{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,9]],"date-time":"2026-11-09T22:35:14Z","timestamp":1794263714868,"version":"4.9.0"},"reference-count":49,"publisher":"Oxford University Press (OUP)","license":[{"start":{"date-parts":[[2023,1,1]],"date-time":"2023-01-01T00:00:00Z","timestamp":1672531200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/pages\/standard-publication-reuse-rights"}],"funder":[{"DOI":"10.13039\/100000055","name":"National Institute on Deafness and Other Communication Disorders","doi-asserted-by":"publisher","award":["1R21DC016990-01"],"award-info":[{"award-number":["1R21DC016990-01"]}],"id":[{"id":"10.13039\/100000055","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100017653","name":"Eastern Michigan University","doi-asserted-by":"publisher","award":["1R21DC016990-01"],"award-info":[{"award-number":["1R21DC016990-01"]}],"id":[{"id":"10.13039\/100017653","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2023,1,1]]},"abstract":"<jats:title>Abstract<\/jats:title><jats:p>In behavioral experiments, rats perceive sodium carbonate (Na2CO3) as super salty. In fact, when the dissociated Na+ ions are accounted for, rats perceive Na2CO3 as 5\u00d7 saltier than equinormal concentrations of NaCl. The chorda tympani nerve (CT) responds to salts through at least two receptor mechanisms and is a model system for understanding how salt taste is transmitted to the brain. Here, we recorded CT nerve activity to a broad range of NaCl (3\u2013300 mM) and Na2CO3 (3\u2013300 mN) to investigate why Na2CO3 tastes so salty to rats. Benzamil, a specific epithelial sodium channel (ENaC) antagonist, was used to determine the relative contribution of apical ENaCs in Na2CO3 transduction. The benzamil-insensitive component of CT nerve responses was enhanced by increasing the adapted tongue temperature from 23\u00b0C to 30\u00b0C. Na2CO3 solutions are alkaline, so we compared neural responses (with and without benzamil) to 100 mM NaCl alone (6.2 pH) and at a pH (11.2 pH) that matched 100 mN Na2CO3. As expected, NaCl responses increased progressively with increasing concentration and temperature. Responses to 3 mN Na2CO3 were greater than 3 mM NaCl with and without benzamil, but the shape of the first log-fold range of was relatively flat. Adjusting the pH of NaCl to 11.2 abolished the thermal enhancement of 100 mN NaCl through the benzamil-insensitive pathway. Rinsing Na2CO3 off the tongue resulted in robust aftertaste that was concentration dependent, thermally sensitive, and benzamil-insensitive. Responses to alkaline NaCl did not recapitulate Na2CO3 responses or aftertaste, suggesting multiple transduction mechanisms for the cations (2Na+) and anion (CO3-2).<\/jats:p>","DOI":"10.1093\/chemse\/bjad015","type":"journal-article","created":{"date-parts":[[2023,5,24]],"date-time":"2023-05-24T22:04:09Z","timestamp":1684965849000},"source":"Crossref","is-referenced-by-count":1,"title":["Analysis of the rat chorda tympani nerve response to \u201csuper salty\u201d sodium carbonate"],"prefix":"10.1093","volume":"48","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-6729-4539","authenticated-orcid":false,"given":"Joseph M","family":"Breza","sequence":"first","affiliation":[{"name":"Department of Psychology, Program in Neuroscience, Eastern Michigan University , Ypsilanti, MI 48197 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0001-0479-8940","authenticated-orcid":false,"given":"Steven J","family":"St. John","sequence":"additional","affiliation":[{"name":"Department of Psychology, Rollins College , Winter Park, FL , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2023,5,24]]},"reference":[{"key":"2023081009313744500_CIT0001","doi-asserted-by":"crossref","first-page":"7795","DOI":"10.1523\/JNEUROSCI.0597-20.2020","article-title":"Optogenetic