 5829:Lamellar corpuscle 5753:Photomorphogenesis 5615:nociception (pain) 5307:Sensory processing 5016:The New York Times 4942:10.1242/jeb.152777 4719:10.1007/pl00000880 4469:10.3758/BF03196369 4199:10.1007/BF00200803 3961:10.1038/nn0709-813 3679:Munger, S (2010). 3512:Fulle, HJ (1995). 2747:. Annual Reviews. 2088:Scientific Reports 1839:Ann-Sophie Barwich 1489:olfactory transfer 1459:and even the flu ( 1254:Deilephila elpenor 1180:volatile compounds 1172:chemical receptors 1063:(analogous to the 997:olfactory tubercle 966:lateral inhibition 902: 874:lateral inhibition 432:Westermarck effect 304:quantum tunnelling 223: 6780: 6779: 6750: 6749: 6728: 6727: 6639:Entorhinal cortex 6585:Olfactory trigone 6463: 6462: 6448:Biases and errors 6443: 6442: 6379:Somatoparaphrenia 6348:Pain dissociation 6193:Ménière's disease 6125:Cortical deafness 6003:Visual impairment 5982: 5981: 5844:Bulbous corpuscle 5834:Tactile corpuscle 5802:sensory receptors 5794: 5793: 5677: 5676: 5630: 5629: 5595:Olfaction (smell) 5549:Vestibular cortex 5531:Cerebral cortices 5368: 5367: 5355:Motion perception 5189:Media related to 5111:978-0-7506-7287-0 4367:978-0-08-045046-9 4002:(18): 1561–1565. 3812:(13): 4654–4664. 3691:(16): 1438–1444. 3571:Omura, M (2015). 3469:(7103): 645–650. 3064:10.1111/ibi.12544 2885:(16): 1601–1605. 2745:Knowable Magazine 2694:(23): 2061–2066. 2651:10.1002/hbm.20686 2139:Turin, L (1996). 2108:10.1038/srep25103 1664:978-1-4292-3719-2 1602:978-0-87893-572-7 1570:Harper, Douglas. 1503:Olfactory fatigue 1294:Machine olfaction 1118:vomeronasal organ 1050:binocular rivalry 1009:entorhinal cortex 802:adenylate cyclase 650:red-bellied lemur 473:Some animals use 440:prefrontal cortex 114: 113: 109: 39:. Olfaction uses 6810: 6767:Olfactory system 6696:Stria terminalis 6676:Habenular nuclei 6672:Stria medullaris 6651:Prepyriform area 6619: 6618: 6601: 6600: 6490: 6483: 6476: 6467: 6466: 6110:Auditory agnosia 6042:Optic neuropathy 5991: 5990: 5859:Stretch receptor 5807: 5806: 5705:Magnetoreception 5700:Electroreception 5688: 5687: 5610:mechanoreception 5559:Gustatory cortex 5554:Olfactory cortex 5388: 5387: 5379: 5378: 5297:Sensory receptor 5280: 5279: 5252: 5245: 5238: 5229: 5228: 5188: 5173: 5172: 5162: 5122: 5116: 5115: 5097: 5088: 5087: 5069: 5063: 5062: 5060: 5058: 5047:Nadia Berenstein 5038: 5032: 5031: 5029: 5027: 5007: 5001: 5000: 4964: 4955: 4954: 4944: 4935:(9): 1554–1557. 4920: 4914: 4913: 4903: 4878:(3): R120–R129. 4863: 4854: 4853: 4835: 4807: 4801: 4800: 4760: 4749: 4748: 4730: 4698: 4692: 4691: 4689: 4687: 4681: 4642: 4633: 4627: 4626: 4624: 4622: 4589:(8): 1667–1679. 4574: 4565: 4564: 4562: 4560: 4554: 4539: 4522:(9): 1225–1232. 4513: 4504: 4498: 4497: 4471: 4447: 4441: 4440: 4430: 4390: 4384: 4383: 4381: 4379: 4352: 4346: 4345: 4343: 4341: 4293: 4282: 4281: 4233: 4227: 4226: 4178: 4172: 4171: 4131: 4125: 4124: 4088: 4079: 4078: 4068: 4044: 4038: 4037: 4027: 3987: 3981: 3980: 3944: 3938: 3937: 3927: 3895: 3889: 3888: 3878: 3846: 3840: 3839: 3829: 3797: 3791: 3790: 3782: 3776: 3775: 3765: 3748:(6): 1196–1203. 3733: 3727: 3726: 3716: 3676: 3670: 3669: 3659: 3649: 3617: 3611: 3610: 3600: 3568: 3562: 3561: 3551: 3541: 3524:(8): 3571–3575. 3509: 3503: 3502: 3458: 3452: 3451: 3449: 3447: 3441: 3434: 3425: 3419: 3418: 3412: 3402: 3396: 3395: 3393: 3381: 3375: 3374: 3364: 3354: 3330: 3324: 3323: 3313: 3303: 3293: 3269: 3263: 3262: 3252: 3212: 3206: 3205: 3203: 3201: 3181: 3175: 3174: 3172: 3170: 3150: 3144: 3143: 3141: 3139: 3119: 3110: 3109: 3107: 3105: 3089: 3083: 3082: 3080: 3078: 3072: 3049: 3040: 3034: 3033: 3023: 3014:(4): 1281–1290. 