{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T10:07:18Z","timestamp":1788948438604,"version":"build-2803163510"},"reference-count":209,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T00:00:00Z","timestamp":1788912000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Pharmacol."],"abstract":"<jats:p>Detection and discrimination of chemosensory stimuli (usually known as odorants) by the olfactory system exert a crucial role in animal survival. Olfactory receptors (ORs) are a large family of G protein-coupled receptors (GPCRs) mainly expressed by the main olfactory neuroepithelium (OE) at the nasal level and can sense different odorants affecting the electric activity of olfactory sensory neurons (OSNs) eventually generating the sense of smell. Recent evidence suggests that the olfactory mucosa, namely, the OE plus the olfactory mucus, plays a critical role as an immune barrier. Healthy olfactory pathways need intact function and anatomy within the nose itself, the OE, the olfactory bulb, and the central brain structures in charge of olfactory processing. Olfactory loss influences nutrition, safety, and social relationships, and subjects with olfactory impairment have been revealed to be at greater risk for overall mortality. Of note, ORs are not only expressed within the OE but an ecnomotopic expression has also been identified, serving different physiological functions, such as glucose and lipid metabolism and wound healing. It should be noted that ligands of ecnomotopic ORs may originate from naturally occurring or synthetic odor compounds present in the foods and\/or food additives, as well as from endogenous compounds produced by host and\/or microbiome cells. This review summarizes the anatomy, physiology, and immunology of the human olfactory system. Then, it delves into the ecnomotopic OR expression and signaling pathways. Finally, it reports recent data highlighting how the OR expression can be modulated offering guidance and future directions for clinical translatability.<\/jats:p>","DOI":"10.3389\/fphar.2026.1905572","type":"journal-article","created":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T09:16:25Z","timestamp":1788945385000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Physiopathology of nasal and ecnomotopic olfactory receptors: focus on molecular pathways and potential pharmacological modulation"],"prefix":"10.3389","volume":"17","author":[{"given":"Amedeo","family":"Amedei","sequence":"first","affiliation":[{"name":"Department of Clinical and Experimental Medicine, University of Florence","place":["Florence, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Cinzia","family":"Parolini","sequence":"additional","affiliation":[{"name":"Department of Pharmacological and Biomolecular Sciences, Rodolfo Paoletti, Universit\u00e0 degli Studi di Milano","place":["Milano, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,9,9]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"1827","DOI":"10.1161\/CIRCULATIONAHA.125.078927","article-title":"Olfactory receptor activation reduces platelet reactivity and arterial thrombosis through actin cytoskeleton remodeling","volume":"153","author":"Aggarwal","year":"2026","journal-title":"Circulation"},{"key":"B2","doi-asserted-by":"publisher","first-page":"e51227","DOI":"10.1371\/journal.pone.0051227","article-title":"A physiological increase of insulin in the olfactory bulb decreases detection of a learned aversive odor and abolishes food odor-induced sniffing behavior in rats","volume":"7","author":"Aim\u00e9","year":"2012","journal-title":"PLoS One"},{"key":"B3","doi-asserted-by":"publisher","first-page":"2911","DOI":"10.1021\/acs.jcim.3c00380","article-title":"Machine learning-based modeling of olfactory receptors in their inactive state: human OR51E2 as a case study","volume":"63","author":"Alfonso-Prieto","year":"2023","journal-title":"J. Chem. Inf. Model"},{"key":"B4","doi-asserted-by":"publisher","first-page":"101158","DOI":"10.1016\/j.jbior.2026.101158","article-title":"Ecnomotopic expression of olfactory receptors is affected by human apolipoproteins A-IMilano and A-II: evidence from liver microarray analyses","volume":"100","author":"Amedei","year":"2026","journal-title":"Adv. Biol. Regul."},{"key":"B5","doi-asserted-by":"publisher","first-page":"2428","DOI":"10.3390\/biomedicines13102428","article-title":"Potential and future therapeutic applications of eicosapentaenoic\/docosahexaenoic acid and probiotics in chronic low-grade inflammation","volume":"13","author":"Amedei","year":"2025","journal-title":"Biomedicines"},{"key":"B6","doi-asserted-by":"publisher","first-page":"149","DOI":"10.3233\/jad-2005-7208","article-title":"Olfactory involvement in aging and Alzheimer's disease: an autopsy study","volume":"7","author":"Attems","year":"2005","journal-title":"J. Alzheimers Dis."},{"key":"B7","doi-asserted-by":"publisher","first-page":"5621","DOI":"10.1523\/JNEUROSCI.4630-06.2007","article-title":"Multiple modes of synaptic excitation of olfactory bulb granule cells","volume":"27","author":"Balu","year":"2007","journal-title":"J. Neurosci."},{"key":"B8","doi-asserted-by":"publisher","first-page":"631","DOI":"10.1002\/glia.23744","article-title":"Sublamina-specific organization of the blood brain barrier in the mouse olfactory nerve layer","volume":"68","author":"Beiersdorfer","year":"2020","journal-title":"Glia"},{"key":"B9","doi-asserted-by":"publisher","first-page":"548","DOI":"10.3390\/life14050548","article-title":"Role of ectopic olfactory receptors in the regulation of the cardiovascular-kidney-metabolic axis","volume":"14","author":"Beito","year":"2024","journal-title":"Life (Basel)"},{"key":"B10","doi-asserted-by":"publisher","first-page":"185","DOI":"10.1016\/j.cell.2017.05.034","article-title":"Enterochromaffin cells are gut chemosensors that couple to sensory neural pathways","volume":"170","author":"Bellono","year":"2017","journal-title":"Cell"},{"key":"B11","doi-asserted-by":"publisher","first-page":"1931","DOI":"10.1126\/science.1109886","article-title":"Elementary response of olfactory receptor neurons to odorants","volume":"308","author":"Bhandawat","year":"2005","journal-title":"Science"},{"key":"B12","doi-asserted-by":"publisher","first-page":"e0213545","DOI":"10.1371\/journal.pone.0213545","article-title":"Vertebrate odorant binding proteins as antimicrobial humoral components of innate immunity for pathogenic microorganisms","volume":"14","author":"Bianchi","year":"2019","journal-title":"PLoS One"},{"key":"B13","doi-asserted-by":"publisher","first-page":"742","DOI":"10.1038\/s41586-023-05798-y","article-title":"Structural basis of odorant recognition by a human odorant receptor","volume":"615","author":"Billesb\u00f8lle","year":"2023","journal-title":"Nature"},{"key":"B14","doi-asserted-by":"publisher","first-page":"158","DOI":"10.1016\/j.molmet.2019.09.001","article-title":"Single cell transcriptomic profiling of large intestinal enteroendocrine cells in mice - identification of selective stimuli for insulin-like peptide-5 and glucagon-like peptide-1 co-expressing cells","volume":"29","author":"Billing","year":"2019","journal-title":"Mol. Metab."},{"key":"B15","doi-asserted-by":"publisher","first-page":"343","DOI":"10.1016\/j.conb.2005.05.014","article-title":"Regulation of cyclic nucleotide-gated channels","volume":"15","author":"Bradley","year":"2005","journal-title":"Curr. Opin. Neurobiol."},{"key":"B16","doi-asserted-by":"publisher","first-page":"589","DOI":"10.1007\/s00441-021-03467-y","article-title":"Modulation of olfactory signal detection in the olfactory epithelium: focus on the internal and external environment, and the emerging role of the immune system","volume":"384","author":"Bryche","year":"2021","journal-title":"Cell Tissue Res."