stimulation of type I GAD65(+) cells in taste buds activates gustatory neurons and drives appetitive licking behavior in sodium-depleted mice","volume":"40","author":"Baumer-Harrison","year":"2020","journal-title":"J Neurosci"},{"key":"2023081009313744500_CIT0002","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1085\/jgp.38.2.133","article-title":"A theory of taste stimulation","volume":"38","author":"Beidler","year":"1954","journal-title":"J Gen Physiol"},{"key":"2023081009313744500_CIT0003","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1111\/j.2517-6161.1995.tb02031.x","article-title":"Controlling the false discovery rate: a practical and powerful approach to multiple testing","volume":"57","author":"Benjamini","year":"1995","journal-title":"J R Stat Soc Series B Methodol"},{"key":"2023081009313744500_CIT0004","doi-asserted-by":"crossref","first-page":"R1027","DOI":"10.1152\/ajpregu.00843.2009","article-title":"Learning-based recovery from perceptual impairment in salt discrimination after permanently altered peripheral gustatory input","volume":"299","author":"Blonde","year":"2010","journal-title":"Am J Physiol Regul Integr Comp Physiol"},{"key":"2023081009313744500_CIT0005","doi-asserted-by":"crossref","first-page":"131","DOI":"10.1093\/chemse\/8.2.131","article-title":"Rat neurophysiological taste responses to salt solutions","volume":"8","author":"Boudreau","year":"1983","journal-title":"Chem Senses"},{"key":"2023081009313744500_CIT0006","doi-asserted-by":"crossref","first-page":"2405","DOI":"10.1152\/jn.00114.2012","article-title":"Acetic acid modulates spike rate and spike latency to salt in peripheral gustatory neurons of rats","volume":"108","author":"Breza","year":"2012","journal-title":"J Neurophysiol"},{"key":"2023081009313744500_CIT0007","doi-asserted-by":"crossref","first-page":"1632","DOI":"10.1152\/jn.00621.2011","article-title":"Anion size modulates salt taste in rats","volume":"107","author":"Breza","year":"2012","journal-title":"J Neurophysiol"},{"key":"2023081009313744500_CIT0008","doi-asserted-by":"crossref","first-page":"674","DOI":"10.1152\/jn.00793.2005","article-title":"Temperature modulates taste responsiveness and stimulates gustatory neurons in the rat geniculate ganglion","volume":"95","author":"Breza","year":"2006","journal-title":"J Neurophysiol"},{"key":"2023081009313744500_CIT0009","doi-asserted-by":"crossref","first-page":"1771","DOI":"10.1152\/jn.00785.2009","article-title":"Response latency to lingual taste stimulation distinguishes neuron types within the geniculate ganglion","volume":"103","author":"Breza","year":"2010","journal-title":"J Neurophysiol"},{"key":"2023081009313744500_CIT0010","volume-title":"Centers for Disease Control and Prevention","author":"CDC","year":"2020"},{"key":"2023081009313744500_CIT0011","doi-asserted-by":"crossref","first-page":"443","DOI":"10.1126\/science.1174601","article-title":"The taste of carbonation","volume":"326","author":"Chandrashekar","year":"2009","journal-title":"Science"},{"key":"2023081009313744500_CIT0012","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1037\/0033-2909.112.1.155","article-title":"A power primer","volume":"112","author":"Cohen","year":"1992","journal-title":"Psychol Bull"},{"key":"2023081009313744500_CIT0013","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1016\/0006-8993(90)91817-Z","article-title":"The anion in salt taste: a possible role for paracellular pathways","volume":"535","author":"Elliott","year":"1990","journal-title":"Brain Res"},{"key":"2023081009313744500_CIT0014","doi-asserted-by":"crossref","first-page":"1229","DOI":"10.1037\/0735-7044.114.6.1229","article-title":"Sodium taste detectability in rats is independent of anion size: the psychophysical characteristics of the transcellular sodium taste transduction