2999: 2993: 2992: 2990: 2988: 2968: 2962: 2961: 2951: 2919: 2913: 2912: 2902: 2870: 2864: 2863: 2861: 2859: 2839: 2833: 2832: 2830: 2828: 2772: 2766: 2765: 2763: 2761: 2756: 2736: 2730: 2729: 2719: 2679: 2673: 2672: 2662: 2645:(8): 2571–2580. 2630: 2624: 2623: 2595: 2589: 2588: 2552: 2546: 2545: 2509: 2503: 2502: 2466: 2460: 2459: 2457: 2455: 2435: 2429: 2428: 2418: 2401:(8): 3013–3021. 2386: 2380: 2379: 2351: 2342: 2341: 2305: 2296: 2295: 2269: 2258: 2256: 2220: 2214: 2213: 2177: 2171: 2170: 2160: 2136: 2130: 2129: 2119: 2079: 2073: 2072: 2062: 2030: 2024: 2023: 2005: 1999: 1998: 1988: 1978: 1961:(6): 3324–3327. 1946: 1940: 1939: 1929: 1905: 1899: 1898: 1896: 1894: 1888: 1865: 1847: 1835: 1829: 1828: 1826: 1824: 1800: 1794: 1793: 1785: 1779: 1778: 1776: 1774: 1726: 1720: 1719: 1709: 1699: 1675: 1669: 1668: 1648: 1640: 1634: 1633: 1630:10.1002/ffj.3249 1613: 1607: 1606: 1588: 1582: 1581: 1567: 1555: 1532: 1245:carboxylic acids 1166:Insect olfaction 1097:olfactory cortex 1031:emotional memory 1027:long-term memory 956:also project to 933:olfactory cortex 929:cribriform plate 918: 861:protein kinase A 834:chloride channel 798:second messenger 794:action potential 724:retronasal smell 697:Olfactory system 680: 509:odorant receptor 469:Guiding movement 428:incest avoidance 414:genes (known as 333:Retronasal smell 296:vibration theory 243: 202:History of study 157:olfactory system 106:edit on Wikidata 103: 64:Olfactory system 33: 21: 20: 6818: 6817: 6813: 6812: 6811: 6809: 6808: 6807: 6783: 6782: 6781: 6776: 6746: 6724: 6704: 6680: 6660: 6629:Piriform cortex 6608: 6589: 6580:Olfactory tract 6543:Olfactory nerve 6537: 6499: 6494: 6464: 6459: 6439: 6388: 6357: 6326: 6278: 6245: 6197: 6159: 6091: 6082:Stereoblindness 6023:Color blindness 5978: 5955: 5932: 5904: 5863: 5811:Mechanoreceptor 5800: 5790: 5786:Machine hearing 5781:Computer vision 5776:Robotic sensing 5762: 5739: 5673: 5626: 5568: 5544:Auditory cortex 5525: 5472: 5435:Sensory systems 5429: 5364: 5316: 5274: 5272: 5265: 5256: 5208:Smells Database 5182: 5177: 5176: 5123: 5119: 5112: 5098: 5091: 5084: 5070: 5066: 5056: 5054: 5039: 5035: 5025: 5023: 5008: 5004: 4965: 4958: 4921: 4917: 4872:Current Biology 4864: 4857: 4808: 4804: 4761: 4752: 4699: 4695: 4685: 4683: 4679: 4657:10.1038/nn.2262 4640: 4634: 4630: 4620: 4618: 4575: 4568: 4558: 4556: 4552: 4528:10.1038/nn.3775 4511: 4505: 4501: 4448: 4444: 4399:Chemical Senses 4391: 4387: 4377: 4375: 4368: 4354: 4353: 4349: 4339: 4337: 4294: 4285: 4234: 4230: 4179: 4175: 4132: 4128: 4089: 4082: 4053:Chemical Senses 4045: 4041: 3996:Current Biology 3988: 3984: 3945: 3941: 3896: 3892: 3867:10.1038/nn.4219 3847: 3843: 3798: 3794: 3783: 3779: 3734: 3730: 3685:Current Biology 3677: 3673: 3618: 3614: 3569: 3565: 3510: 3506: 3459: 3455: 3445: 3443: 3439: 3432: 3426: 3422: 3403: 3399: 3382: 3378: 3331: 3327: 3270: 3266: 3213: 3209: 3199: 3197: 3182: 3178: 3168: 3166: 3151: 3147: 3137: 3135: 3120: 3113: 3103: 3101: 3090: 3086: 3076: 3074: 3070: 3047: 3041: 3037: 3000: 2996: 2986: 2984: 2969: 2965: 2928:Chemical Senses 2920: 2916: 2879:Current Biology 2871: 2867: 2857: 2855: 2840: 2836: 2826: 2824: 2773: 2769: 2759: 2757: 2737: 2733: 2688:Current Biology 2680: 2676: 2631: 2627: 2596: 2592: 2553: 2549: 2510: 2506: 2467: 2463: 2453: 2451: 2436: 2432: 2387: 2383: 2352: 2345: 2306: 2299: 2284: 2270: 2261: 2221: 2217: 2178: 2174: 2145:Chemical Senses 2137: 2133: 2080: 2076: 2031: 2027: 2020: 2006: 2002: 1947: 1943: 1906: 1902: 1892: 1890: 1886: 1845: 1836: 1832: 1822: 1820: 1801: 1797: 1786: 1782: 1772: 1770: 1727: 1723: 1690:(15): 1559–67. 