},{"key":"B17","doi-asserted-by":"publisher","first-page":"175","DOI":"10.1016\/0092-8674(91)90418-x","article-title":"A novel multigene family may encode odorant receptors: a molecular basis for odor recognition","volume":"65","author":"Buck","year":"1991","journal-title":"Cell"},{"key":"B18","doi-asserted-by":"publisher","first-page":"58","DOI":"10.1111\/imr.13066","article-title":"Immune dynamics in the CNS and its barriers during homeostasis and disease","volume":"306","author":"Buckley","year":"2022","journal-title":"Immunol. Rev."},{"key":"B19","doi-asserted-by":"publisher","first-page":"2823","DOI":"10.1038\/jid.2014.273","article-title":"A synthetic sandalwood odorant induces wound-healing processes in human keratinocytes via the olfactory receptor OR2AT4","volume":"134","author":"Busse","year":"2014","journal-title":"J. Invest Dermatol."},{"key":"B20","doi-asserted-by":"publisher","first-page":"615","DOI":"10.1093\/chemse\/bjz050","article-title":"Commensal bacteria regulate gene expression and differentiation in vertebrate olfactory systems through transcription factor REST","volume":"44","author":"Casadei","year":"2019","journal-title":"Chem. Senses"},{"key":"B21","doi-asserted-by":"publisher","first-page":"bjae033","DOI":"10.1093\/chemse\/bjae033","article-title":"Subjective cognitive and olfactory impairments predict different prospective dementia outcomes","volume":"49","author":"Cedres","year":"2024","journal-title":"Chem. Senses"},{"key":"B22","doi-asserted-by":"publisher","first-page":"240","DOI":"10.1038\/nature15721","article-title":"Oxygen regulation of breathing through an olfactory receptor activated by lactate","volume":"527","author":"Chang","year":"2015","journal-title":"Nature"},{"key":"B23","doi-asserted-by":"publisher","first-page":"1441","DOI":"10.1212\/WNL.0000000000004382","article-title":"Olfaction and incident Parkinson disease in US white and black older adults","volume":"89","author":"Chen","year":"","journal-title":"Neurology"},{"key":"B24","doi-asserted-by":"publisher","first-page":"8089","DOI":"10.1073\/pnas.1620664114","article-title":"Acute inflammation regulates neuroregeneration through the NF-\u03baB pathway in olfactory epithelium","volume":"114","author":"Chen","year":"","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B25","doi-asserted-by":"publisher","first-page":"2104","DOI":"10.1002\/jcp.25929","article-title":"The diversified function and potential therapy of ectopic olfactory receptors in non-olfactory tissues","volume":"233","author":"Chen","year":"2018","journal-title":"J. Cell Physiol."},{"key":"B26","doi-asserted-by":"publisher","first-page":"501","DOI":"10.1016\/j.stem.2019.08.011","article-title":"Chronic inflammation directs an olfactory stem cell functional switch from neuroregeneration to immune defense","volume":"25","author":"Chen","year":"2019","journal-title":"Cell Stem Cell"},{"key":"B27","doi-asserted-by":"publisher","first-page":"8105","DOI":"10.1038\/s41467-023-43983-9","article-title":"Understanding the molecular mechanisms of odorant binding and activation of the human OR52 family","volume":"14","author":"Choi","year":"2023","journal-title":"Nat. Commun."},{"key":"B28","doi-asserted-by":"publisher","first-page":"320","DOI":"10.1177\/0301006616663216","article-title":"The olfactory mosaic: bringing an olfactory network together for odor perception","volume":"46","author":"Courtiol","year":"2017","journal-title":"Perception"},{"key":"B29","doi-asserted-by":"publisher","first-page":"253","DOI":"10.1530\/EJE-12-0814","article-title":"Olfactory receptor 51E1 protein as a potential novel tissue biomarker for small intestine neuroendocrine carcinomas","volume":"168","author":"Cui","year":"2013","journal-title":"Eur. J. Endocrinol."},{"key":"B30","doi-asserted-by":"publisher","first-page":"1195","DOI":"10.1101\/gr.220111.116","article-title":"Microbiota regulate intestinal epithelial gene expression by suppressing the transcription factor hepatocyte nuclear factor 4 alpha","volume":"27","author":"Davidson","year":"2017","journal-title":"Genome Res."},{"key":"B31","doi-asserted-by":"publisher","first-page":"499","DOI":"10.1038\/s41586-024-08126-0","article-title":"Engineered odorant receptors illuminate the basis of odour discrimination","volume":"635","author":"De March","year":"2024","journal-title":"Nature"},{"key":"B32","doi-asserted-by":"publisher","first-page":"2043","DOI":"10.1523\/JNEUROSCI.5317-08.2009","article-title":"External tufted cells drive the output of olfactory bulb glomeruli","volume":"29","author":"De Saint Jan","year":"2009","journal-title":"J. Neurisci"},{"key":"B33","doi-asserted-by":"publisher","first-page":"1098212","DOI":"10.3389\/fimmu.2022.1098212","article-title":"Neurotrophic and immunomodulatory effects of olfactory ensheathing cells as a strategy for neuroprotection and regeneration","volume":"13","author":"Denaro","year":"2022","journal-title":"Front. Immunol."},{"key":"B34","doi-asserted-by":"publisher","first-page":"182","DOI":"10.1212\/WNL.0000000000001132","article-title":"Olfactory deficits predict cognitive decline and Alzheimer dementia in an urban community","volume":"84","author":"Devanard","year":"2015","journal-title":"Neurology"},{"key":"B35","doi-asserted-by":"publisher","first-page":"1402","DOI":"10.3390\/ijms20061402","article-title":"Beyond the flavour: the potential druggability of chemosensory G protein-coupled receptors","volume":"20","author":"Di Pizio","year":"2019","journal-title":"Int. J. Mol. Sci."},{"key":"B36","doi-asserted-by":"publisher","first-page":"1183","DOI":"10.1002\/glia.24527","article-title":"ACKR3 in olfactory glia cells shapes the immune defense of the olfactory mucosa","volume":"72","author":"Dietz","year":"2024","journal-title":"Glia"},{"key":"B37","doi-asserted-by":"publisher","first-page":"155","DOI":"10.1146\/annurev-psych-042023-101155","article-title":"What does the human olfactory system do, and how does it Do it?","volume":"75","author":"Dike\u00e7ligil","year":"2024","journal-title":"Annu. Rev. Psycol"},{"key":"B38","doi-asserted-by":"publisher","first-page":"304","DOI":"10.1038\/s44319-023-00013-5","article-title":"The olfactory receptor Olfr78 promotes differentiation of enterochromaffin cells in the mouse colon","volume":"25","author":"Dinsart","year":"2024","journal-title":"EMBO Rep."},{"key":"B39","doi-asserted-by":"publisher","first-page":"e700","DOI":"10.1212\/WNL.0000000000006919","article-title":"Impaired olfaction is associated with cognitive decline and neurodegeneration in the brain","volume":"92","author":"Dintica","year":"2019","journal-title":"Neurology"},{"key":"B40","doi-asserted-by":"publisher","first-page":"1080","DOI":"10.1093\/gerona\/glx081","article-title":"The prevalence of anosmia and associated factors among U.S. Black and white older adults","volume":"72","author":"Dong","year":"2017","journal-title":"J. Gerontol. A Biol. Sci. Med. Sci."},{"key":"B41","doi-asserted-by":"publisher","first-page":"20","DOI":"10.3389\/fpsyg.2014.00020","article-title":"The influences of age on olfaction: a review","volume":"5","author":"Doty","year":"2014","journal-title":"Front. Psychol."},{"key":"B42","doi-asserted-by":"publisher","first-page":"295","DOI":"10.1007\/s40291-026-00832-x","article-title":"The emerging role of olfactory receptors: from genomics to precision medicine","volume":"30","author":"Dubey","year":"2026","journal-title":"Mol. Diagn Ther."},{"key":"B43","doi-asserted-by":"publisher","first-page":"1444","DOI":"10.1038\/nm.4432","article-title":"Asprosin is a centrally acting orexigenic hormone","volume":"23","author":"Duerrschmid","year":"2017","journal-title":"Nat. Med."},{"key":"B44","doi-asserted-by":"publisher","first-page":"323","DOI":"10.1038\/s41593-020-0587-9","article-title":"Single-cell analysis of olfactory neurogenesis and differentiation in adult humans","volume":"23","author":"Durante","year":"2020","journal-title":"Nat. Neurosci."