{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,1]],"date-time":"2026-11-01T08:38:02Z","timestamp":1793522282365,"version":"4.5.0"},"reference-count":39,"publisher":"Springer Science and Business Media LLC","license":[{"start":{"date-parts":[[2026,4,29]],"date-time":"2026-04-29T00:00:00Z","timestamp":1777420800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,4,29]],"date-time":"2026-04-29T00:00:00Z","timestamp":1777420800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100012330","name":"Okayama University","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100012330","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["J Plant Res"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>\n                    Stomatal movements are essential for balancing photosynthetic carbon dioxide uptake with water conservation and defense against pathogens. These processes are controlled by complex signaling networks in guard cells, in which calcium ions (Ca\n                    <jats:sup>2+<\/jats:sup>\n                    ) act as a ubiquitous second messenger. Although stimulus-specific Ca\n                    <jats:sup>2+<\/jats:sup>\n                    signatures have been well documented, how these signals are decoded into distinct physiological responses remains a central question in plant biology. Increasing evidence highlights calcium-dependent protein kinases (CDPKs) as key signal decoders in guard cell signaling. This mini-review summarizes recent advances in our understanding of how CDPKs perceive and translate Ca\n                    <jats:sup>2+<\/jats:sup>\n                    fluctuations into stomatal responses. We focus on the roles of CDPKs in signaling pathways triggered by diverse stimuli, including phytohormones such as abscisic acid ABA, jasmonates, and salicylic acid, as well as biotic cues such as microbe- or pathogen-associated molecular patterns (MAMPs\/PAMPs) and pathogen infection. We also discuss how gaseous signals and metabolic cues are integrated into CDPK-mediated pathways. In addition to their established role as downstream decoders of Ca\n                    <jats:sup>2+<\/jats:sup>\n                    signals, emerging studies suggest that CDPKs can act upstream of Ca\n                    <jats:sup>2+<\/jats:sup>\n                    oscillations and may also function through Ca\n                    <jats:sup>2+<\/jats:sup>\n                    -independent mechanisms. Together, these findings highlight the context-dependent and integrative roles of CDPKs in regulating stomatal behavior, contributing to plant fitness under fluctuating environmental conditions.\n                  <\/jats:p>","DOI":"10.1007\/s10265-026-01703-6","type":"journal-article","created":{"date-parts":[[2026,4,29]],"date-time":"2026-04-29T03:29:52Z","timestamp":1777433392000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":4,"title":["CDPKs as Ca2+ signaling decoders in guard cell signaling"],"prefix":"10.1007","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-2235-0779","authenticated-orcid":false,"given":"Izumi C.","family":"Mori","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,4,29]]},"reference":[{"key":"1703_CR1","doi-asserted-by":"publisher","first-page":"3372","DOI":"10.1093\/jxb\/erac105","volume":"73","author":"C Allan","year":"2022","unstructured":"Allan C, Morris RJ, Meisrimler CN (2022) Encoding, transmission, decoding, and specificity of calcium signals in plants. J Exp Bot 73:3372\u20133385","journal-title":"J Exp Bot"},{"key":"1703_CR2","doi-asserted-by":"publisher","first-page":"2338","DOI":"10.1126\/science.289.5488.2338","volume":"289","author":"GJ Allen","year":"2000","unstructured":"Allen GJ, Chu SP, Harrington CL, Schumacher K, Shimazaki CT, Vafeados D, Kemper A, Hawke SD, Tallman G, Tsien RY, Harper JF, Chory J, Schroeder JI (2000) Alteration of stimulus-specific guard cell calcium oscillations and stomatal closing in Arabidopsis det3 mutant. Science 