1676: 1672: 1665: 1641: 1637: 1614: 1610: 1603: 1589: 1585: 1568: 1564: 1559: 1558: 1533: 1529: 1524: 1477:Electronic nose 1473: 1461:influenza virus 1371: 1357:Henning's prism 1344: 1296: 1290: 1269: 1263: 1238:compounds both 1206:Polybia sericea 1194:) and locating 1168: 1162: 1109: 1081: 1069:mushroom bodies 1005:piriform cortex 916: 910:olfactory nerve 893: 815: 786: 784:Receptor neuron 699: 693: 688: 678: 674: 537: 505: 499: 471: 463:kin recognition 444:piriform cortex 408: 384: 317: 312: 241: 204: 110: 48: 17: 12: 11: 5: 6816: 6806: 6805: 6800: 6795: 6778: 6777: 6775: 6774: 6769: 6764: 6758: 6756: 6752: 6751: 6748: 6747: 6745: 6744: 6738: 6736: 6730: 6729: 6726: 6725: 6723: 6722: 6712: 6710: 6706: 6705: 6703: 6702: 6688: 6686: 6682: 6681: 6679: 6678: 6668: 6666: 6662: 6661: 6659: 6658: 6653: 6648: 6647: 6646: 6641: 6631: 6625: 6623: 6616: 6610: 6609: 6604: 6598: 6591: 6590: 6588: 6587: 6582: 6577: 6576: 6575: 6570: 6563:Olfactory bulb 6560: 6559: 6558: 6547: 6545: 6539: 6538: 6536: 6535: 6530: 6525: 6524: 6523: 6518: 6507: 6505: 6501: 6500: 6493: 6492: 6485: 6478: 6470: 6461: 6460: 6458: 6457: 6451: 6449: 6445: 6444: 6441: 6440: 6438: 6437: 6432: 6427: 6422: 6417: 6412: 6407: 6402: 6396: 6394: 6390: 6389: 6387: 6386: 6381: 6376: 6371: 6365: 6363: 6362:Proprioception 6359: 6358: 6356: 6355: 6350: 6345: 6340: 6334: 6332: 6328: 6327: 6325: 6324: 6319: 6314: 6309: 6304: 6299: 6294: 6288: 6286: 6280: 6279: 6277: 6276: 6271: 6266: 6261: 6255: 6253: 6247: 6246: 6244: 6243: 6238: 6233: 6228: 6223: 6218: 6213: 6207: 6205: 6199: 6198: 6196: 6195: 6190: 6185: 6180: 6175: 6169: 6167: 6161: 6160: 6158: 6157: 6152: 6147: 6142: 6137: 6132: 6127: 6122: 6117: 6112: 6107: 6101: 6099: 6093: 6092: 6090: 6089: 6084: 6079: 6074: 6069: 6064: 6059: 6054: 6049: 6044: 6039: 6030: 6025: 6020: 6015: 6010: 6005: 5999: 5997: 5988: 5984: 5983: 5980: 5979: 5977: 5976: 5971: 5965: 5963: 5957: 5956: 5954: 5953: 5948: 5942: 5940: 5938:Thermoreceptor 5934: 5933: 5931: 5930: 5925: 5920: 5918:Taste receptor 5914: 5912: 5906: 5905: 5903: 5902: 5897: 5892: 5887: 5882: 5877: 5871: 5869: 5865: 5864: 5862: 5861: 5856: 5851: 5846: 5841: 5836: 5831: 5826: 5821: 5815: 5813: 5804: 5796: 5795: 5792: 5791: 5789: 5788: 5783: 5778: 5772: 5770: 5764: 5763: 5761: 5760: 5755: 5749: 5747: 5741: 5740: 5738: 5737: 5732: 5727: 5722: 5717: 5712: 5707: 5702: 5696: 5694: 5685: 5679: 5678: 5675: 5674: 5672: 5671: 5666: 5661: 5656: 5651: 5646: 5644:Proprioception 5640: 5638: 5632: 5631: 5628: 5627: 5625: 5624: 5623: 5622: 5617: 5612: 5602: 5597: 5592: 5587: 5582: 5576: 5574: 5570: 5569: 5567: 5566: 5561: 5556: 5551: 5546: 5541: 5535: 5533: 5527: 5526: 5524: 5523: 5518: 5516:Trigeminal (V) 5513: 5508: 5503: 5498: 5493: 5487: 5485: 5474: 5473: 5471: 5470: 5465: 5460: 5455: 5450: 5445: 5439: 5437: 5431: 5430: 5428: 5427: 5422: 5417: 5412: 5407: 5402: 5396: 5394: 5392:Sensory organs 5385: 5376: 5370: 5369: 5366: 5365: 5363: 5362: 5357: 5352: 5347: 5342: 5337: 5332: 5326: 5324: 5318: 5317: 5315: 5314: 5309: 5304: 5299: 5294: 5288: 5286: 5277: 5267: 5266: 5255: 5254: 5247: 5240: 5232: 5226: 5225: 5220: 5210: 5205: 5200: 5181: 5180:External links 5178: 5175: 5174: 5117: 5110: 5089: 5082: 5064: 5033: 5002: 4956: 4915: 4855: 4802: 4750: 4713:(4): 546–555. 4693: 4651:(2): 210–220. 4628: 4566: 4499: 4462:(2): 244–264. 4442: 4405:(7): 537–546. 4385: 4366: 4347: 4283: 4248:(7): 258–264. 4228: 4193:(5): 379–392. 4173: 4146:(5): 586–605. 4126: 4080: 4059:(5): 577–584. 4039: 3982: 3955:(7): 813–814. 3939: 3890: 3861:(2): 813–814. 3841: 3792: 3777: 3728: 3671: 3612: 3563: 3504: 3453: 3420: 3397: 3391:10.1101/022103 3376: 3325: 3264: 3207: 3176: 3145: 3111: 3084: 3058:(2): 379–389. 