},{"key":"B45","doi-asserted-by":"publisher","first-page":"7595","DOI":"10.1073\/pnas.92.16.7595","article-title":"Disruption of cellular signaling pathways by daunomycin through destabilization of nonlamellar membrane structures","volume":"92","author":"Escriba","year":"1995","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B46","doi-asserted-by":"publisher","first-page":"836","DOI":"10.1016\/j.bbamem.2006.09.001","article-title":"Lipid-protein interactions in GPCR-associated signaling","volume":"1768","author":"Escrib\u00e1","year":"2007","journal-title":"Biochim. Biophys. Acta"},{"key":"B47","doi-asserted-by":"publisher","first-page":"847","DOI":"10.1002\/bies.201400016","article-title":"Microbes and animal olfactory communication: where do we go from here?","volume":"36","author":"Ezenwa","year":"2014","journal-title":"Bioessays"},{"key":"B48","doi-asserted-by":"publisher","first-page":"108923","DOI":"10.1016\/j.neuropharm.2021.108923","article-title":"Olfaction in the context of obesity and diabetes: insights from animal models to humans","volume":"206","author":"Faour","year":"2022","journal-title":"Neuropharmacology"},{"key":"B49","doi-asserted-by":"publisher","first-page":"446","DOI":"10.3390\/md23110446","article-title":"Marine bioactive components and chronic neuroinflammation: focus on neurodegenerative disease","volume":"23","author":"Favari","year":"2025","journal-title":"Mar. Drugs"},{"key":"B50","doi-asserted-by":"publisher","first-page":"121","DOI":"10.1186\/1471-2164-7-121","article-title":"Widespread ectopic expression of olfactory receptor genes","volume":"7","author":"Feldmesser","year":"2006","journal-title":"BMC Genom"},{"key":"B51","doi-asserted-by":"publisher","first-page":"163","DOI":"10.3389\/fnagi.2016.00163","article-title":"Olfactory receptors in non-chemosensory organs: the nervous system in health and disease","volume":"8","author":"Ferrer","year":"2016","journal-title":"Front. Aging Neurosci."},{"key":"B52","doi-asserted-by":"publisher","first-page":"2154","DOI":"10.1002\/cne.25325","article-title":"Integrated age-related immunohistological changes occur in human olfactory epithelium and olfactory bulb","volume":"530","author":"Fitzek","year":"2022","journal-title":"J. Comp. Neurol."},{"key":"B53","doi-asserted-by":"publisher","first-page":"24687","DOI":"10.1038\/srep24687","article-title":"Olfactory epithelium changes in germfree mice","volume":"6","author":"Fran\u00e7ois","year":"2016","journal-title":"Sci. Rep."},{"key":"B54","doi-asserted-by":"publisher","first-page":"113826","DOI":"10.1016\/j.yexcr.2023","article-title":"Microarray analysis identifies human apoA-IMilano and apoA-II as determinants of the liver gene expression related to lipid and energy metabolism","volume":"433","author":"Ganzetti","year":"2023","journal-title":"Exp. Cell Res."},{"key":"B55","doi-asserted-by":"publisher","first-page":"e0202754","DOI":"10.1371\/journal.pone.0202754","article-title":"Microvillous cells in the olfactory epithelium express elements of the solitary chemosensory cell transduction signaling cascade","volume":"13","author":"Genovese","year":"2018","journal-title":"PLoS One"},{"key":"B56","doi-asserted-by":"publisher","first-page":"307","DOI":"10.1093\/molbev\/msg013","article-title":"Population differences in the human functional olfactory repertoire","volume":"20","author":"Gilad","year":"2003","journal-title":"Mol. Biol. Evol."},{"key":"B57","doi-asserted-by":"publisher","first-page":"11044","DOI":"10.1016\/j.abb.2025.110404","article-title":"The odorant (R)-(-)-carvone promotes glucose-stimulated insulin secretion via the olfactory receptor Olfr1259 in pancreatic \u03b2-TC6 cells","volume":"768","author":"Gong","year":"2025","journal-title":"Arch. Biochem. Biophys."},{"key":"B58","doi-asserted-by":"publisher","first-page":"649","DOI":"10.1016\/j.devcel.2009.09.004","article-title":"MOR23 promotes muscle regeneration and regulates cell adhesion and migration","volume":"17","author":"Griffin","year":"2009","journal-title":"Dev. Cell"},{"key":"B59","doi-asserted-by":"publisher","first-page":"104971","DOI":"10.1016\/j.tifs.2025.104971","article-title":"A systematic review of the structure and function of human olfactory receptors and key technologies involved","volume":"159","author":"Gu","year":"2025","journal-title":"Trends Food Sci. and Technol."},{"key":"B60","doi-asserted-by":"publisher","first-page":"551","DOI":"10.1093\/chemse\/26.5.551","article-title":"Parallel-distributed processing in olfactory cortex: new insights from morphological and physiological analysis of neuronal circuitry","volume":"26","author":"Haberly","year":"2001","journal-title":"Chem. Senses"},{"key":"B61","doi-asserted-by":"publisher","first-page":"1251","DOI":"10.1210\/js.2018-00165","article-title":"Olfactory Receptor OR51E1 mediates GLP-1 secretion in human and rodent enteroendocrine L cells","volume":"2","author":"Han","year":"2018","journal-title":"J. Endocr. Soc."},{"key":"B62","doi-asserted-by":"publisher","first-page":"1251","DOI":"10.1126\/science.aad2456","article-title":"Single-cell transcriptomics reveals receptor transformations during olfactory neurogenesis","volume":"350","author":"Hanchate","year":"2015","journal-title":"Science"},{"key":"B63","doi-asserted-by":"publisher","first-page":"115","DOI":"10.1186\/1471-2202-9-115","article-title":"Is TrpM5 a reliable marker for chemosensory cells? Multiple types of microvillous cells in the main olfactory epithelium of mice","volume":"9","author":"Hansen","year":"2008","journal-title":"BMC Neurosci."},{"key":"B64","doi-asserted-by":"publisher","first-page":"2431","DOI":"10.1093\/brain\/awf251","article-title":"Clinical correlates of selective pathology in the amygdala of patients with Parkinson's disease","volume":"125","author":"Harding","year":"2002","journal-title":"Brain"},{"key":"B65","doi-asserted-by":"publisher","first-page":"1848","DOI":"10.1002\/glia.20899","article-title":"Olfactory ensheathing cells: nitric oxide production and innate immunity","volume":"57","author":"Harris","year":"2009","journal-title":"Glia"},{"key":"B66","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1038\/s41577-020-00477-9","article-title":"Regulation of immune responses by the airway epithelial cell landscape","volume":"21","author":"Hewitt","year":"2021","journal-title":"Nat. Rev. Immunol."},{"key":"B67","doi-asserted-by":"publisher","first-page":"1333","DOI":"10.1002\/ar.22735","article-title":"Odorant-binding proteins and xenobiotic metabolizing enzymes: implications in olfactory perireceptor events","volume":"296","author":"Heydel","year":"2013","journal-title":"Anat. Rec. Hob."},{"key":"B68","doi-asserted-by":"crossref","first-page":"469","DOI":"10.1002\/(SICI)1096-9861(19981102)400:4<469::AID-CNE3>3.0.CO;2-8","article-title":"Adult olfactory epithelium contains multipotent progenitors that give rise to neurons and non-neural cells","volume":"400","author":"Huard","year":"1998","journal-title":"J. Comp. Neurol."},{"key":"B69","doi-asserted-by":"publisher","first-page":"833","DOI":"10.1111\/febs.15217","article-title":"The mutual interplay of gut microbiota, diet and human disease","volume":"287","author":"Illiano","year":"2020","journal-title":"FEBS J."},{"key":"B70","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1016\/j.crstbi.2020.10.001","article-title":"A two-stage computational approach to predict novel ligands for a chemosensory receptor","volume":"2","author":"Jabeen","year":"2020","journal-title":"Curr. Res. Struct. Biol."},{"key":"B71","doi-asserted-by":"publisher","first-page":"1010","DOI":"10.3390\/brainsci11081010","article-title":"Volumetry of olfactory structures in mild cognitive impairment and Alzheimer\u2019s disease: a systematic review and a meta-analysis","volume":"11","author":"Jobin","year":"2021","journal-title":"Brain Sci."},{"key":"B72","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1007\/s00395-017-0600-y","article-title":"Medium-chain fatty acids modulate myocardial function via a cardiac odorant receptor","volume":"112","author":"Jovancevic","year":"2017","journal-title":"Basic Res. Cardiol."