289:2338\u20132342","journal-title":"Science"},{"key":"1703_CR3","doi-asserted-by":"publisher","first-page":"1053","DOI":"10.1038\/35082575","volume":"411","author":"GJ Allen","year":"2001","unstructured":"Allen GJ, Chu SP, Harrington CL, Schumacher K, Hoffmann T, Tang YY, Grill E, Schroeder JI (2001) A defined range of guard cell calcium oscillation parameters encodes stomatal movements. Nature 411:1053\u20131057","journal-title":"Nature"},{"key":"1703_CR5","first-page":"952","volume":"53","author":"B Brandt","year":"2012","unstructured":"Brandt B, Brodsky DE, Xue S, Negi J, Iba K, Kangasj\u00e4rvi J, Ghassemian M, Stephan AB, Hu H, Schroeder JI (2012) Reconstitution of abscisic acid activation of SLAC1 anion channel by CPK6\/21\/23 and OST1 kinases and PP2CA phosphatase in Xenopus oocytes. Plant Cell Physiol 53:952\u2013961","journal-title":"Plant Cell Physiol"},{"key":"1703_CR6","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1042\/BJ20091147","volume":"425","author":"TA De Falco","year":"2010","unstructured":"De Falco TA, Moeder W, Yoshioka K (2010) Breaking the code: Ca2+ sensors in plant signalling. Biochem J 425:27\u201340","journal-title":"Biochem J"},{"key":"1703_CR7","doi-asserted-by":"publisher","first-page":"162","DOI":"10.1111\/nph.13435","volume":"208","author":"AG Deger","year":"2015","unstructured":"Deger AG, Scherzer S, Nuhkat M, Kedzierska J, Kollist H, Brosch\u00e9 M, Unyayar S, Boudsocq M, Hedrich R, Roelfsema MRG (2015) Guard cell SLAC1-type anion channels mediate flagellin-induced stomatal closure. New Phytol 208:162\u2013173","journal-title":"New Phytol"},{"key":"1703_CR8","first-page":"2188","volume":"22","author":"D Geiger","year":"2010","unstructured":"Geiger D, Scherzer S, Mumm P, Stange A, Marten I, Bauer H, Ache P, Matschi S, Liese A, Al-Rasheid KAS, Romeis T, Hedrich R (2010) Guard cell anion channel SLAC1 is regulated by CDPK protein kinases with distinct Ca2+ affinities. Plant Cell 22:2188\u20132199","journal-title":"Plant Cell"},{"key":"1703_CR9","doi-asserted-by":"publisher","DOI":"10.3390\/ijms22126304","volume":"22","author":"MM Hasan","year":"2021","unstructured":"Hasan MM, Rahman MA, Skalicky M, Alabdallah NM, Waseem M, Jahan MS, Ahammed GJ, El-Mogy MM, Abou El-Yazied A, Ibrahim MFM, Fang XW (2021) Ozone induced stomatal regulations, MAPK and phytohormone signaling in plants. Int J Mol Sci 22:6304","journal-title":"Int J Mol Sci"},{"key":"1703_CR10","doi-asserted-by":"publisher","first-page":"2054","DOI":"10.1016\/j.biochi.2011.05.019","volume":"93","author":"K Hashimoto","year":"2011","unstructured":"Hashimoto K, Kudla J (2011) Calcium decoding mechanisms in plants. Biochimie 93:2054\u20132059","journal-title":"Biochimie"},{"key":"1703_CR11","first-page":"963","volume":"156","author":"K Held","year":"2011","unstructured":"Held K, Pascaud F, Eckert C, Gajdanowicz P, Hashimoto K, Corratg\u00e9-Faillie C, Offenborn JN, Lacombe B, Dreyer I, Thibaud J-B (2011) Calcium-dependent modulation and plasma membrane targeting of the Arabidopsis potassium channel KAT1 by the protein kinase CPK13. Plant Physiol 156:963\u2013975","journal-title":"Plant Physiol"},{"key":"1703_CR12","doi-asserted-by":"publisher","first-page":"1313","DOI":"10.1111\/jipb.13675","volume":"66","author":"W Hu","year":"2024","unstructured":"Hu W, Cheng J, Lu M, Fang T, Zhu Y, Li Z, Wang X, Wang Y, Guo Y, Yang S, Gong Z (2024) Ca2+-independent ZmCPKs is inhibited by Ca2+-dependent ZmCPK17 during drought response in maize. J Integr Plant Biol 66:1313\u20131333","journal-title":"J Integr Plant Biol"},{"key":"1703_CR13","doi-asserted-by":"publisher","first-page":"177","DOI":"10.1111\/nph.15985","volume":"224","author":"S Huang","year":"2019","unstructured":"Huang S, Waadt R, Nuhkat M, Kollist H, Hedrich R, Roelfsema MRG (2019) Calcium signals in guard cells enhance the efficiency by which abscisic acid triggers stomatal closure. New Phytol 224:177\u2013187","journal-title":"New Phytol"},{"key":"1703_CR14","doi-asserted-by":"crossref","first-page":"6304","DOI":"10.3390\/ijms22126304","volume":"22","author":"C