3035: 2994: 2963: 2934:(7): 585–593. 2914: 2865: 2834: 2787:(1): 191–213. 2767: 2731: 2674: 2625: 2606:(3): 279–295. 2590: 2563:(3): 445–448. 2547: 2520:(4): 384–388. 2504: 2477:(2): 100–107. 2461: 2430: 2381: 2362:(1): 423–452. 2343: 2297: 2282: 2259: 2237:10.1038/nn1215 2231:(4): 337–338. 2215: 2188:(3): 367–385. 2172: 2151:(6): 773–791. 2131: 2074: 2025: 2018: 2000: 1941: 1920:(1): 175–187. 1900: 1856:(4): 749–753. 1830: 1795: 1780: 1741:(6): 415–416. 1721: 1670: 1663: 1635: 1624:(5): 342–361. 1608: 1601: 1583: 1561: 1560: 1557: 1556: 1526: 1525: 1523: 1520: 1519: 1518: 1513: 1505: 1500: 1495: 1490: 1484: 1479: 1472: 1469: 1441: 1440: 1434: 1425: 1419: 1418: 1417: 1411: 1399: 1393: 1387: 1381: 1370: 1367: 1366: 1365: 1358: 1355: 1343: 1342:Classification 1340: 1332:IT Corporation 1292:Main article: 1289: 1286: 1262: 1259: 1190:partners (via 1164:Main article: 1161: 1158: 1142:olfactory bulb 1108: 1105: 1089:olfactory bulb 1080: 1077: 1065:olfactory bulb 978:norepinephrine 962:granular cells 925:olfactory bulb 892: 889: 865:photoreceptors 813: 804:to synthesize 785: 782: 695:Main article: 692: 689: 687: 684: 673: 670: 648:, such as the 646:strepsirrhines 536: 533: 501:Main article: 498: 495: 470: 467: 407: 404: 383: 380: 316: 313: 311: 308: 289:odotope theory 269:odor reception 267:that code for 227:Eleanor Gamble 203: 200: 165:olfactory bulb 118:sense of smell 112: 111: 102: 96: 95: 90: 84: 83: 79: 78: 71: 67: 66: 61: 55: 54: 50: 49: 41:chemoreceptors 34: 26: 25: 15: 9: 6: 4: 3: 2: 6815: 6804: 6801: 6799: 6798:Limbic system 6796: 6794: 6791: 6790: 6788: 6773: 6770: 6768: 6765: 6763: 6760: 6759: 6757: 6753: 6743: 6740: 6739: 6737: 6735: 6731: 6721: 6717: 6714: 6713: 6711: 6707: 6701: 6697: 6693: 6690: 6689: 6687: 6683: 6677: 6673: 6670: 6669: 6667: 6663: 6657: 6654: 6652: 6649: 6645: 6642: 6640: 6637: 6636: 6635: 6632: 6630: 6627: 6626: 6624: 6620: 6617: 6615: 6611: 6607: 6602: 6599: 6596: 6592: 6586: 6583: 6581: 6578: 6574: 6571: 6569: 6566: 6565: 6564: 6561: 6557: 6554: 6553: 6552: 6549: 6548: 6546: 6544: 6540: 6534: 6531: 6529: 6526: 6522: 6519: 6517: 6514: 6513: 6512: 6509: 6508: 6506: 6502: 6498: 6491: 6486: 6484: 6479: 6477: 6472: 6471: 6468: 6456: 6453: 6452: 6450: 6446: 6436: 6433: 6431: 6428: 6426: 6423: 6421: 6420:Hallucination 6418: 6416: 6415:Derealization 6413: 6411: 6408: 6406: 6403: 6401: 6398: 6397: 6395: 6391: 6385: 6382: 6380: 6377: 6375: 6372: 6370: 6369:Asomatognosia 6367: 6366: 6364: 6360: 6354: 6351: 6349: 6346: 6344: 6341: 6339: 6336: 6335: 6333: 6329: 6323: 6320: 6318: 6315: 6313: 6310: 6308: 6307:Hyperesthesia 6305: 6303: 6300: 6298: 6295: 6293: 6292:Astereognosis 6290: 6289: 6287: 6285: 6281: 6275: 6272: 6270: 6267: 6265: 6262: 6260: 6257: 6256: 6254: 6252: 6248: 6242: 6239: 6237: 6234: 6232: 6229: 6227: 6224: 6222: 6219: 6217: 6214: 6212: 6209: 6208: 6206: 6204: 6200: 6194: 6191: 6189: 6188:Labyrinthitis 6186: 6184: 6181: 6179: 6176: 6174: 6171: 6170: 6168: 6166: 6162: 6156: 6153: 6151: 6148: 6146: 6143: 6141: 6138: 6136: 6133: 6131: 6128: 6126: 6123: 6121: 6118: 6116: 6113: 6111: 6108: 6106: 6103: 6102: 6100: 6098: 6094: 6088: 6085: 6083: 6080: 6078: 6075: 6073: 6070: 6068: 6065: 6063: 6060: 6058: 6055: 6053: 6050: 6048: 6045: 6043: 6040: 6038: 6034: 6031: 6029: 6026: 6024: 6021: 6019: 6016: 6014: 6011: 6009: 6006: 6004: 6001: 6000: 5998: 5996: 5992: 5989: 5985: 5975: 5972: 5970: 5967: 5966: 5964: 5962: 5958: 5952: 5949: 5947: 5944: 5943: 5941: 5939: 5935: 5929: 5926: 5924: 5921: 5919: 5916: 5915: 5913: 5911: 5910:Chemoreceptor 5907: 5901: 5898: 5896: 5893: 5891: 5888: 5886: 5883: 5881: 5878: 5876: 5873: 