},{"key":"B73","doi-asserted-by":"publisher","first-page":"281","DOI":"10.1016\/j.tem.2022.01.004","article-title":"Olfactory system and energy metabolism: a two-way street","volume":"33","author":"Jovanovic","year":"2022","journal-title":"Trends Endocrinol. Metab."},{"key":"B74","doi-asserted-by":"publisher","first-page":"4230","DOI":"10.1111\/febs.15606","article-title":"Challenges and possible solutions for decoding extranasal olfactory receptors","volume":"288","author":"Kalra","year":"2021","journal-title":"FEBS J."},{"key":"B75","doi-asserted-by":"publisher","first-page":"e0113170","DOI":"10.1371\/journal.pone.0113170","article-title":"Deep sequencing of the murine olfactory receptor neuron transcriptome","volume":"10","author":"Kanageswaran","year":"2015","journal-title":"PLoS One"},{"key":"B76","doi-asserted-by":"publisher","first-page":"e70107","DOI":"10.1096\/fj.202401468RR","article-title":"MOR23 deficiency exacerbates hepatic steatosis in mice","volume":"38","author":"Kang","year":"2024","journal-title":"FASEB J."},{"key":"B77","doi-asserted-by":"publisher","first-page":"154","DOI":"10.1177\/0898264309353066","article-title":"Prevalence and neurodegenerative or other associations with olfactory impairment in an older community","volume":"22","author":"Karpa","year":"2010","journal-title":"J. Aging Health"},{"key":"B78","doi-asserted-by":"publisher","first-page":"1806","DOI":"10.1523\/JNEUROSCI.4723-04.2005","article-title":"Structural basis for a broad but selective ligand spectrum of a mouse olfactory receptor: mapping the odorant-binding site","volume":"25","author":"Katada","year":"2005","journal-title":"J. Neurosci."},{"key":"B79","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1016\/j.neuroscience.2014.04.050","article-title":"Stereological quantification of olfactory receptor neurons in mice","volume":"272","author":"Kawagishi","year":"2014","journal-title":"Neuroscinece"},{"key":"B80","doi-asserted-by":"publisher","first-page":"511","DOI":"10.1093\/chemse\/bjz046","article-title":"Differential expression of mucins in Murine Olfactory versus respiratory epithelium","volume":"44","author":"Kennel","year":"2019","journal-title":"Chen Senses"},{"key":"B81","doi-asserted-by":"publisher","first-page":"C502","DOI":"10.1152\/ajpcell.00454.2019","article-title":"The olfactory G protein-coupled receptor (Olfr-78\/OR51E2) modulates the intestinal response to colitis","volume":"318","author":"Kotlo","year":"2020","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"B82","doi-asserted-by":"publisher","first-page":"e03038","DOI":"10.1128\/AEM.03038-18","article-title":"The indoor-air microbiota of pig farms drives the composition of the pig farmers\u2019 nasal microbiota in a season-dependent and farm-specific manner","volume":"85","author":"Kraemer","year":"2019","journal-title":"Appl. Environ. Microbiol."},{"key":"B83","doi-asserted-by":"publisher","first-page":"574082","DOI":"10.3389\/fphys.2020.574082","article-title":"The sensing liver: localization and ligands for hepatic Murine Olfactory and taste receptors","volume":"11","author":"Kurtz","year":"2020","journal-title":"Front. Physiol."},{"key":"B84","doi-asserted-by":"publisher","first-page":"1176","DOI":"10.1111\/j.1365-2826.2008.01777.x","article-title":"Expression of insulin system in the olfactory epithelium: first approaches to its role and regulation","volume":"20","author":"Lacroix","year":"2008","journal-title":"J. Neuroendocrinol."},{"key":"B85","doi-asserted-by":"publisher","first-page":"327","DOI":"10.1212\/WNL.0000000000004159","article-title":"Odor identification as a biomarker of preclinical AD in older adults at risk","volume":"89","author":"Lafaille-Magnan","year":"2017","journal-title":"Neurology"},{"key":"B86","doi-asserted-by":"publisher","first-page":"D1370","DOI":"10.1093\/nar\/gkad886","article-title":"M2OR: a database of olfactory receptor-odorant pairs for understanding the molecular mechanisms of olfaction","volume":"52","author":"Lalis","year":"2024","journal-title":"Nucleic Acid. Res."},{"key":"B87","doi-asserted-by":"publisher","first-page":"440","DOI":"10.4103\/1673-5374.317986","article-title":"Gene and protein expression profiles of olfactory ensheathing cells from olfactory bulb versus olfactory mucosa","volume":"17","author":"Lan","year":"2022","journal-title":"Neural Regen. Res."},{"key":"B88","doi-asserted-by":"publisher","first-page":"1764","DOI":"10.1097\/00005537-200410000-00017","article-title":"A study on the frequency of olfactory dysfunction","volume":"114","author":"Landis","year":"2004","journal-title":"Laryngoscope"},{"key":"B89","doi-asserted-by":"publisher","first-page":"2324","DOI":"10.1523\/JNEUROSCI.4507-09.2010","article-title":"A genetic model of chronic rhinosinusitis-associated olfactory inflammation reveals reversible functional impairment and dramatic neuroepithelial reorganization","volume":"30","author":"Lane","year":"2010","journal-title":"J. Neurosci."},{"key":"B90","doi-asserted-by":"publisher","first-page":"547","DOI":"10.1093\/chemse\/bjh059","article-title":"Demographic and cognitive predictors of cued odor identification: evidence from a population-based study","volume":"29","author":"Larsson","year":"2004","journal-title":"Chem. Senses"},{"key":"B91","doi-asserted-by":"publisher","first-page":"718","DOI":"10.1016\/j.tins.2022.08.003","article-title":"The microbiome-nose-brain axis in health and disease","volume":"45","author":"Lazarini","year":"2022","journal-title":"Trends Neurosci."},{"key":"B92","doi-asserted-by":"publisher","first-page":"2974","DOI":"10.1523\/JNEUROSCI.5067-10.2011","article-title":"Olfactory marker protein is critical for functional maturation of olfactory sensory neurons and development of mother preference","volume":"31","author":"Lee","year":"2011","journal-title":"J. Neurosci."},{"key":"B93","doi-asserted-by":"publisher","first-page":"116","DOI":"10.1038\/s41573-018-0002-3","article-title":"Therapeutic potential of ectopic olfactory and taste receptors","volume":"18","author":"Lee","year":"2019","journal-title":"Nat. Rev. Drug Discov."},{"key":"B94","doi-asserted-by":"publisher","first-page":"1698","DOI":"10.3174\/ajnr.A6697","article-title":"Diagnostic accuracy of MRI-Based morphometric parameters for detecting olfactory nerve dysfunction","volume":"41","author":"Lee","year":"2020","journal-title":"AJNR Am. J. Neuroradiol."},{"key":"B95","doi-asserted-by":"publisher","first-page":"763","DOI":"10.1007\/s00702-021-02347-7","article-title":"Hyposmia may predict development of freezing of gait in Parkinson's disease","volume":"128","author":"Lee","year":"2021","journal-title":"J. Neural Transm. (Vienna)"},{"key":"B96","doi-asserted-by":"publisher","first-page":"1842","DOI":"10.1126\/science.1152837","article-title":"Aversive learning enhances perceptual and cortical discrimination of indiscriminable odor cues","volume":"319","author":"Li","year":"2008","journal-title":"Science"},{"key":"B97","doi-asserted-by":"publisher","first-page":"319","DOI":"10.1016\/j.cmet.2019.05.022","article-title":"OLFR734 mediates glucose metabolism as a receptor of asprosin","volume":"30","author":"Li","year":"2019","journal-title":"Cell Metab."},{"key":"B98","doi-asserted-by":"publisher","first-page":"674","DOI":"10.1111\/joa.12777","article-title":"Olfactory receptor neuronal dendrites become mostly intra-sustentacularly enwrapped upon maturity","volume":"232","author":"Liang","year":"2018","journal-title":"J. Anat."},{"key":"B99","doi-asserted-by":"publisher","first-page":"0266","DOI":"10.1523\/ENEURO.0266-19.2019","article-title":"Sequential maturation of olfactory sensory neurons in the mature olfactory epithelium","volume":"6","author":"Liberia","year":"2019","journal-title":"eNeuro"},{"key":"B100","doi-asserted-by":"publisher","first-page":"689","DOI":"10.2174\/156720511796717212","article-title":"Follow-up study of olfactory deficits, cognitive functions, and volume loss of medial temporal lobe structures in patients with mild cognitive impairment","volume":"8","author":"Lojkowska","year":"2011","journal-title":"Curr. Alzheimer Res."