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Natl. Acad. Sci. U.S.A."],"published-print":{"date-parts":[[2019,12,10]]},"abstract":"<jats:p>\n                    Iron is an essential nutrient for plants, but excess iron is toxic due to its catalytic role in the formation of hydroxyl radicals. Thus, iron uptake is highly regulated and induced only under iron deficiency. The mechanisms of iron uptake in roots are well characterized, but less is known about how plants perceive iron deficiency. We show that a basic helix\u2013loop\u2013helix (bHLH) transcription factor Upstream Regulator of IRT1 (URI) acts as an essential part of the iron deficiency signaling pathway in\n                    <jats:italic>Arabidopsis thaliana<\/jats:italic>\n                    . The\n                    <jats:italic>uri<\/jats:italic>\n                    mutant is defective in inducing Iron-Regulated Transporter1 (IRT1) and Ferric Reduction Oxidase2 (FRO2) and their transcriptional regulators FER-like iron deficiency-induced transcription factor (FIT) and bHLH38\/39\/100\/101 in response to iron deficiency. Chromatin immunoprecipitation followed by sequencing (ChIP-seq) reveals direct binding of URI to promoters of many iron-regulated genes, including\n                    <jats:italic>bHLH38\/39\/100\/101<\/jats:italic>\n                    but not\n                    <jats:italic>FIT<\/jats:italic>\n                    . While URI transcript and protein are expressed regardless of iron status, a phosphorylated form of URI only accumulates under iron deficiency. Phosphorylated URI is subject to proteasome-dependent degradation during iron resupply, and turnover of phosphorylated URI is dependent on the E3 ligase BTS. The subgroup IVc bHLH transcription factors, which have previously been shown to regulate bHLH38\/39\/100\/101, coimmunoprecipitate with URI mainly under Fe-deficient conditions, suggesting that it is the phosphorylated form of URI that is capable of forming heterodimers in vivo. We propose that the phosphorylated form of URI accumulates under Fe deficiency, forms heterodimers with subgroup IVc proteins, and induces transcription of\n                    <jats:italic>bHLH38\/39\/100\/101<\/jats:italic>\n                    . These transcription factors in turn heterodimerize with FIT and drive the transcription of\n                    <jats:italic>IRT1<\/jats:italic>\n                    and\n                    <jats:italic>FRO2<\/jats:italic>\n                    to increase Fe uptake.\n                  <\/jats:p>","DOI":"10.1073\/pnas.1916892116","type":"journal-article","created":{"date-parts":[[2019,11,27]],"date-time":"2019-11-27T18:05:19Z","timestamp":1574877919000},"page":"24933-24942","update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":197,"title":["The iron deficiency response in\n                    <i>Arabidopsis thaliana<\/i>\n                    requires the phosphorylated transcription factor URI"],"prefix":"10.1073","volume":"116","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-6415-1201","authenticated-orcid":false,"given":"Sun A.","family":"Kim","sequence":"first","affiliation":[{"name":"Department of Biological Sciences, Dartmouth College, Hanover, NH 03755;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ian S.","family":"LaCroix","sequence":"additional","affiliation":[{"name":"Norris Cotton Cancer Center, Geisel School of Medicine at Dartmouth, Lebanon, NH 03756;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Scott A.","family":"Gerber","sequence":"additional","affiliation":[{"name":"Norris Cotton Cancer Center, Geisel School of Medicine at Dartmouth, Lebanon, NH 03756;"},{"name":"Department of Molecular and Systems Biology, Geisel School of Medicine at Dartmouth, Lebanon, NH 03756"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4718-4666","authenticated-orcid":false,"given":"Mary Lou","family":"Guerinot","sequence":"additional","affiliation":[{"name":"Department of Biological Sciences, Dartmouth College, Hanover, NH 03755;"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"341","published-online":{"date-parts":[[2019,11,27]]},"reference":[{"key":"e_1_3_4_1_2","doi-asserted-by":"crossref","first-page":"108","DOI":"10.1016\/0014-5793(92)80911-Y","article-title":"Biologically relevant metal ion-dependent hydroxyl radical generation. 