5872: 5870: 5868:Photoreceptor 5866: 5860: 5857: 5855: 5854:Slit sensilla 5852: 5850: 5847: 5845: 5842: 5840: 5837: 5835: 5832: 5830: 5827: 5825: 5822: 5820: 5817: 5816: 5814: 5812: 5808: 5805: 5803: 5797: 5787: 5784: 5782: 5779: 5777: 5774: 5773: 5771: 5769: 5765: 5759: 5756: 5754: 5751: 5750: 5748: 5746: 5742: 5736: 5733: 5731: 5728: 5726: 5723: 5721: 5718: 5716: 5713: 5711: 5708: 5706: 5703: 5701: 5698: 5697: 5695: 5693: 5689: 5686: 5684: 5680: 5670: 5669:Visceral pain 5667: 5665: 5662: 5660: 5657: 5655: 5652: 5650: 5647: 5645: 5642: 5641: 5639: 5637: 5633: 5621: 5620:thermoception 5618: 5616: 5613: 5611: 5608: 5607: 5606: 5603: 5601: 5598: 5596: 5593: 5591: 5588: 5586: 5583: 5581: 5578: 5577: 5575: 5571: 5565: 5562: 5560: 5557: 5555: 5552: 5550: 5547: 5545: 5542: 5540: 5539:Visual cortex 5537: 5536: 5534: 5532: 5528: 5522: 5519: 5517: 5514: 5512: 5509: 5507: 5504: 5502: 5501:Olfactory (I) 5499: 5497: 5494: 5492: 5489: 5488: 5486: 5484: 5483:spinal nerves 5480: 5475: 5469: 5466: 5464: 5461: 5459: 5456: 5454: 5451: 5449: 5446: 5444: 5441: 5440: 5438: 5436: 5432: 5426: 5423: 5421: 5418: 5416: 5413: 5411: 5408: 5406: 5403: 5401: 5398: 5397: 5395: 5393: 5389: 5386: 5384: 5380: 5377: 5375: 5371: 5361: 5358: 5356: 5353: 5351: 5348: 5346: 5343: 5341: 5340:Consciousness 5338: 5336: 5333: 5331: 5328: 5327: 5325: 5323: 5319: 5313: 5310: 5308: 5305: 5303: 5300: 5298: 5295: 5293: 5290: 5289: 5287: 5285: 5281: 5278: 5276: 5268: 5264: 5260: 5253: 5248: 5246: 5241: 5239: 5234: 5233: 5230: 5224: 5221: 5218: 5214: 5211: 5209: 5206: 5204: 5201: 5199: 5196: 5195: 5194: 5192: 5187: 5170: 5166: 5161: 5156: 5152: 5148: 5144: 5140: 5136: 5132: 5128: 5121: 5113: 5107: 5103: 5096: 5094: 5085: 5083:9783750176669 5079: 5075: 5068: 5052: 5048: 5044: 5037: 5021: 5017: 5013: 5006: 4998: 4994: 4990: 4986: 4982: 4978: 4974: 4970: 4963: 4961: 4952: 4948: 4943: 4938: 4934: 4930: 4926: 4919: 4911: 4907: 4902: 4897: 4893: 4889: 4885: 4881: 4877: 4873: 4869: 4862: 4860: 4851: 4847: 4843: 4839: 4834: 4829: 4825: 4821: 4817: 4813: 4806: 4798: 4794: 4790: 4786: 4782: 4778: 4774: 4770: 4766: 4759: 4757: 4755: 4746: 4742: 4738: 4734: 4729: 4724: 4720: 4716: 4712: 4708: 4704: 4697: 4678: 4674: 4670: 4666: 4662: 4658: 4654: 4650: 4646: 4639: 4632: 4616: 4612: 4608: 4604: 4600: 4596: 4592: 4588: 4584: 4580: 4573: 4571: 4551: 4547: 4543: 4538: 4533: 4529: 4525: 4521: 4517: 4510: 4503: 4495: 4491: 4487: 4483: 4479: 4475: 4470: 4465: 4461: 4457: 4453: 4446: 4438: 4434: 4429: 4424: 4420: 4416: 4412: 4408: 4404: 4400: 4396: 4389: 4373: 4369: 4363: 4359: 4358: 4351: 4335: 4331: 4327: 4323: 4319: 4315: 4311: 4307: 4303: 4299: 4292: 4290: 4288: 4279: 4275: 4271: 4267: 4263: 4259: 4255: 4251: 4247: 4243: 4239: 4232: 4224: 4220: 4216: 4212: 4208: 4204: 4200: 4196: 4192: 4188: 4184: 4177: 4169: 4165: 4161: 4157: 4153: 4149: 4145: 4141: 4137: 4130: 4122: 4118: 4114: 4110: 4106: 4102: 4098: 4094: 4087: 4085: 4076: 4072: 4067: 4062: 4058: 4054: 4050: 4043: 4035: 4031: 4026: 4021: 4017: 4013: 4009: 4005: 4001: 3997: 3993: 3986: 3978: 3974: 3970: 3966: 3962: 3958: 3954: 3950: 3943: 3935: 3931: 3926: 3921: 3917: 3913: 3909: 3905: 3901: 3894: 3886: 3882: 3877: 3872: 3868: 3864: 3860: 3856: 3852: 3845: 3837: 3833: 3828: 3823: 3819: 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2085: 2078: 2070: 2066: 2061: 2056: 2052: 2048: 2045:(7): 629–33. 