pathway","volume":"114","author":"Geran","year":"2000","journal-title":"Behav Neurosci"},{"key":"2023081009313744500_CIT0015","doi-asserted-by":"crossref","first-page":"779","DOI":"10.1037\/0735-7044.121.4.779","article-title":"Amiloride-insensitive units of the chorda tympani nerve are necessary for normal ammonium chloride detectability in the rat","volume":"121","author":"Geran","year":"2007","journal-title":"Behav Neurosci"},{"key":"2023081009313744500_CIT0016","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1080\/03670244.1973.9990329","article-title":"Blood pressure and dietary salt in human populations","volume":"2","author":"Gleiberman","year":"1973","journal-title":"Ecol Food Nutr"},{"key":"2023081009313744500_CIT0017","doi-asserted-by":"crossref","first-page":"403","DOI":"10.1126\/science.6691151","article-title":"Salt taste transduction occurs through an amiloride-sensitive sodium transport pathway","volume":"223","author":"Heck","year":"1984","journal-title":"Science"},{"key":"2023081009313744500_CIT0018","doi-asserted-by":"crossref","first-page":"934","DOI":"10.1038\/nature05084","article-title":"The cells and logic for mammalian sour taste detection","volume":"442","author":"Huang","year":"2006","journal-title":"Nature"},{"key":"2023081009313744500_CIT0019","doi-asserted-by":"crossref","first-page":"426","DOI":"10.1016\/S0140-6736(87)90127-9","article-title":"The dominance of salt in manufactured food in the sodium intake of affluent societies","volume":"1","author":"James","year":"1987","journal-title":"Lancet"},{"key":"2023081009313744500_CIT0020","volume-title":"Salt: a world history","author":"Kurlansky","year":"2003"},{"key":"2023081009313744500_CIT0021","doi-asserted-by":"crossref","first-page":"1942","DOI":"10.1523\/JNEUROSCI.2947-15.2016","article-title":"Amiloride-insensitive salt taste is mediated by two populations of type III taste cells with distinct transduction mechanisms","volume":"36","author":"Lewandowski","year":"2016","journal-title":"J Neurosci"},{"key":"2023081009313744500_CIT0022","doi-asserted-by":"crossref","first-page":"727","DOI":"10.1093\/chemse\/bjw082","article-title":"Temperature influences chorda tympani nerve responses to sweet, salty, sour, umami, and bitter stimuli in mice","volume":"41","author":"Lu","year":"2016","journal-title":"Chem Senses"},{"key":"2023081009313744500_CIT0023","doi-asserted-by":"crossref","first-page":"309","DOI":"10.1016\/S0006-8993(96)01118-3","article-title":"Temperature and amiloride alter taste nerve responses to Na+, K+, and NH+4 salts in rats","volume":"744","author":"Lundy","year":"1997","journal-title":"Brain Res"},{"key":"2023081009313744500_CIT0024","doi-asserted-by":"crossref","first-page":"2970","DOI":"10.1152\/jn.1999.82.6.2970","article-title":"Gustatory neuron types in rat geniculate ganglion","volume":"82","author":"Lundy","year":"1999","journal-title":"J Neurophysiol"},{"key":"2023081009313744500_CIT0025","doi-asserted-by":"crossref","first-page":"C1005","DOI":"10.1152\/ajpcell.2001.281.3.C1005","article-title":"Decrease in rat taste receptor cell intracellular pH is the proximate stimulus in sour taste transduction","volume":"281","author":"Lyall","year":"2001","journal-title":"Am J Physiol Cell Physiol"},{"key":"2023081009313744500_CIT0026","doi-asserted-by":"crossref","first-page":"793","DOI":"10.1085\/jgp.20028656","article-title":"Modulation of rat chorda tympani NaCl responses and intracellular Na+ activity in polarized taste receptor cells by pH","volume":"120","author":"Lyall","year":"2002","journal-title":"J Gen Physiol"},{"key":"2023081009313744500_CIT0027","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1113\/jphysiol.2004.065656","article-title":"The mammalian amiloride-insensitive non-specific salt taste receptor is a vanilloid receptor-1 