{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,7]],"date-time":"2026-03-07T03:56:52Z","timestamp":1772855812956,"version":"3.50.1"},"reference-count":28,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2025,12,18]],"date-time":"2025-12-18T00:00:00Z","timestamp":1766016000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"},{"start":{"date-parts":[[2025,12,18]],"date-time":"2025-12-18T00:00:00Z","timestamp":1766016000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"}],"funder":[{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["21K06681"],"award-info":[{"award-number":["21K06681"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["19K07190"],"award-info":[{"award-number":["19K07190"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["19K07110"],"award-info":[{"award-number":["19K07110"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["21K06723"],"award-info":[{"award-number":["21K06723"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Neurochem Res"],"published-print":{"date-parts":[[2026,2]]},"DOI":"10.1007\/s11064-025-04635-0","type":"journal-article","created":{"date-parts":[[2025,12,18]],"date-time":"2025-12-18T09:31:10Z","timestamp":1766050270000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["l-Theanine Uptake via Solute Carrier Family 38 Member 1 Inhibits Neuroblastoma Cell Growth"],"prefix":"10.1007","volume":"51","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-9232-770X","authenticated-orcid":false,"given":"Koichi","family":"Kawada","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yuri","family":"Matsushima","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ruka","family":"Nakamura","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Megumi","family":"Nishio","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yuka","family":"Watanabe","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Shuhei","family":"Araki","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Tamika","family":"Sudo","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Sayuki","family":"Yoshikawa","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Mitsuhiro","family":"Shibata","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Toshihiko","family":"Kinjo","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0315-0456","authenticated-orcid":false,"given":"Kyosuke","family":"Uno","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yutaro","family":"Higashiura","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Akiko","family":"Yamamuro","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6767-7628","authenticated-orcid":false,"given":"Yuki","family":"Ishimaru","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6827-9701","authenticated-orcid":false,"given":"Yasuhiro","family":"Yoshioka","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Sadaaki","family":"Maeda","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2565-2067","authenticated-orcid":false,"given":"Nobuyuki","family":"Kuramoto","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"297","published-online":{"date-parts":[[2025,12,18]]},"reference":[{"key":"4635_CR1","doi-asserted-by":"publisher","first-page":"1583","DOI":"10.1007\/s00726-014-1713-3","volume":"46","author":"M Narukawa","year":"2014","unstructured":"Narukawa M, Toda Y, Nakagita T, Hayashi Y, Misaka T (2014) L-theanine elicits umami taste via the T1R1\u2009+\u2009T1R3 umami taste receptor. Amino Acids 46:1583\u20131587. https:\/\/doi.org\/10.1007\/s00726-014-1713-3","journal-title":"Amino Acids"},{"key":"4635_CR2","doi-asserted-by":"publisher","first-page":"667","DOI":"10.1023\/a:1022490806093","volume":"23","author":"H Yokogoshi","year":"1998","unstructured":"Yokogoshi H, Kobayashi M, Mochizuki M, Terashima T (1998) Effect of theanine, \u03b3-glutamylethylamide, on brain monoamines and striatal dopamine release in conscious rats. Neurochem Res 23:667\u2013673. https:\/\/doi.org\/10.1023\/a:1022490806093","journal-title":"Neurochem Res"},{"key":"4635_CR3","doi-asserted-by":"publisher","first-page":"1846","DOI":"10.1002\/jnr.21637","volume":"86","author":"T