2044: 2040: 2036: 2029: 2021: 2019:0-205-42651-4 2015: 2011: 2010:Biopsychology 2004: 1996: 1992: 1987: 1982: 1977: 1972: 1968: 1964: 1960: 1956: 1952: 1945: 1937: 1933: 1928: 1923: 1919: 1915: 1911: 1904: 1885: 1881: 1877: 1873: 1869: 1864: 1859: 1855: 1851: 1844: 1840: 1834: 1818: 1814: 1810: 1806: 1799: 1791: 1784: 1768: 1764: 1760: 1756: 1752: 1748: 1744: 1740: 1736: 1732: 1725: 1717: 1713: 1708: 1703: 1698: 1693: 1689: 1685: 1681: 1674: 1666: 1660: 1656: 1652: 1647: 1639: 1631: 1627: 1623: 1619: 1612: 1604: 1598: 1594: 1587: 1579: 1578: 1573: 1566: 1562: 1553: 1549: 1545: 1541: 1537: 1531: 1527: 1517: 1514: 1512: 1510: 1506: 1504: 1501: 1499: 1496: 1494: 1491: 1488: 1485: 1483: 1480: 1478: 1475: 1474: 1468: 1466: 1462: 1458: 1454: 1450: 1446: 1438: 1435: 1433: 1429: 1426: 1423: 1420: 1415: 1412: 1409: 1406: 1405: 1403: 1400: 1397: 1394: 1391: 1388: 1385: 1382: 1379: 1376: 1375: 1374: 1363: 1359: 1356: 1353: 1349: 1348: 1347: 1339: 1337: 1334:waste ponds, 1333: 1329: 1325: 1321: 1317: 1313: 1308: 1306: 1302: 1295: 1285: 1283: 1278: 1274: 1268: 1258: 1256: 1255: 1250: 1246: 1241: 1237: 1232: 1230: 1229: 1224: 1220: 1215: 1210: 1208: 1207: 1202: 1197: 1193: 1189: 1185: 1181: 1177: 1173: 1167: 1157: 1155: 1151: 1147: 1143: 1139: 1133: 1131: 1127: 1123: 1119: 1115: 1104: 1100: 1098: 1094: 1090: 1085: 1076: 1074: 1070: 1066: 1062: 1061:antennal lobe 1058: 1053: 1051: 1047: 1042: 1038: 1036: 1035:limbic system 1032: 1028: 1023: 1018: 1014: 1010: 1006: 1002: 998: 994: 990: 985: 983: 979: 975: 971: 970:acetylcholine 967: 963: 959: 954: 950: 946: 942: 938: 934: 930: 926: 922: 915: 914:cranial nerve 911: 907: 897: 888: 886: 882: 877: 875: 870: 866: 862: 858: 853: 849: 847: 843: 839: 835: 831: 827: 823: 819: 811: 807: 803: 799: 795: 791: 781: 779: 775: 771: 766: 764: 760: 756: 752: 748: 744: 740: 736: 732: 727: 725: 721: 716: 712: 708: 704: 698: 683: 669: 665: 663: 659: 655: 651: 647: 643: 639: 635: 632: 627: 624: 623:Grizzly bears 620: 618: 614: 610: 606: 602: 598: 596: 592: 591: 586: 582: 578: 574: 570: 566: 562: 558: 554: 550: 546: 542: 532: 530: 526: 522: 518: 514: 510: 504: 494: 492: 488: 484: 480: 476: 466: 464: 460: 456: 451: 449: 445: 441: 437: 433: 429: 423: 421: 420:immune system 417: 413: 403: 401: 397: 393: 389: 379: 377: 373: 369: 365: 361: 357: 353: 349: 345: 340: 338: 334: 330: 326: 322: 307: 305: 301: 297: 292: 290: 286: 282: 277: 274: 270: 266: 262: 258: 254: 250: 249:Linda B. Buck 245: 239: 234: 232: 228: 220: 219: 214: 213: 208: 199: 197: 193: 189: 185: 181: 176: 174: 170: 166: 162: 158: 154: 150: 146: 141: 139: 135: 131: 127: 126:special sense 123: 119: 107: 101: 97: 94: 91: 89: 85: 80: 76: 72: 68: 65: 62: 60: 56: 51: 46: 42: 38: 32: 27: 22: 19: 6720:Hypothalamus 6700:Hypothalamus 6573:mitral cells 6504:Microanatomy 6496: 6353:Phantom pain 6338:Hyperalgesia 6312:Hypoesthesia 6130:Hearing loss 5951:TRP channels 5928:Osmoreceptor 5895:Photopigment 5819:Baroreceptor 5758:Gravitropism 5730:Frog hearing 5710:Echolocation 5594: 5506:Facial (VII) 5216: 5183: 5134: 5130: 5120: 5101: 5073: 5067: 5055:. Retrieved 5046: 5036: 5024:. Retrieved 5015: 5005: 4972: 4968: 4932: 4928: 4918: 4875: 4871: 4815: 4811: 4805: 4772: 4768: 4710: 4706: 4696: 4684:. Retrieved 4648: 4644: 4631: 4619:. Retrieved 4586: 4582: 4557:. Retrieved 4519: 4515: 4502: 4459: 4455: 4445: 4402: 4398: 4388: 4376:. Retrieved 4356: 4350: 4338:. 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Retrieved 2443: 2433: 2398: 2394: 2384: 2359: 2355: 2316:(2): 92–96. 