},{"key":"B101","doi-asserted-by":"publisher","first-page":"70","DOI":"10.1016\/j.molmet.2018.03.004","article-title":"Enterochromaffin 5-HT cells - a major target for GLP-1 and gut microbial metabolites","volume":"11","author":"Lund","year":"2018","journal-title":"Mol. Metab."},{"key":"B102","doi-asserted-by":"publisher","first-page":"2449","DOI":"10.1073\/pnas.041604898","article-title":"Activation and targeting of extracellular signal-regulated kinases by beta-arrestin scaffolds","volume":"98","author":"Luttrell","year":"2001","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B103","doi-asserted-by":"publisher","first-page":"713","DOI":"10.1016\/s0092-8674(00)80581-4","article-title":"Combinatorial receptor codes for odors","volume":"96","author":"Malnic","year":"1999","journal-title":"Cell"},{"key":"B104","doi-asserted-by":"publisher","first-page":"114","DOI":"10.1016\/j.vph.2015.08.006","article-title":"Beta2-adrenergic activity modulates vascular tone regulation in lecithin:cholesterol acyltransferase knockout mice","volume":"74","author":"Manzini","year":"2015","journal-title":"Vasc. Pharmacol."},{"key":"B105","doi-asserted-by":"publisher","first-page":"1460","DOI":"10.1111\/j.1476-5381.2011.01429.x","article-title":"Rosuvastatin does not affect human apolipoprotein A-I expression in genetically modified mice: a clue to the disputed effect of statins on HDL","volume":"164","author":"Marchesi","year":"2011","journal-title":"Br. J. Pharmacol."},{"key":"B106","doi-asserted-by":"publisher","first-page":"243","DOI":"10.1016\/s0092-8674(02)00700-6","article-title":"Representation of the glomerular olfactory map in the Drosophila brain","volume":"109","author":"Marin","year":"2002","journal-title":"Cell"},{"key":"B107","doi-asserted-by":"publisher","first-page":"368","DOI":"10.1016\/s0165-6147(00)01678-3","article-title":"G-protein-coupled receptors and signaling networks: emerging paradigms","volume":"22","author":"Marinissen","year":"2001","journal-title":"Trends Pharmacol. Sci."},{"key":"B108","doi-asserted-by":"publisher","first-page":"9902","DOI":"10.1523\/JNEUROSCI.2897-06.2006","article-title":"Beta-arrestin2-mediated internalization of mammalian odorant receptors","volume":"26","author":"Mashukova","year":"2006","journal-title":"J. Neurosci."},{"key":"B109","doi-asserted-by":"publisher","first-page":"19","DOI":"10.1016\/j.neuroscience.2026.01.039","article-title":"Nose-to-brain axis: mechanistic links between nasal microbiome dysbiosis, neuroinflammation, and brain disorders","volume":"598","author":"Mathias","year":"2026","journal-title":"Neuroscience"},{"key":"B110","doi-asserted-by":"publisher","first-page":"16","DOI":"10.3390\/metabo14010016","article-title":"The role of the olfactory System in obesity and metabolism in humans: a systematic review and meta-analysis","volume":"14","author":"Matiashova","year":"2023","journal-title":"Metabolites"},{"key":"B111","doi-asserted-by":"publisher","first-page":"1739","DOI":"10.1152\/physrev.00013.2017","article-title":"Human olfactory receptors: novel cellular functions outside of the nose","volume":"98","author":"Ma\u00dfberg","year":"2018","journal-title":"Physiol. Rev."},{"key":"B112","doi-asserted-by":"publisher","first-page":"1038","DOI":"10.1161\/CIRCRESAHA.119.315175","article-title":"Migratory and dancing macrophage subsets in atherosclerotic lesions","volume":"125","author":"McArdle","year":"2019","journal-title":"Circ. Res."},{"key":"B113","doi-asserted-by":"publisher","first-page":"0","DOI":"10.1152\/physiol.00037.2024","article-title":"Olfactory development and dysfunction: involvement of Microglia","volume":"40","author":"Meller","year":"2025","journal-title":"Physiol. (Bethesda)"},{"key":"B114","doi-asserted-by":"publisher","first-page":"77","DOI":"10.1016\/j.tins.2013.11.004","article-title":"Aging in the olfactory system","volume":"37","author":"Mobley","year":"2014","journal-title":"Trends Neurosci."},{"key":"B115","doi-asserted-by":"publisher","first-page":"5791","DOI":"10.1021\/bi00185a017","article-title":"Fluorescence studies of the location and membrane accessibility of the palmitoylation sites of rhodopsin","volume":"33","author":"Moench","year":"1994","journal-title":"Biochemistry"},{"key":"B116","doi-asserted-by":"publisher","first-page":"e152373","DOI":"10.1172\/JCI152373","article-title":"Platelet olfactory receptor activation limits platelet reactivity and growth of aortic aneurysms","volume":"132","author":"Morrell","year":"2022","journal-title":"J. Clin. Invest"},{"key":"B117","doi-asserted-by":"publisher","first-page":"eabb1817","DOI":"10.1126\/sciimmunol.abb1817","article-title":"T cell engagement of cross-presenting microglia protects the brain from a nasal virus infection","volume":"5","author":"Moseman","year":"2020","journal-title":"Sci. Immunol."},{"key":"B118","doi-asserted-by":"publisher","first-page":"234","DOI":"10.3390\/microorganisms14010234","article-title":"Clinical and immunological perspectives on the nasal microbiome\u2019s role in olfactory function and dysfunction","volume":"14","author":"Mukhtar","year":"2026","journal-title":"Microorganisms"},{"key":"B119","doi-asserted-by":"publisher","first-page":"736","DOI":"10.1038\/srep00736","article-title":"Rapid Nipah virus entry into the central nervous system of hamsters via the olfactory route","volume":"2","author":"Munster","year":"2012","journal-title":"Sci. Rep."},{"key":"B120","doi-asserted-by":"publisher","first-page":"e45266","DOI":"10.1371\/journal.pone.0045266","article-title":"Neuropeptide Y enhances olfactory mucosa responses to odorant in hungry rats","volume":"7","author":"Negroni","year":"2012","journal-title":"PLoS One"},{"key":"B121","doi-asserted-by":"publisher","first-page":"16218","DOI":"10.1074\/jbc.M109.012096","article-title":"Activation of an olfactory receptor inhibits proliferation of prostate cancer cells","volume":"284","author":"Neuhaus","year":"2009","journal-title":"J. Biol. Chem."},{"key":"B122","doi-asserted-by":"publisher","first-page":"107","DOI":"10.1186\/1479-7364-4-2-107","article-title":"Evolutionary dynamics of olfactory receptor genes in chordates: interaction between environments and genomic contents","volume":"4","author":"Niimura","year":"2009","journal-title":"Hum. Genomics"},{"key":"B123","doi-asserted-by":"publisher","first-page":"e44278","DOI":"10.7554\/eLife.44278","article-title":"Anatomical basis and physiological role of cerebrospinal fluid transport through the murine cribriform plate","volume":"8","author":"Norwood","year":"2019","journal-title":"Elife"},{"key":"B124","doi-asserted-by":"publisher","first-page":"685","DOI":"10.1091\/mbc.12.3.685","article-title":"Segregation of heterotrimeric G proteins in cell surface microdomains. G(q) binds caveolin to concentrate in caveolae, whereas g(i) and G (s) target lipid rafts by default","volume":"12","author":"Oh","year":"2001","journal-title":"Mol. Biol. Cell"},{"key":"B125","doi-asserted-by":"publisher","first-page":"42","DOI":"10.1186\/s12862-020-01607-6","article-title":"A unified nomenclature for vertebrate olfactory receptors","volume":"20","author":"Olender","year":"2020","journal-title":"BMC Evol. Biol."},{"key":"B126","doi-asserted-by":"publisher","first-page":"e155506","DOI":"10.1172\/JCI155506","article-title":"Aging-related olfactory loss is associated with olfactory stem cell transcriptional alterations in humans","volume":"132","author":"Oliva","year":"2022","journal-title":"J. Clin. Invest"},{"key":"B127","doi-asserted-by":"publisher","first-page":"W507","DOI":"10.1093\/nar\/gkae305","article-title":"Pred-O3, a web server to predict molecules, olfactory receptors and odor relationships","volume":"52","author":"Ollitrault","year":"2024","journal-title":"Nucleic Acid. Res."},{"key":"B128","doi-asserted-by":"publisher","first-page":"496","DOI":"10.1007\/s10519-009-9289-5","article-title":"Odor identification deficit as a predictor of five-year global cognitive change: interactive effects with age and ApoE-epsilon4","volume":"39","author":"Olofsson","year":"2009","journal-title":"Behav. Genet."