Kakuda","year":"2008","unstructured":"Kakuda T, Hinoi E, Abe A, Nozawa A, Ogura M, Yoneda Y (2008) Theanine, an ingredient of green tea, inhibits [3H]glutamine transport in neurons and astroglia in rat brain. J Neurosci Res 86:1846\u20131856. https:\/\/doi.org\/10.1002\/jnr.21637","journal-title":"J Neurosci Res"},{"key":"4635_CR4","doi-asserted-by":"publisher","first-page":"436","DOI":"10.1080\/07315724.2014.926153","volume":"34","author":"TP Rao","year":"2015","unstructured":"Rao TP, Ozeki M, Juneja LR (2015) In search of a safe natural sleep aid. J Am Coll Nutr 34:436\u2013447. https:\/\/doi.org\/10.1080\/07315724.2014.926153","journal-title":"J Am Coll Nutr"},{"key":"4635_CR5","doi-asserted-by":"publisher","first-page":"287","DOI":"10.1271\/bbb.64.287","volume":"64","author":"T Kakuda","year":"2000","unstructured":"Kakuda T, Nozawa A, Unno T, Okamura N, Okai O (2000) Inhibiting effects of theanine on caffeine stimulation evaluated by EEG in the rat. Biosci Biotechnol Biochem 64:287\u2013293. https:\/\/doi.org\/10.1271\/bbb.64.287","journal-title":"Biosci Biotechnol Biochem"},{"key":"4635_CR6","doi-asserted-by":"publisher","first-page":"292","DOI":"10.1016\/j.jphs.2014.12.018","volume":"127","author":"T Takarada","year":"2015","unstructured":"Takarada T, Nakamichi N, Kakuda T, Nakazato R, Kokubo H, Ikeno S et al (2015) Daily oral intake of theanine prevents the decline of 5-bromo-2\u2019-deoxyuridine incorporation in hippocampal dentate gyrus with concomitant alleviation of behavioral abnormalities in adult mice with severe traumatic stress. J Pharmacol Sci 127:292\u2013297. https:\/\/doi.org\/10.1016\/j.jphs.2014.12.018","journal-title":"J Pharmacol Sci"},{"key":"4635_CR7","doi-asserted-by":"publisher","first-page":"7110","DOI":"10.1002\/ptr.7110","volume":"35","author":"S Shojaei-Zarghani","year":"2021","unstructured":"Shojaei-Zarghani S, Rafraf M, Yari-Khosroushahi A (2021) Theanine and cancer: a systematic review of the literature. Phytother Res 35:7110. https:\/\/doi.org\/10.1002\/ptr.7110","journal-title":"Phytother Res"},{"key":"4635_CR8","doi-asserted-by":"publisher","DOI":"10.1155\/2017\/9747256","volume":"2017","author":"Q Yan","year":"2017","unstructured":"Yan Q, Tong H, Tang S, Tan Z, Han X, Zhou C (2017) L-theanine administration modulates the absorption of dietary nutrients and expression of transporters and receptors in the intestinal mucosa of rats. Biomed Res Int 2017:9747256. https:\/\/doi.org\/10.1155\/2017\/9747256","journal-title":"Biomed Res Int"},{"key":"4635_CR9","doi-asserted-by":"publisher","first-page":"1768","DOI":"10.1271\/bbb.60.1768","volume":"60","author":"S Kitaoka","year":"1996","unstructured":"Kitaoka S, Hayashi H, Yokogoshi H, Suzuki Y (1996) Transmural potential changes associated with the in vitro absorption of theanine in the Guinea pig intestine. Biosci Biotechnol Biochem 60:1768\u20131771. https:\/\/doi.org\/10.1271\/bbb.60.1768","journal-title":"Biosci Biotechnol Biochem"},{"key":"4635_CR10","doi-asserted-by":"publisher","first-page":"89","DOI":"10.1016\/j.bbrep.2015.09.021","volume":"5","author":"T Takarada","year":"2015","unstructured":"Takarada T, Nakamichi N, Nakazato R, Kakuda T, Kokubo H, Ikeno S et al (2015) Possible activation by the green tea amino acid theanine of mammalian target of Rapamycin signaling in undifferentiated neural progenitor cells in vitro. Biochem Biophys Rep 5:89\u201395. https:\/\/doi.org\/10.1016\/j.bbrep.2015.09.021","journal-title":"Biochem Biophys Rep"},{"key":"4635_CR11","doi-asserted-by":"publisher","first-page":"e48270","DOI":"10.1371\/journal.pone.0048270","volume":"7","author":"M Ogura","year":"2012","unstructured":"Ogura M, Kakuda T, Takarada T, Nakamichi N, Fukumori R, Kim YH et al (2012) Promotion of both proliferation and neuronal differentiation in pluripotent P19 cells with stable overexpression of the glutamine transporter slc38a1. PLoS One 7:e48270. https:\/\/doi.org\/10.1371\/journal.pone.0048270","journal-title":"PLoS One"},{"key":"4635_CR12","doi-asserted-by":"publisher","first-page":"877","DOI":"10.1101\/gad.189365.112","volume":"26","author":"CV Dang","year":"2012","unstructured":"Dang