variant","volume":"558","author":"Lyall","year":"2004","journal-title":"J Physiol"},{"issue":"1","key":"2023081009313744500_CIT0028","doi-asserted-by":"crossref","first-page":"34","DOI":"10.1037\/h0082791","article-title":"The relative Effectiveness of salt stimuli for the rat","volume":"23","author":"Morrison","year":"1969","journal-title":"Can J Psychol"},{"issue":"1","key":"2023081009313744500_CIT0029","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1016\/0031-9384(72)90124-2","article-title":"Detectability and preference for sodium chloride and sodium carbonate","volume":"8","author":"Morrison","year":"1972","journal-title":"Physiol Behav"},{"issue":"1","key":"2023081009313744500_CIT0030","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1016\/0031-9384(72)90125-4","article-title":"Taste control over sodium intake in sodium deficient rats","volume":"8","author":"Morrison","year":"1972","journal-title":"Physiol Behav"},{"key":"2023081009313744500_CIT0031","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1016\/0006-8993(88)90777-9","article-title":"Amiloride inhibition of responses of rat single chorda tympani fibers to chemical and electrical tongue stimulations","volume":"451","author":"Ninomiya","year":"1988","journal-title":"Brain Res"},{"key":"2023081009313744500_CIT0032","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1113\/jphysiol.1968.sp008650","article-title":"Multiple sensitivity of chordat typani fibres of the rat and hamster to gustatory and thermal stimuli","volume":"199","author":"Ogawa","year":"1968","journal-title":"J Physiol (Lond)"},{"key":"2023081009313744500_CIT0033","doi-asserted-by":"crossref","first-page":"472","DOI":"10.1038\/nature11905","article-title":"High salt recruits aversive taste pathways","volume":"494","author":"Oka","year":"2013","journal-title":"Nature"},{"key":"2023081009313744500_CIT0034","doi-asserted-by":"crossref","first-page":"527","DOI":"10.1016\/S0002-8703(76)80337-7","article-title":"Epidemiologic evidence on the etiology of human hypertension and its possible prevention","volume":"91","author":"Page","year":"1976","journal-title":"Am Heart J"},{"key":"2023081009313744500_CIT0035","doi-asserted-by":"crossref","first-page":"453","DOI":"10.1085\/jgp.101.3.453","article-title":"Anion modulation of taste responses in sodium-sensitive neurons of the hamster chorda tympani nerve","volume":"101","author":"Rehnberg","year":"1993","journal-title":"J Gen Physiol"},{"key":"2023081009313744500_CIT0036","first-page":"R1732","article-title":"Amiloride-sensitive sodium signals and salt appetite: multiple gustatory pathways","volume":"276","author":"Roitman","year":"1999","journal-title":"Am J Physiol"},{"key":"2023081009313744500_CIT0037","doi-asserted-by":"crossref","first-page":"813","DOI":"10.1093\/chemse\/bjl024","article-title":"Detection of NaCl and KCl in TRPV1 knockout mice","volume":"31","author":"Ruiz","year":"2006","journal-title":"Chem Senses"},{"key":"2023081009313744500_CIT0038","doi-asserted-by":"crossref","first-page":"R1195","DOI":"10.1152\/ajpregu.00154.2012","article-title":"Contribution of the TRPV1 channel to salt taste quality in mice as assessed by conditioned taste aversion generalization and chorda tympani nerve responses","volume":"303","author":"Smith","year":"2012","journal-title":"Am J Physiol Regul Integr Comp Physiol"},{"key":"2023081009313744500_CIT0039","first-page":"R169","article-title":"Salt taste discrimination after bilateral section of the chorda tympani or glossopharyngeal nerves","volume":"263","author":"Spector","year":"1992","journal-title":"Am J Physiol"},{"key":"2023081009313744500_CIT0040","first-page":"R820","article-title":"Chemospecific deficits in taste detection after selective gustatory deafferentation in rats","volume":"258","author":"Spector","year":"1990","journal-title":"Am