{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,20]],"date-time":"2026-10-20T11:43:45Z","timestamp":1792496625257,"version":"4.5.0"},"reference-count":52,"publisher":"American Chemical Society (ACS)","issue":"20","license":[{"start":{"date-parts":[[2026,8,13]],"date-time":"2026-08-13T00:00:00Z","timestamp":1786579200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-029"},{"start":{"date-parts":[[2026,8,13]],"date-time":"2026-08-13T00:00:00Z","timestamp":1786579200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-037"},{"start":{"date-parts":[[2026,8,13]],"date-time":"2026-08-13T00:00:00Z","timestamp":1786579200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-045"}],"funder":[{"DOI":"10.13039\/501100001700","name":"Ministry of Education, Culture, Sports, Science and Technology","doi-asserted-by":"publisher","award":["26H01084"],"award-info":[{"award-number":["26H01084"]}],"id":[{"id":"10.13039\/501100001700","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["26H01084"],"award-info":[{"award-number":["26H01084"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100025019","name":"Support for Pioneering Research Initiated by the Next Generation","doi-asserted-by":"publisher","award":["JPMJSP2106"],"award-info":[{"award-number":["JPMJSP2106"]}],"id":[{"id":"10.13039\/501100025019","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100025019","name":"Support for Pioneering Research Initiated by the Next Generation","doi-asserted-by":"publisher","award":["JPMJSP2180"],"award-info":[{"award-number":["JPMJSP2180"]}],"id":[{"id":"10.13039\/501100025019","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Kunwakai Research Foundation"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,10,20]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Here, we performed biochemical and genetic characterization of a GppA\/PPX-family phosphohydrolase from Arabidopsis thaliana (AtGPPA\/PPX). Sixteen splice variants of AtGPPA\/PPX are deposited in public databases and can be classified into two types based on differences in their N-terminal sequences. Green fluorescent protein fusions with either N-terminus indicated that neither possesses a transit peptide, suggesting that all AtGPPA\/PPX splice variants localize to the cytosol. Biochemical analysis of the long splice variant of AtGPPA\/PPX demonstrated phosphatase activity toward guanosine 5\u2032-triphosphate 3\u2032-diphosphate (pppGpp) and long-chain polyphosphate in vitro, although its substrate affinity and catalytic efficiency were lower than those ofEscherichia coli PPX. Heterologous expression of AtGPPA\/PPX in anEscherichia coligppA mutant reduced intracellular pppGpp levels relative to the mutant control, indicating that AtGPPA\/PPX possesses pppGpp dephosphorylation activity in vivo. Loss-of-function mutants of AtGPPA\/PPX exhibited no overt growth defects under either standard or nitrogen-starvation conditions. However, chlorophyll fluorescence imaging revealed delayed and attenuated nonphotochemical quenching after actinic-light illumination without changes in the PSII quantum yield, suggesting faster electron transport and reduced need for thermal dissipation. Together, these results suggest that the cytosol-localized AtGPPA\/PPX functions as a phosphatase with unknown physiological substrate(s) and plays a key role in photosynthetic acclimation to fluctuating light conditions.<\/jats:p>","DOI":"10.1021\/acs.biochem.6c00467","type":"journal-article","created":{"date-parts":[[2026,8,13]],"date-time":"2026-08-13T16:02:31Z","timestamp":1786636951000},"page":"3273-3285","source":"Crossref","is-referenced-by-count":0,"title":["A Cytosolic\n                    <i>Arabidopsis<\/i>\n                    Homologue\nof Bacterial GppA\/PPX Retains Polyphosphate and pppGpp Phosphatase\nActivities and Regulates Photosynthetic Acclimation"],"prefix":"10.1021","volume":"65","author":[{"given":"Takanari","family":"Nemoto","sequence":"first","affiliation":[{"name":"Tokyo Institute of Technology , , ,","place":["Yokohama, Japan, 226-8501"]},{"name":"Institute of Science Tokyo , , ,","place":["Yokohama, Japan, 226-8501"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuto","family":"Omata","sequence":"additional","affiliation":[{"name":"Tokyo Institute of Technology , , ,","place":["Yokohama, Japan, 226-8501"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Masataka","family":"Inazu","sequence":"additional","affiliation":[{"name":"Tokyo