CV (2012) Links between metabolism and cancer. Genes Dev 26:877\u2013890. https:\/\/doi.org\/10.1101\/gad.189365.112","journal-title":"Genes Dev"},{"key":"4635_CR13","doi-asserted-by":"publisher","first-page":"752","DOI":"10.1016\/j.tibs.2018.05.003","volume":"43","author":"P Kandasamy","year":"2018","unstructured":"Kandasamy P, Gyimesi G, Kanai Y, Hediger MA (2018) Amino acid transporters revisited: new views in health and disease. Trends Biochem Sci 43:752\u2013789. https:\/\/doi.org\/10.1016\/j.tibs.2018.05.003","journal-title":"Trends Biochem Sci"},{"key":"4635_CR14","doi-asserted-by":"publisher","first-page":"108","DOI":"10.1016\/j.mam.2013.01.001","volume":"34","author":"Y Kanai","year":"2013","unstructured":"Kanai Y, Cl\u00e9men\u00e7on B, Simonin A, Leuenberger M, Lochner M, Weisstanner M et al (2013) The SLC1 high-affinity glutamate and neutral amino acid transporter family. Mol Aspects Med 34:108\u2013120. https:\/\/doi.org\/10.1016\/j.mam.2013.01.001","journal-title":"Mol Aspects Med"},{"key":"4635_CR15","doi-asserted-by":"publisher","first-page":"2531","DOI":"10.1016\/j.bbamcr.2015.12.017","volume":"1863","author":"YD Bhutia","year":"2016","unstructured":"Bhutia YD, Ganapathy V (2016) Glutamine transporters in mammalian cells and their functions in physiology and cancer. Biochim Biophys Acta 1863:2531\u20132539. https:\/\/doi.org\/10.1016\/j.bbamcr.2015.12.017","journal-title":"Biochim Biophys Acta"},{"key":"4635_CR16","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1126\/science.aab2674","volume":"351","author":"RL Wolfson","year":"2016","unstructured":"Wolfson RL, Chantranupong L, Saxton RA, Shen K, Scaria SM, Cantor JR et al (2016) Sestrin2 is a leucine sensor for the mTORC1 pathway. Science 351:43\u201348. https:\/\/doi.org\/10.1126\/science.aab2674","journal-title":"Science"},{"key":"4635_CR17","volume-title":"Basic Neurochemistry","author":"S Brady","year":"2011","unstructured":"Brady S, Siegel G, Albers RW, Price D (2011) Basic Neurochemistry. Academic Press, Cambridge"},{"key":"4635_CR18","first-page":"1989","volume":"26","author":"SG Kim","year":"2006","unstructured":"Kim SG, Kim HH, Kim HK, Kim CH, Chun HS, Kanai Y et al (2006) Differential expression and functional characterization of system L amino acid transporters in human normal osteoblast cells and osteogenic sarcoma cells. Anticancer Res 26:1989\u20131996","journal-title":"Anticancer Res"},{"key":"4635_CR19","doi-asserted-by":"publisher","first-page":"368","DOI":"10.1254\/jphs.12124fp","volume":"119","author":"N Khunweeraphong","year":"2012","unstructured":"Khunweeraphong N, Nagamori S, Wiriyasermkul P, Nishinaka Y, Wongthai P, Ohgaki R et al (2012) Establishment of stable cell lines with high expression of heterodimers of human 4F2hc and human amino acid transporter LAT1 or LAT2 and delineation of their differential interaction with \u03b1-alkyl moieties. J Pharmacol Sci 119:368\u2013380. https:\/\/doi.org\/10.1254\/jphs.12124fp","journal-title":"J Pharmacol Sci"},{"key":"4635_CR20","doi-asserted-by":"publisher","first-page":"19745","DOI":"10.1074\/jbc.274.28.19745","volume":"274","author":"H Segawa","year":"1999","unstructured":"Segawa H, Fukasawa Y, Miyamoto K, Takeda E, Endou H, Kanai Y (1999) Identification and functional characterization of a Na+-independent neutral amino acid transporter with broad substrate selectivity. J Biol Chem 274:19745\u201319751. https:\/\/doi.org\/10.1074\/jbc.274.28.19745","journal-title":"J Biol Chem"},{"key":"4635_CR21","doi-asserted-by":"publisher","unstructured":"Kanai Y (2021) Amino acid transporter LAT1 (SLC7A5) as a molecular target for cancer diagnosis and therapeutics. Pharmacol Ther 107964. https:\/\/doi.org\/10.1016\/j.pharmthera.2021.107964","DOI":"10.1016\/j.pharmthera.2021.107964"},{"key":"4635_CR22","doi-asserted-by":"publisher","first-page":"2230","DOI":"10.1271\/bbb.120519","volume":"76","author":"S Yamamoto","year":"2012","unstructured":"Yamamoto S, Kimura T, Tachiki T, Anzai N, Sakurai T, Ushimaru M (2012) The involvement of L-type amino acid transporters in theanine transport. Biosci Biotechnol Biochem 