},{"key":"B129","doi-asserted-by":"publisher","first-page":"214","DOI":"10.1126\/science.abg3067","article-title":"Olfactory receptor 2 in vascular macrophages drives atherosclerosis by NLRP3-dependent IL-1 production","volume":"375","author":"Orecchioni","year":"2022","journal-title":"Science"},{"key":"B130","doi-asserted-by":"publisher","first-page":"731","DOI":"10.1001\/archotol.1992.01880070061012","article-title":"Human olfactory biopsy. The influence of age and receptor distribution","volume":"118","author":"Paik","year":"1992","journal-title":"Arch. Otolaryngol. Head. Neck Surg."},{"key":"B131","doi-asserted-by":"publisher","first-page":"bjab045","DOI":"10.1093\/chemse\/bjab045","article-title":"Olfactory loss and aging: connections with health and well-being","volume":"46","author":"Papazian","year":"2021","journal-title":"Chem. Senses"},{"key":"B132","doi-asserted-by":"publisher","first-page":"167761","DOI":"10.1016\/j.bbadis.2025.167761","article-title":"Sepsis and high-density lipoproteins: pathophysiology and potential new therapeutic targets","volume":"1871","author":"Parolini","year":"","journal-title":"Biocheim Biophys. Acta Mol. Basis Dis."},{"key":"B133","doi-asserted-by":"publisher","first-page":"116844","DOI":"10.1016\/j.bcp.2025.116844","article-title":"Pathophysiology of bone remodelling cycle: role of immune system and lipids","volume":"235","author":"Parolini","year":"","journal-title":"Biochem. Pharmacol."},{"key":"B134","doi-asserted-by":"publisher","first-page":"47174","DOI":"10.31083\/FBL47174","article-title":"The microbiome epoch: cracking the cardiovascular-neurodegenerative disorder code","volume":"31","author":"Parolini","year":"2026","journal-title":"Front. Biosci. (Landmark Ed.)"},{"key":"B135","doi-asserted-by":"publisher","first-page":"1363","DOI":"10.1111\/j.1478-3231.2012.02855.x","article-title":"Reduced biliary sterol output with no change in total faecal excretion in mice expressing a human apolipoprotein A-I variant","volume":"32","author":"Parolini","year":"2012","journal-title":"Liver Int."},{"key":"B136","doi-asserted-by":"publisher","DOI":"10.2310\/7290.2014.00012","article-title":"Magnetic resonance imaging visualization of vulnerable atherosclerotic plaques at the brachiocephalic artery of apolipoprotein E knockout mice by the blood-pool contrast agent B22956\/1","volume":"13","author":"Parolini","year":"2014","journal-title":"Mol. Imaging"},{"key":"B137","doi-asserted-by":"publisher","first-page":"253","DOI":"10.1038\/s41586-020-2451-1","article-title":"Structure and flexibility in cortical representations of odour space","volume":"583","author":"Pashkovski","year":"2020","journal-title":"Nature"},{"key":"B138","doi-asserted-by":"publisher","first-page":"283","DOI":"10.1002\/mds.870100309","article-title":"The anterior olfactory nucleus in Parkinson\u2019s disease","volume":"10","author":"Pearce","year":"1995","journal-title":"Mov. Disord."},{"key":"B139","doi-asserted-by":"publisher","first-page":"2550","DOI":"10.1002\/glia.23870","article-title":"Insights into olfactory ensheathing cell development from a laser-microdissection and transcriptome-profiling approach","volume":"68","author":"Perera","year":"2020","journal-title":"Glia"},{"key":"B140","doi-asserted-by":"publisher","first-page":"87","DOI":"10.1007\/s12016-012-8343-x","article-title":"Smell and autoimmunity: a comprehensive review","volume":"45","author":"Perricone","year":"2013","journal-title":"Clin. Rev Allergy Immunol."},{"key":"B141","doi-asserted-by":"publisher","first-page":"1249","DOI":"10.1136\/gut.33.9.1249","article-title":"Portal and peripheral blood short chain fatty acid concentrations after caecal lactulose instillation at surgery","volume":"33","author":"Peters","year":"1992","journal-title":"Gut"},{"key":"B142","doi-asserted-by":"publisher","first-page":"406","DOI":"10.1002\/cne.23686","article-title":"Novel role of cystic fibrosis transmembrane conductance regulator in maintaining adult mouse olfactory neuronal homeostasis","volume":"523","author":"Pfister","year":"2015","journal-title":"J. Comp. Neurol."},{"key":"B143","doi-asserted-by":"publisher","first-page":"e53462","DOI":"10.1371\/journal.pone.0053462","article-title":"Bioluminescent imaging and histopathologic characterization of WEEV neuroinvasion in outbred CD-1 mice","volume":"8","author":"Phillips","year":"2013","journal-title":"PLoS One"},{"key":"B144","doi-asserted-by":"publisher","first-page":"1943","DOI":"10.3390\/genes12121943","article-title":"High olfactory receptor-rich 11q11 copy number in girls and African American Children","volume":"12","author":"Phillips","year":"2021","journal-title":"Genes (Basel)"},{"key":"B145","doi-asserted-by":"publisher","first-page":"2059","DOI":"10.1073\/pnas.0812859106","article-title":"Functional expression of the olfactory signaling system in the kidney","volume":"106","author":"Pluznick","year":"2009","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B146","doi-asserted-by":"publisher","first-page":"4410","DOI":"10.1073\/pnas.1215927110","article-title":"Olfactory receptor responding to gut microbiota-derived signals plays a role in renin secretion and blood pressure regulation","volume":"110","author":"Pluznick","year":"2013","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B147","doi-asserted-by":"publisher","first-page":"17013","DOI":"10.1038\/nrdp.2017.13","article-title":"Parkinson disease","volume":"3","author":"Poewe","year":"2017","journal-title":"Nat. Rev. Dis. Prim."},{"key":"B148","doi-asserted-by":"publisher","first-page":"850","DOI":"10.1016\/j.neuron.2009.05.022","article-title":"Odor representations in olfactory cortex: \u201csparse\u201d coding, global inhibition, and oscillations","volume":"62","author":"Poo","year":"2009","journal-title":"Neuron"},{"key":"B149","doi-asserted-by":"publisher","first-page":"1287","DOI":"10.1016\/j.neuroscience.2009.05.043","article-title":"Nutritional status modulates behavioural and olfactory bulb Fos responses to isoamyl acetate or food odour in rats: roles of orexins and leptin","volume":"162","author":"Prud'homme","year":"2009","journal-title":"Neuroscience"},{"key":"B150","doi-asserted-by":"publisher","first-page":"1497","DOI":"10.3233\/JAD-160842","article-title":"Odor identification screening improves diagnostic classification in incipient Alzheimer's Disease","volume":"55","author":"Quarmley","year":"2017","journal-title":"J. Alzheimers Dis."},{"key":"B151","doi-asserted-by":"publisher","first-page":"5424","DOI":"10.1080\/10408398.2021.1885007","article-title":"Human ectopic olfactory receptors and their food originated ligands: a review","volume":"62","author":"Raka","year":"2022","journal-title":"Crit. Rev. Food Sci. Nutr."},{"key":"B152","doi-asserted-by":"publisher","first-page":"141","DOI":"10.3389\/fcvm.2019.00141","article-title":"Shear-Dependent platelet aggregation: mechanisms and therapeutic opportunities","volume":"6","author":"Rana","year":"2019","journal-title":"Front. Cardiovasc Med."},{"key":"B153","doi-asserted-by":"publisher","first-page":"9","DOI":"10.12688\/wellcomeopenres.10646.1","article-title":"Revisiting olfactory receptors as putative drivers of cancer","volume":"2","author":"Ranzani","year":"2017","journal-title":"Wellcome Open Res."},{"key":"B154","doi-asserted-by":"publisher","first-page":"318","DOI":"10.3389\/fnins.2015.00318","article-title":"Impaired sense of smell and altered olfactory system in RAG-1(-\u2215-) immunodeficient mice","volume":"9","author":"Rattazzi","year":"2015","journal-title":"Front. Neurosci."},{"key":"B155","doi-asserted-by":"publisher","first-page":"1143","DOI":"10.1016\/j.otc.2004.06.003","article-title":"Chronic sinusitis and olfactory dysfunction","volume":"37","author":"Raviv","year":"2004","journal-title":"Otolaryngol. Clin. North Am."