{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,2,12]],"date-time":"2026-02-12T14:13:21Z","timestamp":1770905601357,"version":"3.50.1"},"reference-count":42,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2017,10,2]],"date-time":"2017-10-02T00:00:00Z","timestamp":1506902400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2017,10,2]],"date-time":"2017-10-02T00:00:00Z","timestamp":1506902400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title><jats:p>Stomatal regulation is important for water transpiration from plants. Stomatal opening and closing are controlled by many transporter proteins in guard cells. AtABCG22 is a member of the ATP-binding cassette (ABC) transporters and is a stomatal regulator; however, the function of AtABCG22 has not yet been determined fully, although a mutant phenotype included a significant effect on stomatal status. Here, we further investigated the function of the <jats:italic>AtABCG22<\/jats:italic> gene and its functional relationships with other subfamily genes. Among close family members, we found a functional relationship of stomatal phenotypes with <jats:italic>AtABCG21<\/jats:italic>, which is also expressed specifically in guard cells. Based on an analysis of double mutants, adding the <jats:italic>atabcg21<\/jats:italic> mutation to <jats:italic>atabcg22<\/jats:italic> mutant partially suppressed the open-stomata phenotype of <jats:italic>atabcg22<\/jats:italic>. Multiple-mutant analyses indicated that this suppression was independent of abscisic acid signaling in guard cells. We also found that <jats:italic>atabcg22<\/jats:italic> mutant showed a unique time course-dependent phenotype, being defective in maintenance of stomatal status after initial stomatal opening elicited by light signaling. The function of AtABCG22 and its relationship with AtABCG21 in stomatal regulation are considered.<\/jats:p>","DOI":"10.1038\/s41598-017-12643-6","type":"journal-article","created":{"date-parts":[[2017,9,26]],"date-time":"2017-09-26T10:21:32Z","timestamp":1506421292000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":16,"title":["Functional relationship of AtABCG21 and AtABCG22 in stomatal regulation"],"prefix":"10.1038","volume":"7","author":[{"given":"Takashi","family":"Kuromori","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Eriko","family":"Sugimoto","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Haruka","family":"Ohiraki","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Kazuko","family":"Yamaguchi-Shinozaki","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Kazuo","family":"Shinozaki","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"297","published-online":{"date-parts":[[2017,10,2]]},"reference":[{"key":"12643_CR1","doi-asserted-by":"publisher","first-page":"3089","DOI":"10.1105\/tpc.007906","volume":"14","author":"AC Mustilli","year":"2002","unstructured":"Mustilli, A. C., Merlot, S., Vavasseur, A., Fenzi, F. & Giraudat, J. Arabidopsis OST1 protein kinase mediates the regulation of stomatal aperture by abscisic acid and acts upstream of reactive oxygen species production. Plant Cell \n                           14, 3089\u20133099 (2002).","journal-title":"Plant Cell"},{"key":"12643_CR2","doi-asserted-by":"publisher","first-page":"1473","DOI":"10.1093\/pcp\/pcf188","volume":"43","author":"R Yoshida","year":"2002","unstructured":"Yoshida, R. et al. ABA-activated SnRK2 protein kinase is required for dehydration stress signaling in Arabidopsis. Plant Cell Physiol \n                           43, 1473\u20131483 (2002).","journal-title":"Plant Cell Physiol"},{"key":"12643_CR3","doi-asserted-by":"publisher","first-page":"2325","DOI":"10.1016\/j.febslet.2007.04.008","volume":"581","author":"S Pandey","year":"2007","unstructured":"Pandey, S., Zhang, W. & Assmann, S. M. Roles of ion channels and transporters in guard cell signal transduction. FEBS Lett. \n                           581, 2325\u20132336 (2007).","journal-title":"FEBS Lett."},{"key":"12643_CR4","doi-asserted-by":"publisher","first-page":"561","DOI":"10.1146\/annurev-arplant-042809-112226","volume":"61","author":"TH Kim","year":"2010","unstructured":"Kim, T. H., B\u00f6hmer, M., Hu, H., Nishimura, N. & Schroeder, J. I. Guard cell signal transduction network: advances in understanding abscisic acid, CO2, and Ca2+ signaling. Annu. Rev. Plant Biol. \n                           61, 561\u2013591 (2010).","journal-title":"Annu. Rev. Plant Biol."