J Physiol"},{"key":"2023081009313744500_CIT0041","doi-asserted-by":"crossref","first-page":"450","DOI":"10.1037\/0735-7044.111.2.450","article-title":"Chorda tympani transection and selective desalivation differentially disrupt two-lever salt discrimination performance in rats","volume":"111","author":"St John","year":"1997","journal-title":"Behav Neurosci"},{"key":"2023081009313744500_CIT0042","first-page":"R141","article-title":"Salt discriminability is related to number of regenerated taste buds after chorda tympani nerve section in rats","volume":"269","author":"St John","year":"1995","journal-title":"Am J Physiol"},{"key":"2023081009313744500_CIT0043","doi-asserted-by":"crossref","first-page":"647","DOI":"10.1093\/chemse\/bjx043","article-title":"Sodium Carbonate is Saltier Than Sodium Chloride to Sodium-Depleted Rats","volume":"42","author":"St John","year":"2017","journal-title":"Chem Senses"},{"key":"2023081009313744500_CIT0044","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1016\/0031-9384(95)00056-O","article-title":"Taste quality profiles for fifteen organic and inorganic salts","volume":"58","author":"van der Klaauw","year":"1995","journal-title":"Physiol Behav"},{"key":"2023081009313744500_CIT0045","doi-asserted-by":"crossref","first-page":"117","DOI":"10.1093\/chemse\/bjx076","article-title":"Aging decreases chorda-tympani nerve responses to NaCl and alters morphology of fungiform taste pores in rats","volume":"43","author":"Whiddon","year":"2018","journal-title":"Chem Senses"},{"issue":"5","key":"2023081009313744500_CIT0046","doi-asserted-by":"crossref","first-page":"e65439","DOI":"10.1371\/journal.pone.0065439","article-title":"Statistical analysis and decoding of neural activity in the rodent geniculate ganglion using a metric-based inference system","volume":"8","author":"Wu","year":"2013","journal-title":"PLoS One"},{"key":"2023081009313744500_CIT0047","doi-asserted-by":"crossref","first-page":"724","DOI":"10.1126\/science.1948054","article-title":"The anion paradox in sodium taste reception: resolution by voltage- clamp studies","volume":"254","author":"Ye","year":"1991","journal-title":"Science"},{"key":"2023081009313744500_CIT0048","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1152\/jn.1993.70.1.167","article-title":"Voltage dependence of the rat chorda tympani response to Na+ salts: implications for the functional organization of taste receptor cells","volume":"70","author":"Ye","year":"1993","journal-title":"J Neurophysiol"},{"key":"2023081009313744500_CIT0049","doi-asserted-by":"crossref","first-page":"927","DOI":"10.1038\/nn.4575","article-title":"The cellular mechanism for water detection in the mammalian taste system","volume":"20","author":"Zocchi","year":"2017","journal-title":"Nat Neurosci"}],"container-title":["Chemical Senses"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/chemse\/advance-article-pdf\/doi\/10.1093\/chemse\/bjad015\/50433841\/bjad015.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/chemse\/article-pdf\/doi\/10.1093\/chemse\/bjad015\/51079281\/bjad015.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/chemse\/article-pdf\/doi\/10.1093\/chemse\/bjad015\/51079281\/bjad015.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,10,21]],"date-time":"2024-10-21T01:26:01Z","timestamp":1729473961000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/chemse\/article\/doi\/10.1093\/chemse\/bjad015\/7177645"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,1,1]]},"references-count":49,"URL":"https:\/\/doi.org\/10.1093\/chemse\/bjad015","relation":{},"ISSN":["0379-864X","1464-3553"],"issn-type":[{"value":"0379-864X","type":"print"},{"value":"1464-3553","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2023,1,1]]},"published":{"date-parts":[[2023,1,1]]},"article-number":"bjad015"}}