},{"key":"B156","doi-asserted-by":"publisher","first-page":"553","DOI":"10.1016\/j.neuron.2005.01.012","article-title":"Mechanism of the excitatory cl\u2212 response in Mouse Olfactory receptor neurons","volume":"45","author":"Reisert","year":"2005","journal-title":"Neuron"},{"key":"B157","doi-asserted-by":"publisher","first-page":"117403","DOI":"10.1016\/j.biopha.2024.117403","article-title":"Ecnomotopic olfactory receptors in metabolic regulation","volume":"179","author":"Ren","year":"2024","journal-title":"Biomed. Pharmacother."},{"key":"B158","doi-asserted-by":"publisher","first-page":"226","DOI":"10.1016\/j.nbd.2016.12.013","article-title":"The olfactory bulb as the entry site for prion-like propagation in neurodegenerative diseases","volume":"109","author":"Rey","year":"2018","journal-title":"Neurobiol. Dis."},{"key":"B159","doi-asserted-by":"publisher","first-page":"377","DOI":"10.1016\/j.bbalip.2013.09.001","article-title":"Fatty aldehyde and fatty alcohol metabolism: review and importance for epidermal structure and function","volume":"1814","author":"Rizzo","year":"2014","journal-title":"Biochim. Biophys. Acta"},{"key":"B160","doi-asserted-by":"publisher","first-page":"13","DOI":"10.4193\/Rhino11.126","article-title":"Assessment of chemosensory function using electroencephalographic techniques","volume":"50","author":"Rombaux","year":"2012","journal-title":"Rhinology"},{"key":"B161","doi-asserted-by":"publisher","first-page":"566","DOI":"10.1016\/j.cell.2016.02.063","article-title":"Asprosin, a fasting induced glucogenic protein hormone","volume":"165","author":"Romere","year":"2016","journal-title":"Cell"},{"key":"B162","doi-asserted-by":"publisher","first-page":"449","DOI":"10.1016\/j.stem.2019.09.006","article-title":"Smelling danger: olfactory stem cells control immune defense during chronic inflammation","volume":"25","author":"Rustenhoven","year":"2019","journal-title":"Cell Stem Cell"},{"key":"B163","doi-asserted-by":"publisher","first-page":"eaax0396","DOI":"10.1126\/sciadv.aax0396","article-title":"A transcriptomic atlas of mammalian olfactory mucosae reveals an evolutionary influence on food odor detection in humans","volume":"5","author":"Saraiva","year":"2019","journal-title":"Sci. Adv."},{"key":"B164","doi-asserted-by":"publisher","first-page":"a011148","DOI":"10.1101\/cshperspect.a011148","article-title":"The cyclic AMP pathway","volume":"4","author":"Sassone-Corsi","year":"2012","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"B165","doi-asserted-by":"publisher","first-page":"2898","DOI":"10.1152\/jn.91169.2008","article-title":"Modulation of spontaneous and odorant-evoked activity of rat olfactory sensory neurons by two anorectic peptides, insulin and leptin","volume":"101","author":"Savigner","year":"2009","journal-title":"J. Neurophysiol."},{"key":"B166","doi-asserted-by":"publisher","first-page":"e326591","DOI":"10.1161\/CIRCRESAHA.125.326591","article-title":"Olfr2 promotes recruitment of monocytes via CX3CR1 in abdominal aortic aneurysm","volume":"138","author":"Schelemei","year":"2026","journal-title":"Circ. Res."},{"key":"B167","doi-asserted-by":"publisher","first-page":"731","DOI":"10.1001\/jamaoto.2016.0927","article-title":"Mucous cytokine levels in chronic rhinosinusitis-associated olfactory loss","volume":"142","author":"Schlosser","year":"2016","journal-title":"JAMA Otolaryngol. Head. Neck Surg."},{"key":"B168","doi-asserted-by":"publisher","first-page":"3975","DOI":"10.1128\/JVI.06828-11","volume":"86","author":"Schrauwen","year":"2012","journal-title":"J. Virol."},{"key":"B169","doi-asserted-by":"publisher","first-page":"710","DOI":"10.1093\/gerona\/glw036","article-title":"Sensory impairments and risk of mortality in older adults","volume":"72","author":"Schubert","year":"2017","journal-title":"J. Gerontol. A Biol. Sci. Med. Sci."},{"key":"B170","doi-asserted-by":"publisher","first-page":"32","DOI":"10.3389\/fgene.2013.00032","article-title":"Sex bias in copy number variation of olfactory receptor gene family depends on ethnicity","volume":"4","author":"Shadravan","year":"2013","journal-title":"Front. Genet."},{"key":"B171","doi-asserted-by":"publisher","first-page":"891","DOI":"10.2174\/1570159X17666181206095626","article-title":"Sense of smell: structural, functional, mechanistic advancements and challenges in human olfactory research","volume":"17","author":"Sharma","year":"2019","journal-title":"Curr. Neuropharmacol."},{"key":"B172","doi-asserted-by":"publisher","first-page":"D678","DOI":"10.1093\/nar\/gkab763","article-title":"OlfactionBase: a repository to explore odors, odorants, olfactory receptors and odorant-receptor interactions","volume":"50","author":"Sharma","year":"2022","journal-title":"Nucleic Acid. Res."},{"key":"B173","doi-asserted-by":"publisher","first-page":"1615","DOI":"10.1007\/s11010-024-05024-x","article-title":"New insights into the roles of olfactory receptors in cardiovascular disease","volume":"479","author":"Shi","year":"2024","journal-title":"Mol. Cell Biochem."},{"key":"B174","doi-asserted-by":"publisher","first-page":"703","DOI":"10.1093\/gbe\/evs039","article-title":"Transcriptomes of mouse olfactory epithelium reveal sexual differences in odorant detection","volume":"4","author":"Shiao","year":"2012","journal-title":"Genome Biol. Evol."},{"key":"B175","doi-asserted-by":"publisher","first-page":"77","DOI":"10.1016\/j.neulet.2009.02.006","article-title":"Regional differences in the severity of Lewy body pathology across the olfactory cortex","volume":"453","author":"Silveira-Moriyama","year":"2009","journal-title":"Neurosci. Lett."},{"key":"B176","doi-asserted-by":"publisher","first-page":"17","DOI":"10.1016\/B978-0-444-63855-7.00002-2","article-title":"Anatomy of the olfactory system","volume":"164","author":"Smith","year":"2019","journal-title":"Handb. Clin. Neurol."},{"key":"B177","doi-asserted-by":"publisher","first-page":"135","DOI":"10.3233\/JAD-2009-1020","article-title":"Olfactory dysfunction is associated with subjective memory complaints in community-dwelling elderly individuals","volume":"17","author":"Sohrabi","year":"2009","journal-title":"J. Alzheimers Dis."},{"key":"B178","doi-asserted-by":"publisher","first-page":"e13382","DOI":"10.1111\/jne.13382","article-title":"The olfactory bulb: a neuroendocrine spotlight on feeding and metabolism","volume":"36","author":"Stark","year":"2024","journal-title":"J. Neuroendocrinol."},{"key":"B179","doi-asserted-by":"publisher","first-page":"11776","DOI":"10.1073\/pnas.0903304106","article-title":"ANO2 is the cilial calcium-activated chloride channel that may mediate olfactory amplification","volume":"106","author":"Stephan","year":"2009","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B180","doi-asserted-by":"publisher","first-page":"279","DOI":"10.1016\/0165-3806(92)90207-d","article-title":"The role of olfactory bulb norepinephrine in early olfactory learning","volume":"70","author":"Sullivan","year":"1992","journal-title":"Brain Rev. Dev. Brain Rev."},{"key":"B181","doi-asserted-by":"publisher","first-page":"5205","DOI":"10.1038\/ncomms6205","article-title":"Nasal immunity is an ancient arm of the mucosal immune system of vertebrates","volume":"5","author":"Tacchi","year":"2014","journal-title":"Nat. Commun."},{"key":"B182","doi-asserted-by":"publisher","first-page":"115954","DOI":"10.1016\/j.ab.2025.115954","article-title":"A sensitive split luciferase assay for detecting olfactory receptor response to odorants","volume":"707","author":"Tanazawa","year":"2025","journal-title":"Anal. Biochem."},{"key":"B183","doi-asserted-by":"publisher","first-page":"1405","DOI":"10.3390\/microorganisms10071405","article-title":"Nasal Microbiota, olfactory health, neurological disorders and Aging-A review","volume":"10","author":"Thangaleela","year":"2022","journal-title":"Microorganisms"},{"key":"B184","doi-asserted-by":"publisher","first-page":"1275","DOI":"10.3233\/JAD-210636","article-title":"Olfaction, cognitive impairment, and PET biomarkers in community-dwelling older adults","volume":"86","author":"Tian","year":"2022","journal-title":"J. Alzheimers Dis."