},{"key":"12643_CR5","doi-asserted-by":"publisher","first-page":"1243","DOI":"10.2307\/2442367","volume":"67","author":"B Galatis","year":"1980","unstructured":"Galatis, B. & Mitrakos, K. The ultrastructural cytology of the differentiating guard cells of vigna sinensis. Amer. J. Bot. \n                           67, 1243\u20131261 (1980).","journal-title":"Amer. J. 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Plant Biol. \n                           58, 347\u2013375 (2007).","journal-title":"Annu. Rev. Plant Biol."},{"key":"12643_CR9","doi-asserted-by":"publisher","DOI":"10.1199\/tab.0153","volume":"9","author":"J Kang","year":"2011","unstructured":"Kang, J. et al. Plant ABC Transporters. Arabidopsis Book \n                           9, e0153, https:\/\/doi.org\/10.1199\/tab.0153 (2011).","journal-title":"Arabidopsis Book"},{"key":"12643_CR10","doi-asserted-by":"publisher","first-page":"3569","DOI":"10.1105\/tpc.114.129049","volume":"26","author":"V Yadav","year":"2014","unstructured":"Yadav, V. et al. ABCG transporters are required for suberin and pollen wall extracellular barriers in Arabidopsis. Plant Cell \n                           26, 3569\u20133588 (2014).","journal-title":"Plant Cell"},{"key":"12643_CR11","doi-asserted-by":"publisher","first-page":"1863","DOI":"10.1007\/s00299-016-2001-3","volume":"35","author":"S Yim","year":"2016","unstructured":"Yim, S. et al. Postmeiotic development of pollen surface layers requires two Arabidopsis ABCG-type transporters. Plant Cell Rep. \n                           35, 1863\u20131873 (2016).","journal-title":"Plant Cell Rep."},{"key":"12643_CR12","doi-asserted-by":"publisher","first-page":"276","DOI":"10.1104\/pp.16.01556","volume":"174","author":"F Fedi","year":"2017","unstructured":"Fedi, F. et al. Awake1, an ABC-Type Transporter, Reveals an Essential Role for Suberin in the Control of Seed Dormancy. Plant Physiol. \n                           174, 276\u2013283 (2017).","journal-title":"Plant Physiol."},{"key":"12643_CR13","doi-asserted-by":"publisher","first-page":"310","DOI":"10.1105\/tpc.113.118935","volume":"26","author":"H Choi","year":"2014","unstructured":"Choi, H. et al. The role of Arabidopsis ABCG9 and ABCG31 ATP binding cassette transporters in pollen fitness and the deposition of steryl glycosides on the pollen coat. Plant Cell \n                           26, 310\u2013324 (2014).","journal-title":"Plant Cell"},{"key":"12643_CR14","doi-asserted-by":"publisher","first-page":"702","DOI":"10.1126\/science.1102331","volume":"306","author":"JA Pighin","year":"2004","unstructured":"Pighin, J. A. et al. Plant cuticular lipid export requires an ABC transporter. Science \n                           306, 702\u2013704 (2004).","journal-title":"Science"},{"key":"12643_CR15","doi-asserted-by":"publisher","first-page":"485","DOI":"10.1111\/j.1365-313X.2007.03252.x","volume":"52","author":"D Bird","year":"2007","unstructured":"Bird, D. et al. Characterization of Arabidopsis ABCG11\/WBC11, an ATP binding cassette (ABC) transporter that is required for cuticular lipid secretion. Plant J. \n                           52, 485\u2013498 (2007).","journal-title":"Plant J."},{"key":"12643_CR16","doi-asserted-by":"publisher","first-page":"1345","DOI":"10.1104\/pp.107.105676","volume":"145","author":"D Panikashvili","year":"2007","unstructured":"Panikashvili, D. et al. The Arabidopsis DESPERADO\/AtWBC11 transporter is required for cutin and wax secretion. Plant Physiol. \n                           145, 1345\u20131360 (2007).","journal-title":"Plant Physiol."},{"key":"12643_CR17","doi-asserted-by":"publisher","first-page":"1524","DOI":"10.1093\/pcp\/pcm139","volume":"48","author":"H Ukitsu","year":"2007","unstructured":"Ukitsu, H. et al. Cytological and biochemical analysis of COF1, an Arabidopsis mutant of an ABC transporter gene. 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Plant Cell \n                           26, 4483\u20134498 (2014).","journal-title":"Plant Cell"},{"key":"12643_CR30","doi-asserted-by":"publisher","first-page":"885","DOI":"10.1111\/j.1365-313X.2011.04641.x","volume":"67","author":"T Kuromori","year":"2011","unstructured":"Kuromori, T., Sugimoto, E. & Shinozaki, K. Arabidopsis mutants of AtABCG22, an ABC transporter gene, increase water transpiration and drought susceptibility. Plant J. \n                           67, 885\u2013894 (2011).","journal-title":"Plant J."