Institute of Technology , , ,","place":["Yokohama, Japan, 226-8501"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shouta","family":"Nonoyama","sequence":"additional","affiliation":[{"name":"Tokyo Institute of Technology , , ,","place":["Yokohama, Japan, 226-8501"]},{"name":"Institute of Science Tokyo , , ,","place":["Yokohama, Japan, 226-8501"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5349-4927","authenticated-orcid":true,"given":"Shinji","family":"Masuda","sequence":"additional","affiliation":[{"name":"Tokyo Institute of Technology , , ,","place":["Yokohama, Japan, 226-8501"]},{"name":"Institute of Science Tokyo , , ,","place":["Yokohama, Japan, 226-8501"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,8,13]]},"reference":[{"key":"2026102007163868300_cit1","first-page":"1458","article-title":"The stringent\nresponse","volume-title":"Escherichia coli and Salmonella:\nCellular and Molecular Biology","author":"Cashel","year":"1996","edition":"2nd"},{"issue":"5","key":"2026102007163868300_cit2","doi-asserted-by":"publisher","first-page":"298","DOI":"10.1038\/nrmicro3448","article-title":"Recent functional\ninsights into the role of (p)ppGpp in bacterial physiology","volume":"13","author":"Hauryliuk","year":"2015","journal-title":"Nat. Rev. Microbiol"},{"key":"2026102007163868300_cit3","doi-asserted-by":"publisher","first-page":"5980","DOI":"10.1016\/S0021-9258(19)67694-5","article-title":"Residual guanosine\n3\u2018,5\u2018-bispyrophosphate\nsynthetic activity of relA null mutants can be eliminated by spoT\nnull mutations","volume":"266","author":"Xiao","year":"1991","journal-title":"J. Biol. Chem"},{"key":"2026102007163868300_cit4","doi-asserted-by":"publisher","first-page":"35","DOI":"10.1146\/annurev.micro.62.081307.162903","article-title":"p)ppGpp: Still magical?","volume":"62","author":"Potrykus","year":"2008","journal-title":"Annu.\nRev. Microbiol"},{"key":"2026102007163868300_cit5","doi-asserted-by":"publisher","first-page":"315","DOI":"10.1007\/BF00382277","article-title":"Mutants of Escherichia coli defective\nin the degradation of guanosine 5\u2032-triphosphate, 3\u2032-diphosphate\n(pppGpp)","volume":"169","author":"Somerville","year":"1979","journal-title":"Mol. Gen. Genet"},{"key":"2026102007163868300_cit6","doi-asserted-by":"publisher","first-page":"21240","DOI":"10.1074\/jbc.272.34.21240","article-title":"Guanosine tetra- and\npentaphosphate promote accumulation of inorganic polyphosphate in Escherichia coli","volume":"272","author":"Kuroda","year":"1997","journal-title":"J. Biol.\nChem"},{"key":"2026102007163868300_cit7","doi-asserted-by":"publisher","first-page":"6175","DOI":"10.1093\/nar\/gkt302","article-title":"Differential\nregulation by ppGpp versus pppGpp in Escherichia coli","volume":"41","author":"Mechold","year":"2013","journal-title":"Nucleic Acids Res"},{"key":"2026102007163868300_cit8","doi-asserted-by":"publisher","first-page":"93","DOI":"10.1111\/mmi.12172","article-title":"Dose-dependent reduction of replication\nelongation rate by (p)ppGpp in Escherichia coli and Bacillus subtilis","volume":"88","author":"Denapoli","year":"2013","journal-title":"Mol.\nMicrobiol"},{"key":"2026102007163868300_cit9","doi-asserted-by":"publisher","first-page":"1469","DOI":"10.1016\/j.jmb.2007.11.073","article-title":"Structure of the PPX\/GPPA phosphatase\nfrom Aquifex aeolicus in complex with the alarmone\nppGpp","volume":"375","author":"Kristensen","year":"2008","journal-title":"J. Mol. Biol"},{"key":"2026102007163868300_cit10","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0042561","article-title":"The two PPX-GppA\nhomologues from Mycobacterium tuberculosis have distinct biochemical\nactivities","volume":"7","author":"Choi","year":"2012","journal-title":"PLoS One"},{"key":"2026102007163868300_cit11","doi-asserted-by":"publisher","first-page":"564644","DOI":"10.3389\/fmicb.2020.564644","article-title":"Functional\ninsights\ninto the role of gppA in (p)ppGpp metabolism of Vibrio cholerae","volume":"11","author":"Rakshit","year":"2020","journal-title":"Front. 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Sci"},{"key":"2026102007163868300_cit26","doi-asserted-by":"publisher","first-page":"156","DOI":"10.1016\/0968-0004(94)90275-5","article-title":"Yeast protein\ncontrolling inter-organelle communication is related to bacterial\nphosphatases containing the Hsp70-type ATP-binding domain","volume":"19","author":"Koonin","year":"1994","journal-title":"Trends Biochem. Sci"},{"key":"2026102007163868300_cit27","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0023479","article-title":"The RelA\/SpoT\nHomolog (RSH) superfamily:\nDistribution and Ffunctional evolution of ppGpp synthetases and hydrolases\nacross the tree of life","volume":"6","author":"Atkinson","year":"2011","journal-title":"PLoS One"},{"key":"2026102007163868300_cit28","doi-asserted-by":"publisher","first-page":"625","DOI":"10.1007\/s10265-017-0922-8","article-title":"Phylogenetic analysis\nof proteins involved in the stringent response in plant cells","volume":"130","author":"Ito","year":"2017","journal-title":"J. Plant Res"},{"key":"2026102007163868300_cit29","doi-asserted-by":"publisher","first-page":"7495","DOI":"10.1074\/jbc.M311573200","article-title":"Inducible expression, enzymatic activity,\nand origin\nof higher plant homologues of bacterial RelA\/SpoT stress proteins\nin Nicotiana tabacum","volume":"279","author":"Givens","year":"2004","journal-title":"J. Biol. Chem"},{"key":"2026102007163868300_cit30","doi-asserted-by":"publisher","first-page":"35536","DOI":"10.1074\/jbc.M703820200","article-title":"Calcium-activated\n(p)ppGpp synthetase\nin chloroplasts of land plants","volume":"282","author":"Tozawa","year":"2007","journal-title":"J. Biol. Chem"},{"key":"2026102007163868300_cit31","doi-asserted-by":"publisher","DOI":"10.26508\/LSA.201900429","article-title":"Detecting sequence signals in targeting\npeptides using deep learning","volume":"2","author":"Armenteros","year":"2019","journal-title":"Life Sci. 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Natl. Acad. Sci. U. S. A"},{"key":"2026102007163868300_cit43","doi-asserted-by":"publisher","first-page":"511","DOI":"10.1007\/s10265-015-0711-1","article-title":"A highly sensitive\nquantification\nmethod for the accumulation of alarmone ppGpp in Arabidopsis thaliana\nusing UPLC-ESI-qMS\/MS","volume":"128","author":"Ihara","year":"2015","journal-title":"J. Plant Res"},{"key":"2026102007163868300_cit44","doi-asserted-by":"publisher","first-page":"7029","DOI":"10.1073\/pnas.90.15.7029","article-title":"Guanosine\npentaphosphate phosphohydrolase\nof Escherichia coli is a long- chain\nexopolyphosphatase","volume":"90","author":"Keasling","year":"1993","journal-title":"Proc. Natl. Acad. Sci. U.\nS. 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Rep"},{"key":"2026102007163868300_cit50","doi-asserted-by":"publisher","first-page":"507","DOI":"10.1111\/tpj.14642","article-title":"A genetically validated approach for detecting inorganic\npolyphosphates in plants","volume":"102","author":"Zhu","year":"2020","journal-title":"Plant J"},{"key":"2026102007163868300_cit51","doi-asserted-by":"publisher","first-page":"817","DOI":"10.1007\/s00425-017-2780-y","article-title":"Within and beyond the\nstringent response-RSH and (p)ppGpp in plants","volume":"246","author":"Boniecka","year":"2017","journal-title":"Planta"},{"key":"2026102007163868300_cit52","doi-asserted-by":"publisher","first-page":"385","DOI":"10.1093\/plphys\/kiae241","article-title":"Genetically\nmanipulated chloroplast\nstromal phosphate levels alter photosynthetic efficiency","volume":"196","author":"Raju","year":"2024","journal-title":"Plant Physiol"}],"container-title":["Biochemistry"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/pubs.acs.org\/doi\/pdf\/10.1021\/acs.biochem.6c00467","content-type":"application\/pdf","content-version":"vor","intended-application":"unspecified"},{"URL":"https:\/\/pubs.acs.org\/bichaw\/article-pdf\/65\/20\/3273\/66532687\/acs.biochem.6c00467.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/pubs.acs.org\/bichaw\/article-pdf\/65\/20\/3273\/66532687\/acs.biochem.6c00467.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,20]],"date-time":"2026-10-20T11:29:18Z","timestamp":1792495758000},"score":1,"resource":{"primary":{"URL":"https:\/\/pubs.acs.org\/bichaw\/article\/65\/20\/3273\/5254637\/A-Cytosolic-Arabidopsis-Homologue-of-Bacterial"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,13]]},"references-count":52,"journal-issue":{"issue":"20","published-online":{"date-parts":[[2026,8,13]]},"published-print":{"date-parts":[[2026,10,20]]}},"URL":"https:\/\/doi.org\/10.1021\/acs.biochem.6c00467","relation":{},"ISSN":["0006-2960","1520-4995"],"issn-type":[{"value":"0006-2960","type":"print"},{"value":"1520-4995","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,13]]}}}