Ko","year":"2021","unstructured":"Ko C, Kim TH (2021) Calcium signaling in guard cells: An update on Ca2+ sensors, ion channels, and signal crosstalk. Int J Mol Sci 22:6304","journal-title":"Int J Mol Sci"},{"key":"1703_CR15","doi-asserted-by":"publisher","first-page":"1065","DOI":"10.1105\/tpc.106.048884","volume":"19","author":"M Kobayashi","year":"2007","unstructured":"Kobayashi M, Ohura I, Kawakita K, Yokota N, Fujiwara M, Shimamoto K, Doke N, Yoshioka H (2007) Calcium-dependent protein kinases regulate the production of reactive oxygen species by potato NADPH oxidase. Plant Cell 19:1065\u20131080","journal-title":"Plant Cell"},{"key":"1703_CR16","doi-asserted-by":"publisher","first-page":"1471","DOI":"10.1093\/jxb\/ery024","volume":"69","author":"Y Li","year":"2018","unstructured":"Li Y, Xu S, Wang Z, He L, Xu K, Wang G (2018) Glucose triggers stomatal closure mediated by basal signaling through HXK1 and PYR\/RCAR receptors in Arabidopsis. J Exp Bot 69:1471\u20131484","journal-title":"J Exp Bot"},{"key":"1703_CR17","doi-asserted-by":"publisher","DOI":"10.1111\/ppl.70196","volume":"177","author":"X Li","year":"2025","unstructured":"Li X, Wang L, Zhang M, He J, An Y (2025) Role of CPKs in ethylene-induced Arabidopsis stomatal closure and their crosstalk with H2O2 and NO signaling. Physiol Plant 177:e70196","journal-title":"Physiol Plant"},{"key":"1703_CR18","doi-asserted-by":"publisher","first-page":"702","DOI":"10.1104\/pp.104.042069","volume":"135","author":"IC Mori","year":"2004","unstructured":"Mori IC, Schroeder JI (2004) Reactive oxygen species activation of plant Ca2+ channels. A signalling mechanism in polar growth, hormone transduction, stress signalling, and hypothetically mechanotransduction. Plant Physiol 135:702\u2013708","journal-title":"Plant Physiol"},{"key":"1703_CR19","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pbio.0040327","volume":"4","author":"IC Mori","year":"2006","unstructured":"Mori IC, Murata Y, Yang Y, Munemasa S, Wang Y-F, Andreoli S, Tiriac H, Alonso JM, Harper JF, Ecker JR, Kwak JM, Schroeder JI (2006) CDPKs CPK6 and CPK3 function in ABA regulation of guard-cell s-type anion- and Ca2+-permeable channels and stomatal closure. PLoS Biol 4:e327","journal-title":"PLoS Biol"},{"key":"1703_CR20","doi-asserted-by":"publisher","first-page":"553","DOI":"10.1104\/pp.110.162750","volume":"155","author":"S Munemasa","year":"2011","unstructured":"Munemasa S, Hossain MA, Nakamura Y, Mori IC, Murata Y (2011) The Arabidopsis calcium-dependent protein kinase, CPK6, functions as a positive regulator of methyl jasmonate signaling in guard cells. Plant Physiol 155:553\u2013561","journal-title":"Plant Physiol"},{"key":"1703_CR21","doi-asserted-by":"publisher","first-page":"437","DOI":"10.1111\/pce.13406","volume":"42","author":"L Ooi","year":"2019","unstructured":"Ooi L, Matsuura T, Munemasa S, Murata Y, Katsuhara M, Hirayama T, Mori IC (2019) The mechanism of SO2-induced stomatal closure differs from O3 and CO2 responses and is mediated by nonapoptotic cell death in guard cells. Plant Cell Environ 42:437\u2013447","journal-title":"Plant Cell Environ"},{"key":"1703_CR22","first-page":"1076","volume":"66","author":"L Ooi","year":"2025","unstructured":"Ooi L, Matuura T, Mori IC (2025) Sulfur Dioxide-Induced Guard Cell Death aNd Stomatal Closure are Attenuated in Nitrate\/proton Antipoter AtCLCa Mutants 66:1076\u20131085","journal-title":"Sulfur Dioxide-Induced Guard Cell Death aNd Stomatal Closure are Attenuated in Nitrate\/proton Antipoter AtCLCa Mutants"},{"key":"1703_CR23","doi-asserted-by":"publisher","DOI":"10.1016\/j.jplph.2021.153585","volume":"268","author":"X Ou","year":"2022","unstructured":"Ou X, Li T, Zhao Y, Chang Y, Wu L, Chen G, Day B, Jiang K (2022) Calcium-dependent ABA signaling functions in stomatal immunity by regulating rapid SA responses in guard cells. J Plant Physiol 268:153585","journal-title":"J Plant