2313: 2309: 2273: 2228: 2224: 2218: 2185: 2181: 2175: 2148: 2144: 2134: 2091: 2087: 2077: 2042: 2039:EMBO Reports 2038: 2028: 2009: 2003: 1958: 1954: 1944: 1917: 1913: 1903: 1891:. Retrieved 1853: 1849: 1833: 1821:. Retrieved 1812: 1808: 1798: 1789: 1783: 1771:. Retrieved 1738: 1734: 1724: 1687: 1683: 1673: 1650: 1638: 1621: 1617: 1611: 1592: 1586: 1575: 1565: 1551: 1547: 1543: 1534:Coined from 1530: 1508: 1442: 1372: 1345: 1309: 1305:olfactometer 1297: 1270: 1252: 1233: 1226: 1211: 1204: 1203:, including 1174:that enable 1169: 1154:hypothalamus 1134: 1110: 1101: 1093:mitral cells 1086: 1082: 1073:lateral horn 1054: 1043: 1039: 986: 949:Mitral cells 903: 878: 854: 850: 844:. A calcium- 787: 767: 728: 700: 675: 666: 628: 621: 617:Basset Hound 603: 599: 594: 589: 588: 573:king penguin 571:(VOCs) from 538: 506: 479:tracking dog 475:scent trails 472: 452: 424: 409: 398:. Whereas a 385: 358:. The human 352:chemesthesis 341: 318: 298:proposed by 293: 281:shape theory 278: 253:Richard Axel 246: 235: 224: 216: 210: 177: 153:nasal cavity 142: 121: 117: 115: 18: 6533:Tufted cell 6435:Synesthesia 6343:Hypoalgesia 6317:Paresthesia 6302:Formication 6297:CMT disease 6264:Hypergeusia 6087:Visual snow 6062:Photophobia 6057:Papilledema 6047:Oscillopsia 6033:Hemeralopia 5900:Aureochrome 5735:Toad vision 5659:Suffocation 5573:Perceptions 5026:22 February 4833:11336/19586 4818:: 317–333. 4621:19 February 3583:: 114–124. 3291:1411.0165v2 2987:7 September 2977:Nature News 2858:7 September 2848:Nature News 2454:30 December 1893:17 February 1572:"olfaction" 1396:Presbyosmia 1240:sensitively 1196:oviposition 1022:hippocampus 941:micrometers 857:ion channel 703:vertebrates 613:scent trail 609:bloodhounds 605:Scenthounds 595:microsmotic 590:macrosmatic 557:albatrosses 337:mastication 257:Nobel Prize 151:within the 147:binds to a 82:Identifiers 6787:Categories 6511:Epithelium 6455:Pareidolia 6410:Allochiria 6393:Multimodal 6274:Parageusia 6269:Hypogeusia 6241:Phantosmia 6221:Hyperosmia 6165:Vestibular 6145:Palinopsia 6105:Amblyaudia 6052:Palinopsia 6037:Nyctalopia 5961:Nociceptor 5768:Artificial 5491:Optic (II) 5322:Perception 5271:Processes 5263:perception 5137:: 135694. 5074:Der Geruch 4686:1 December 4559:4 November 3345:: e07865. 3284:: e07865. 3104:2 November 1773:10 October 1651:Psychology 1554:"to make". 1522:References 1453:SARS-CoV-1 1445:SARS-CoV-2 1437:Osmophobia 1422:Heterosmia 1414:Phantosmia 1384:Hyperosmia 1330:; and the 1265:See also: 1228:Drosophila 1192:pheromones 1160:In insects 1114:pheromones 1007:, and the 846:calmodulin 828:with some 759:antibodies 715:epithelium 654:pheromones 644:. In some 631:catarrhine 577:carnivores 563:, and the 491:turbulence 487:chemotaxis 455:inbreeding 386:Smell and 368:inhalation 364:exhalation 348:trigeminal 300:Luca Turin 134:pheromones 6793:Olfaction 6568:glomeruli 6251:Gustatory 6203:Olfactory 6067:Photopsia 6013:Amaurosis 5987:Disorders 5880:Cone cell 5799:Types of 5410:Inner ear 5345:Cognition 5335:Awareness 5284:Sensation 5259:Sensation 5151:1872-7972 5057:15 August 4975:: 83–89. 4850:207568844 4775:: 31–41. 4611:207090474 4478:1531-5320 4419:1464-3553 4378:14 August 4340:14 August 4322:2352-1546 4308:: 30–39. 4262:0166-2236 4207:1432-0770 4160:0013-4694 3138:14 August 2817:243966350 2809:1947-5454 2292:882238865 2094:: 25103. 1880:218627484 1823:14 August 1684:Neurology 1544:olfactere 1465:olfaction 1432:body odor 1369:Disorders 1261:In plants 1249:aldehydes 974:serotonin 945:glomeruli 824:(largely 810:G protein 753:, salts, 743:dendrites 656:; a male 638:cetaceans 581:ungulates 549:tubenoses 376:forebrain 238:Lucretius 161:Glomeruli 124:, is the 122:olfaction 75:chemicals 37:carnation 6692:Amygdala 6236:Parosmia 6226:Hyposmia 6216:Dysosmia 6155:Tinnitus 6097:Auditory 6072:Polyopia 6028:Diplopia 5885:Rod cell 5683:Nonhuman 5636:Internal 5477:Sensory 5383:External 5292:Stimulus 5275:concepts 5169:33600902 5051:Archived 5020:Archived 4997:29588018 4951:28468812 4910:25649823 4842:26982441 4797:26416146 