76:2230\u20132235. https:\/\/doi.org\/10.1271\/bbb.120519","journal-title":"Biosci Biotechnol Biochem"},{"key":"4635_CR23","doi-asserted-by":"publisher","first-page":"571","DOI":"10.1016\/j.mam.2012.07.012","volume":"34","author":"HB Schi\u00f6th","year":"2013","unstructured":"Schi\u00f6th HB, Roshanbin S, H\u00e4gglund MG, Fredriksson R (2013) Evolutionary origin of amino acid transporter families SLC32, SLC36 and SLC38 and physiological, pathological and therapeutic aspects. Mol Aspects Med 34:571\u2013585. https:\/\/doi.org\/10.1016\/j.mam.2012.07.012","journal-title":"Mol Aspects Med"},{"key":"4635_CR24","doi-asserted-by":"publisher","first-page":"92","DOI":"10.1007\/s004240100663","volume":"443","author":"A Albers","year":"2001","unstructured":"Albers A, Br\u00f6er A, Wagner CA, Setiawan I, Lang PA, Kranz EU et al (2001) Na+ transport by the neural glutamine transporter ATA1. Pflugers Arch 443:92\u2013101. https:\/\/doi.org\/10.1007\/s004240100663","journal-title":"Pflugers Arch"},{"key":"4635_CR25","doi-asserted-by":"publisher","DOI":"10.1007\/s00726-024-03389-3","volume":"7","author":"S Yamaura","year":"2024","unstructured":"Yamaura S, Sadamori K, Konishi R, Majima T, Mukai A, Uno K et al (2024) Pharmacokinetics of L-theanine and the effect on amino acid composition in mice administered with L-theanine. Amino Acids 7:29. https:\/\/doi.org\/10.1007\/s00726-024-03389-3","journal-title":"Amino Acids"},{"key":"4635_CR26","doi-asserted-by":"publisher","first-page":"931","DOI":"10.1038\/nrd4002","volume":"12","author":"C Gorrini","year":"2013","unstructured":"Gorrini C, Harris IS, Mak TW (2013) Modulation of oxidative stress as an anticancer strategy. Nat Rev Drug Discov 12:931\u2013947. https:\/\/doi.org\/10.1038\/nrd4002","journal-title":"Nat Rev Drug Discov"},{"key":"4635_CR27","doi-asserted-by":"publisher","first-page":"15074","DOI":"10.1038\/ncomms15074","volume":"8","author":"CS Shin","year":"2017","unstructured":"Shin CS, Mishra P, Watrous JD, Carelli V, D\u2019Aurelio M, Jain M et al (2017) The glutamate\/cystine xCT antiporter antagonizes glutamine metabolism and reduces nutrient flexibility. Nat Commun 8:15074. https:\/\/doi.org\/10.1038\/ncomms15074","journal-title":"Nat Commun"},{"key":"4635_CR28","doi-asserted-by":"publisher","DOI":"10.1016\/j.cmp.2025.06.001","author":"K Kawada","year":"2025","unstructured":"Kawada K, Yamaura S, Yamamoto C, Komori K, Kinjo T, Uno K et al (2025) L-theanine inhibits mammalian target of rapamycin-p70S6K pathway-mediated neuroexcitation induced by N-methyl-D-aspartate. Curr Mol Pharmacol. https:\/\/doi.org\/10.1016\/j.cmp.2025.06.001","journal-title":"Curr Mol Pharmacol"}],"container-title":["Neurochemical Research"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s11064-025-04635-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s11064-025-04635-0","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s11064-025-04635-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,3,4]],"date-time":"2026-03-04T10:11:36Z","timestamp":1772619096000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s11064-025-04635-0"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,12,18]]},"references-count":28,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2026,2]]}},"alternative-id":["4635"],"URL":"https:\/\/doi.org\/10.1007\/s11064-025-04635-0","relation":{},"ISSN":["0364-3190","1573-6903"],"issn-type":[{"value":"0364-3190","type":"print"},{"value":"1573-6903","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,12,18]]},"assertion":[{"value":"1 October 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"1 December 2025","order":2,"name":"revised","label":"Revised","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"9 December 2025","order":3,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"18 December 2025","order":4,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare no competing interests.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing Interests"}},{"value":"This study did not involve human participants or live animals.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethical Approval"}}],"article-number":"17"}}