Physiol"},{"key":"1703_CR24","doi-asserted-by":"publisher","first-page":"441","DOI":"10.1104\/pp.18.00321","volume":"178","author":"MY Prodhan","year":"2018","unstructured":"Prodhan MY, Munemasa S, Nahar MNEN, Nakamura Y, Murata Y (2018) Guard cell salicylic acid signaling is integrated into abscisic acid signaling via Ca2+\/CPK-dependent pathway. Plant Physiol 178:441\u2013450","journal-title":"Plant Physiol"},{"key":"1703_CR26","first-page":"1880","volume":"232","author":"C Schulze","year":"2021","unstructured":"Schulze C, Takahashi Y, Skirycz A, Cieslak M, Koziolek C, K\u00fcgler S, Mayer KFX, Fricke W, K\u00f6ster P, Hedrich R (2021) Stomatal cyclic nucleotide-gated channel 2 mediates CO\u2082 signal transduction in guard cells. New Phytol 232:1880\u20131897","journal-title":"New Phytol"},{"key":"1703_CR27","doi-asserted-by":"publisher","first-page":"13577","DOI":"10.1073\/pnas.96.23.13577","volume":"96","author":"G Torsethauger","year":"1999","unstructured":"Torsethauger G, Pell EJ, Assmann SM (1999) Ozone inhibits guard cell K+ channels implicated in stomatal opening. Proc Natl Acad Sci U S A 96:13577\u201313582","journal-title":"Proc Natl Acad Sci U S A"},{"key":"1703_CR28","doi-asserted-by":"publisher","first-page":"240","DOI":"10.1111\/pce.12273","volume":"38","author":"JP Vainonen","year":"2015","unstructured":"Vainonen JP, Kangasj\u00e4rvi J (2015) Plant signalling in acute ozone exposure. Plant Cell Environ 38:240\u2013252","journal-title":"Plant Cell Environ"},{"key":"1703_CR29","doi-asserted-by":"publisher","first-page":"690","DOI":"10.1111\/nph.12087","volume":"197","author":"HJ Whalley","year":"2013","unstructured":"Whalley HJ, Knight MR (2013) Calcium signatures are decoded by plants to give specific gene responses. New Phytol 197:690\u2013693","journal-title":"New Phytol"},{"key":"1703_CR30","doi-asserted-by":"publisher","first-page":"577","DOI":"10.1038\/s41586-020-2032-3","volume":"578","author":"F-H Wu","year":"2020","unstructured":"Wu F-H, Chi Y, Jiang Z, Xu Y, Xie L, Huang F, Wan D, Ni J, Yuan F, Wu X, Zhang Y, Chen J, Zhao Y, Yi K, Zhu M, Liang T, Zhou J, Zhang D-P, Pei Z-M (2020) Hydrogen peroxide sensor HPCA1 is an LRR receptor kinase in Arabidopsis. Nature 578:577\u2013581","journal-title":"Nature"},{"key":"1703_CR31","doi-asserted-by":"publisher","first-page":"591","DOI":"10.1104\/pp.113.224055","volume":"163","author":"W Ye","year":"2013","unstructured":"Ye W, Muroyama D, Munemasa S, Nkamura Y, Mori IC, Murata Y (2013) Calcium-dependent protein kinase CPK6 positively functions in induction by yeast elicitor of stomatal closure and inhibition by yeast elicitor of light-induced stomatal opening in Arabidopsis. Plant Physiol 163:591\u2013599","journal-title":"Plant Physiol"},{"key":"1703_CR32","doi-asserted-by":"publisher","first-page":"20932","DOI":"10.1073\/pnas.1922319117","volume":"117","author":"W Ye","year":"2020","unstructured":"Ye W, Munemasa S, Shinya T, Wu W, Ma T, Lu J, Kinoshita T, Kaku H, Shibuya N, Murata Y (2020) Stomatal immunity against fungal invasion comprises not only chitin-induced stomatal closure but also chitosan-induced guard cell death. Proc Natl Acad Sci U S A 117:20932\u201320942","journal-title":"Proc Natl Acad Sci U S A"},{"key":"1703_CR33","doi-asserted-by":"publisher","first-page":"39","DOI":"10.1007\/s10265-015-0757-0","volume":"129","author":"R Yoshida","year":"2016","unstructured":"Yoshida R, Mori IC, Kamizono N, Shichiri Y, Shimatani T, Miyata F, Honda K, Iwai S (2016) Glutamate functions in stomatal closure in Arabidopsis and fava bean. J Plant Res 129:39\u201349","journal-title":"J Plant Res"},{"key":"1703_CR34","doi-asserted-by":"crossref","first-page":"112","DOI":"10.1016\/j.envexpbot.2015.06.006","volume":"121","author":"L Zhang","year":"2016","unstructured":"Zhang L, Wang X, Liu Y, Zhang C, Ma L, Wang H (2016) O3-induced changes in stomatal movement and reactive oxygen species production are modulated by salicylic acid in *Arabidopsis thaliana*. Environ Exp Bot 121:112\u2013120","journal-title":"Environ Exp Bot"},{"key":"1703_CR35","first-page":"948","volume":"156","author":"R Zhao","year":"2011","unstructured":"Zhao R, Sun H-L, Mei C, Wang X-J, Yan L, Liu R, Zhang X-F, Wang X-F, Zhang D-P (2011) The Arabidopsis Ca2+-dependent protein kinase CPK12 negatively regulates abscisic acid signaling in guard cells. Plant Physiol 156:948\u2013960","journal-title":"Plant Physiol"},{"key":"1703_CR36","doi-asserted-by":"publisher","first-page":"3019","DOI":"10.1105\/tpc.107.050666","volume":"19","author":"S-Y Zhu","year":"2007","unstructured":"Zhu S-Y, Yu X-C, Wang X-J, Zhao R, Li Y, Fan R-C, Shang Y, Du S-Y, Wang X-F, Wu F-Q, Xu Y-H, Zhang X-Y, Zhang D-P (2007) Two calcium-dependent protein kinases, CPK4 and CPK11, regulate abscisic acid signal transduction in Arabidopsis. Plant Cell 19:3019\u20133036","journal-title":"Plant Cell"},{"key":"1703_CR37","doi-asserted-by":"publisher","first-page":"1232","DOI":"10.1104\/pp.110.157545","volume":"154","author":"J-J Zou","year":"2010","unstructured":"Zou J-J, Wei F-J, Wang C, Wu J-J, Ratnasekera D, Liu W-X, Wu W-H (2010) Arabidopsis calcium-dependent protein kinase CPK10 functions in abscisic acid- and Ca2+-mediated stomatal regulation in response to drought stress. Plant Physiol 154:1232\u20131243","journal-title":"Plant Physiol"},{"key":"1703_CR38","doi-asserted-by":"publisher","first-page":"1445","DOI":"10.1105\/tpc.15.00144","volume":"27","author":"J-J Zou","year":"2015","unstructured":"Zou J-J, Li X-D, Ratnasekera D, Wang C, Liu W-X, Song L-F, Zhang W-Z, Wu W-H (2015) Arabidopsis calcium-dependent protein kinase 8 and catalase 3 function in abscisic acid-mediated signaling and H\u2082O\u2082 homeostasis in stomatal guard cells under drought stress. Plant Cell 27:1445\u20131460","journal-title":"Plant Cell"},{"key":"1703_CR40","doi-asserted-by":"publisher","unstructured":"McAinsh MR, Evans NH, Montgomery LT, North KA (2002) Calcium signalling in stomatal responses to pollutants. New Phytol 153: 441-447 https:\/\/doi.org\/10.1046\/j.0028-646X.2001.00336.x","DOI":"10.1046\/j.0028-646X.2001.00336.x"},{"key":"1703_CR41","doi-asserted-by":"publisher","unstructured":"Arnaud D, Hwang I (2015) A sophisticated network of signaling pathways regulates stomatal defenses to bacterial pathogens https:\/\/doi.org\/10.1016\/j.molp.2014.10.012","DOI":"10.1016\/j.molp.2014.10.012"},{"key":"1703_CR42","doi-asserted-by":"publisher","unstructured":"Liu J, Whalley HJ, Knight MR (2015) Combining modelling and experimental approaches to explain how calcium signatures are decoded by calmodulin-binding transcription activators (CAMTAs) to produce specific gene expression responses. New Phytol 208: 174-187 https:\/\/doi.org\/10.1111\/nph.13428","DOI":"10.1111\/nph.13428"}],"container-title":["Journal of Plant Research"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s10265-026-01703-6.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s10265-026-01703-6","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s10265-026-01703-6.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,16]],"date-time":"2026-06-16T22:54:16Z","timestamp":1781650456000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s10265-026-01703-6"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,4,29]]},"references-count":39,"alternative-id":["1703"],"URL":"https:\/\/doi.org\/10.1007\/s10265-026-01703-6","relation":{},"ISSN":["0918-9440","1618-0860"],"issn-type":[{"value":"0918-9440","type":"print"},{"value":"1618-0860","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,4,29]]},"assertion":[{"value":"27 October 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"22 February 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"29 April 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The author declares that he has no conflict of financial interest of personal relationships that could have appeared to influence the work reported in this paper.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Conflict of interest"}}]}}