},{"key":"B185","doi-asserted-by":"publisher","first-page":"9471","DOI":"10.1038\/s41598-017-10379-x","article-title":"Olfactory receptor 10J5 responding to \u03b1-cedrene regulates hepatic steatosis via the cAMP-PKA pathway","volume":"7","author":"Tong","year":"2017","journal-title":"Sci. Rep."},{"key":"B186","doi-asserted-by":"publisher","first-page":"202","DOI":"10.1038\/s41366-018-0176-0","article-title":"\u03b1-Cedrene protects rodents from high-fat diet-induced adiposity via adenylyl cyclase 3","volume":"43","author":"Tong","year":"2019","journal-title":"Int. J. Obes. (Lond)."},{"key":"B187","doi-asserted-by":"publisher","first-page":"1909","DOI":"10.1021\/acschemneuro.0c00249","article-title":"Proinflammatory cytokines in the olfactory Mucosa result in COVID-19 induced anosmia","volume":"11","author":"Torabi","year":"2020","journal-title":"ACS Chem. Neurosci."},{"key":"B188","doi-asserted-by":"publisher","first-page":"307","DOI":"10.1146\/annurev.physiol.010908.163209","article-title":"Sensing odorants and pheromones with chemosensory receptors","volume":"71","author":"Touhara","year":"2009","journal-title":"Annu. Rev. Ohysiol"},{"key":"B189","doi-asserted-by":"publisher","first-page":"9475","DOI":"10.1073\/pnas.1804106115","article-title":"Genetic variation across the human olfactory receptor repertoire alters odor perception","volume":"116","author":"Trimmer","year":"2019","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B190","doi-asserted-by":"publisher","first-page":"151","DOI":"10.1007\/978-1-0716-2364-0_11","article-title":"Isolation, ex vivo culture, and stimulation of tracheal and nasal chemosensory cells","volume":"2505","author":"Ualiyeva","year":"2022","journal-title":"Methods Mol. Biol."},{"key":"B191","doi-asserted-by":"publisher","first-page":"1","DOI":"10.15761\/JTS.1000470","article-title":"How smell regulates metabolism: the role of ectopically expressed olfactory receptors in lipid and glucose homeostasis","volume":"8","author":"Urbani","year":"2022","journal-title":"J. Transl. Sci."},{"key":"B192","doi-asserted-by":"publisher","first-page":"140","DOI":"10.3389\/fnins.2020.00140","article-title":"Mechanisms linking olfactory impairment and risk of mortality","volume":"14","author":"Van Regemorter","year":"2020","journal-title":"Front. Neurosci."},{"key":"B193","doi-asserted-by":"publisher","first-page":"2459","DOI":"10.1007\/s12035-024-04375-2","article-title":"Is nasal dysbiosis a required component for neuroinflammation in major depressive disorder?","volume":"62","author":"V\u00e1squez-P\u00e9rez","year":"2025","journal-title":"Mol. Neurobiol."},{"key":"B194","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1016\/j.parkreldis.2022.09.007","article-title":"Serial olfactory testing for the diagnosis of prodromal Parkinson\u2019s disease in the PARS study","volume":"104","author":"Vaswani","year":"2022","journal-title":"Park. Relat. Disord."},{"key":"B195","doi-asserted-by":"publisher","first-page":"e96333","DOI":"10.1371\/journal.pone.0096333","article-title":"Profiling of olfactory receptor gene expression in whole human olfactory mucosa","volume":"9","author":"Verbeurgt","year":"2014","journal-title":"PLoS One"},{"key":"B196","doi-asserted-by":"publisher","first-page":"e160277","DOI":"10.1172\/jci.insight.160277","article-title":"SARS-CoV-2 infection of sustentacular cells disrupts olfactory signaling pathways","volume":"7","author":"Verma","year":"2022","journal-title":"JCI Insight"},{"key":"B197","doi-asserted-by":"publisher","first-page":"585","DOI":"10.3233\/JAD-201168","article-title":"Olfactory dysfunction is already present with subjective cognitive decline and deepens with disease severity in the Alzheimer\u2019s disease spectrum","volume":"79","author":"Wang","year":"2021","journal-title":"J. Alzheimers Dis."},{"key":"B198","doi-asserted-by":"publisher","first-page":"e10321","DOI":"10.1002\/advs.202510321","article-title":"Inhaling eugenol inhibits NAFLD by activating the hepatic ectopic olfactory receptor Olfr544 and modulating the gut microbiota","volume":"12","author":"Wang","year":"2025","journal-title":"Adv. Sci. (Weinh)"},{"key":"B199","doi-asserted-by":"publisher","first-page":"33","DOI":"10.3389\/fonc.2018.00033","article-title":"Olfactory receptors as biomarkers in human breast carcinoma tissues","volume":"8","author":"Weber","year":"2018","journal-title":"Front. Oncol."},{"key":"B200","doi-asserted-by":"publisher","first-page":"2118","DOI":"10.1016\/j.immuni.2022.08.017","article-title":"Mucosal plasma cells are required to protect the upper airway and brain from infection","volume":"55","author":"Wellford","year":"2022","journal-title":"Immunity"},{"key":"B201","doi-asserted-by":"publisher","first-page":"3698","DOI":"10.1073\/pnas.0812874106","article-title":"Metabolomics analysis reveals large effects of gut microflora on mammalian blood metabolites","volume":"106","author":"Wikoff","year":"2009","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B202","doi-asserted-by":"publisher","first-page":"63","DOI":"10.1093\/chemse\/bjq098","article-title":"Odor identification and mortality in old age","volume":"36","author":"Wilson","year":"2011","journal-title":"Chem. Senses"},{"key":"B203","doi-asserted-by":"publisher","first-page":"103450","DOI":"10.1016\/j.mcn.2019.103450","article-title":"Macrophage migration inhibitory factor and its binding partner HTRA1 are expressed by olfactory ensheathing cells","volume":"102","author":"Wright","year":"2020","journal-title":"Mol. Cell Neurosci."},{"key":"B204","doi-asserted-by":"publisher","first-page":"75","DOI":"10.1016\/j.biocel.2015.03.008","article-title":"Activation of OR1A1 suppresses PPAR-\u03b3 expression by inducing HES-1 in cultured hepatocytes","volume":"64","author":"Wu","year":"2015","journal-title":"Int. J. Biochem. Cell Biol."},{"key":"B205","doi-asserted-by":"publisher","first-page":"4118","DOI":"10.1172\/JCI89344","article-title":"Olfactory receptor 544 reduces adiposity by steering fuel preference toward fats","volume":"127","author":"Wu","year":"2017","journal-title":"J. Clin. Investig."},{"key":"B206","doi-asserted-by":"publisher","first-page":"489","DOI":"10.1016\/j.bbalip.2019.01.004","article-title":"Olfactory receptor 43 reduces hepatic lipid accumulation and adiposity in mice","volume":"1864","author":"Wu","year":"2019","journal-title":"Biochim. Biophys. Acta Mol. Cell Biol. Lipids"},{"key":"B207","doi-asserted-by":"publisher","first-page":"1987782","DOI":"10.1080\/19490976.2021.1987782","article-title":"Activation of ectopic olfactory receptor 544 induces GLP-1 secretion and regulates gut inflammation","volume":"13","author":"Wu","year":"2021","journal-title":"Gut Microbes"},{"key":"B208","doi-asserted-by":"publisher","first-page":"17170","DOI":"10.1038\/s41598-018-35102-2","article-title":"The human olfactory cleft mucus proteome and its age-related changes","volume":"8","author":"Yoshikawa","year":"2018","journal-title":"Sci. Rep."},{"key":"B209","doi-asserted-by":"publisher","first-page":"e2300372","DOI":"10.1002\/mnfr.202300372","article-title":"Inulin supplementation modulates the hepatic transcriptome, metabolome, and ferritin content in ovariectomized rats","volume":"67","author":"Zhao","year":"2023","journal-title":"Mol. Nutr. Food Res."}],"container-title":["Frontiers in Pharmacology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fphar.2026.1905572\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T09:16:27Z","timestamp":1788945387000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fphar.2026.1905572\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,9]]},"references-count":209,"alternative-id":["10.3389\/fphar.2026.1905572"],"URL":"https:\/\/doi.org\/10.3389\/fphar.2026.1905572","relation":{},"ISSN":["1663-9812"],"issn-type":[{"value":"1663-9812","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,9]]},"article-number":"1905572"}}