},{"key":"12643_CR31","doi-asserted-by":"publisher","first-page":"325","DOI":"10.1046\/j.1365-313x.2001.01096.x","volume":"27","author":"S Iuchi","year":"2001","unstructured":"Iuchi, S. et al. Regulation of drought tolerance by gene manipulation of 9-cis-epoxycarotenoid dioxygenase, a key enzyme in abscisic acid biosynthesis in Arabidopsis. Plant J. \n                           27, 325\u2013333 (2001).","journal-title":"Plant J."},{"key":"12643_CR32","doi-asserted-by":"publisher","first-page":"751","DOI":"10.1073\/pnas.0134257100","volume":"100","author":"N Shitan","year":"2003","unstructured":"Shitan, N. et al. Involvement of CjMDR1, a plant multidrug-resistance-type ATP-binding cassette protein, in alkaloid transport in Coptis japonica. Proc. Natl. Acad. Sci. USA \n                           100, 751\u2013756 (2003).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"12643_CR33","doi-asserted-by":"publisher","first-page":"1217","DOI":"10.1038\/ncb1782","volume":"10","author":"M Lee","year":"2008","unstructured":"Lee, M. et al. The ABC transporter AtABCB14 is a malate importer and modulates stomatal response to CO2. Nat. Cell Biol. \n                           10, 1217\u20131223 (2008).","journal-title":"Nat. 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Acetylated 1,3-diaminopropane antagonizes abscisic acid-mediated stomatal closing in Arabidopsis. Plant J. \n                           79, 322\u2013333 (2014).","journal-title":"Plant J."},{"key":"12643_CR37","doi-asserted-by":"publisher","first-page":"657","DOI":"10.1016\/j.molp.2015.01.014","volume":"8","author":"E Merilo","year":"2015","unstructured":"Merilo, E., Jalakas, P., Kollist, H. & Brosch\u00e9, M. The role of ABA recycling and transporter proteins in rapid stomatal responses to reduced air humidity, elevated CO2, and exogenous ABA. Mol. Plant \n                           8, 657\u2013659 (2015).","journal-title":"Mol. Plant"},{"key":"12643_CR38","doi-asserted-by":"publisher","first-page":"1321","DOI":"10.1016\/j.molp.2015.06.006","volume":"8","author":"E Merilo","year":"2015","unstructured":"Merilo, E. et al. Abscisic acid transport and homeostasis in the context of stomatal regulation. Mol. Plant \n                           8, 1321\u20131333 (2015).","journal-title":"Mol. Plant"},{"key":"12643_CR39","doi-asserted-by":"publisher","first-page":"33","DOI":"10.1146\/annurev.cellbio.15.1.33","volume":"15","author":"WR Briggs","year":"1999","unstructured":"Briggs, W. R. & Huala, E. Blue-light photoreceptors in higher plants. Annu. Rev. Cell Dev. Biol. \n                           15, 33\u201362 (1999).","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"12643_CR40","doi-asserted-by":"publisher","first-page":"219","DOI":"10.1146\/annurev.arplant.57.032905.105434","volume":"58","author":"K Shimazaki","year":"2007","unstructured":"Shimazaki, K., Doi, M., Assmann, S. M. & Kinoshita, T. Light regulation of stomatal movement. Annu. Rev. Plant Biol. \n                           58, 219\u2013247 (2007).","journal-title":"Annu. Rev. 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Plant Cell \n                           20, 1693\u20131707 (2008).","journal-title":"Plant Cell"}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-017-12643-6.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-017-12643-6","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-017-12643-6.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,12,23]],"date-time":"2022-12-23T11:28:01Z","timestamp":1671794881000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-017-12643-6"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2017,10,2]]},"references-count":42,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2017,12]]}},"alternative-id":["12643"],"URL":"https:\/\/doi.org\/10.1038\/s41598-017-12643-6","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2017,10,2]]},"assertion":[{"value":"26 May 2017","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"13 September 2017","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"2 October 2017","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare that they have no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing Interests"}}],"article-number":"12501"}}