4745:19372867 4737:11361090 4728:11146473 4677:Archived 4673:30978362 4665:19151709 4615:Archived 4603:23806440 4550:Archived 4546:25086608 4494:43129982 4486:16082803 4437:28641388 4372:Archived 4334:Archived 4330:27989941 4121:22505357 4113:12668289 4075:11418503 4034:19699095 3977:28094328 3969:19554043 3934:18923046 3885:26752161 3836:24672011 3772:27916458 3723:20637621 3666:17724338 3607:25701815 3491:16878137 3437:Archived 3371:26151672 3320:26151672 3259:24653035 3200:23 March 3194:Archived 3163:Archived 3159:BBC News 3098:Archived 3068:Archived 3030:34180521 2981:Archived 2958:22714804 2909:23910657 2852:Archived 2827:13 March 2821:Archived 2760:13 March 2726:17997307 2669:19067327 2620:12810039 2585:42316168 2534:15777804 2499:15621496 2491:16337283 2448:Archived 2425:20181598 2376:11148312 2330:20171264 2245:15034588 2210:12183125 2126:27112241 2069:17603536 1995:12612342 1884:Archived 1872:32413294 1841:(2020). 1817:Archived 1815:: 7–18. 1767:Archived 1755:28504103 1716:25788559 1540:olfactus 1471:See also 1457:MERS-CoV 1408:Parosmia 1402:Dysosmia 1390:Hyposmia 1352:caprylic 1301:sniffing 1236:volatile 1214:antennae 1184:foraging 1146:amygdala 1124:and the 1046:nostrils 1001:amygdala 812:called G 774:sensilla 662:bombykol 658:silkworm 634:primates 561:vultures 525:cilantro 497:Genetics 430:and the 372:thalamus 321:auditory 310:Function 285:molecule 221:, Paris. 149:receptor 70:Function 6755:General 6405:Agnosia 6284:Tactile 6259:Ageusia 6211:Anosmia 6173:Vertigo 6077:Scotoma 6018:Anopsia 5479:cranial 5350:Feeling 5160:7883672 4977:Bibcode 4901:4317791 4880:Bibcode 4777:Bibcode 4537:4146660 4428:5863551 4278:3947494 4270:2475938 4223:7932310 4215:2551392 4025:2901510 4004:Bibcode 3925:2588668 3876:4948943 3827:3965788 3763:5196021 3714:2929674 3693:Bibcode 3657:1964822 3634:Bibcode 3598:4396857 3558:7724600 3526:Bibcode 3499:2864195 3471:Bibcode 3415:265–275 3386:bioRxiv 3362:4491593 3311:4491593 3250:4483192 3229:Bibcode 3221:Science 3169:20 June 3096:. PBS. 3077:21 July 2949:3408771 2887:Bibcode 2789:Bibcode 2717:2148465 2696:Bibcode 2660:6870682 2577:4011726 2542:8568275 2416:2846283 2338:2671577 2253:1073550 2190:Bibcode 2167:8985605 2117:4844992 2096:Bibcode 2060:1905909 1963:Bibcode 1936:1840504 1763:5121325 1707:4408285 1655:166–171 1511:(novel) 1509:Perfume 1378:Anosmia 1223:antenna 1176:insects 1130:flehmen 1057:insects 822:cations 770:insects 755:enzymes 741:on the 709:in the 553:petrels 551:(e.g., 541:mammals 392:rodents 382:Hearing 342:Smell, 261:Mammals 184:anosmia 173:emotion 93:D012903 53:Details 6556:neuron 6521:mucosa 6516:glands 5995:Visual 5946:Cilium 5692:Animal 5664:Nausea 5654:Thirst 5649:Hunger 5521:Spinal 5360:Qualia 5167:  5157:  5149:  5108:  5080:  4995:  4949:  4908:  4898:  4848:  4840:  4795:  4743:  4735:  4725:  4671:  4663:  4609:  4601:  4544:  4534:  4492:  4484:  4476:  4435:  4425:  4417:  4364:  4328:  4320:  4276:  4268:  4260:  4221:  4213:  4205:  4166:  4158:  4119:  4111:  4073:  4032:  4022:  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Index


carnation
chemoreceptors
brain
System
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chemicals
MeSH
D012903
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edit on Wikidata
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pheromones
taste
odor
receptor
nasal cavity
olfactory system
Glomeruli
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memory
emotion
nose
anosmia
upper respiratory infections
traumatic brain injury
neurodegenerative disease

The Lady and the Unicorn

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