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Schmidt, Dissecting iron deficiency-induced proton extrusion in Arabidopsis roots. New Phytol. 183, 1072\u20131084 (2009).","journal-title":"New Phytol."},{"key":"e_1_3_4_4_2","doi-asserted-by":"crossref","first-page":"1405","DOI":"10.1093\/pcp\/pcz076","article-title":"The age of coumarins in plant-microbe interactions","volume":"60","author":"Stringlis I. A.","year":"2019","unstructured":"I. A. Stringlis, R. de Jonge, C. M. J. Pieterse, The age of coumarins in plant-microbe interactions. Plant Cell Physiol. 60, 1405\u20131419 (2019).","journal-title":"Plant Cell Physiol."},{"key":"e_1_3_4_5_2","doi-asserted-by":"crossref","first-page":"694","DOI":"10.1038\/17800","article-title":"A ferric-chelate reductase for iron uptake from soils","volume":"397","author":"Robinson N. J.","year":"1999","unstructured":"N. J. Robinson, C. M. Procter, E. L. Connolly, M. L. Guerinot, A ferric-chelate reductase for iron uptake from soils. 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Li ., The iron deficiency response regulators IAA-LEUCINE RESISTANT3 and bHLH104 possess different targets and have distinct effects on photosynthesis in Arabidopsis. J. Plant Sci. 62, 109\u2013119 (2019).","journal-title":"J. Plant Sci."},{"key":"e_1_3_4_22_2","doi-asserted-by":"crossref","first-page":"1223","DOI":"10.1105\/tpc.001388","article-title":"IRT1, an Arabidopsis transporter essential for iron uptake from the soil and for plant growth","volume":"14","author":"Vert G.","year":"2002","unstructured":"G. Vert ., IRT1, an Arabidopsis transporter essential for iron uptake from the soil and for plant growth. Plant Cell 14, 1223\u20131233 (2002).","journal-title":"Plant Cell"},{"key":"e_1_3_4_23_2","doi-asserted-by":"crossref","first-page":"587","DOI":"10.1023\/A:1019942200164","article-title":"Knock-out of Arabidopsis metal transporter gene IRT1 results in iron deficiency accompanied by cell differentiation defects","volume":"50","author":"Henriques R.","year":"2002","unstructured":"R. Henriques ., Knock-out of Arabidopsis metal transporter gene IRT1 results in iron deficiency accompanied by cell differentiation defects. Plant Mol. Biol. 50, 587\u2013597 (2002).","journal-title":"Plant Mol. Biol."},{"key":"e_1_3_4_24_2","doi-asserted-by":"crossref","first-page":"589","DOI":"10.1046\/j.1365-313X.2002.01381.x","article-title":"The metal ion transporter IRT1 is necessary for iron homeostasis and efficient photosynthesis in Arabidopsis thaliana","volume":"31","author":"Varotto C.","year":"2002","unstructured":"C. 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Hong ., Identification and characterization of transcription factors regulating Arabidopsis HAK5. Plant Cell Physiol. 54, 1478\u20131490 (2013).","journal-title":"Plant Cell Physiol."},{"key":"e_1_3_4_27_2","doi-asserted-by":"crossref","first-page":"srep00090","DOI":"10.1038\/srep00090","article-title":"Proteome-wide post-translational modification statistics: Frequency analysis and curation of the swiss-prot database","volume":"1","author":"Khoury G. A.","year":"2011","unstructured":"G. A. Khoury, R. C. Baliban, C. A. Floudas, Proteome-wide post-translational modification statistics: Frequency analysis and curation of the swiss-prot database. Sci. Rep. 1, srep00090 (2011).","journal-title":"Sci. Rep."},{"key":"e_1_3_4_28_2","doi-asserted-by":"crossref","first-page":"1445","DOI":"10.1093\/protein\/10.12.1445","article-title":"Effects of sulfhydryl regents on the activity of lambda Ser\/Thr phosphoprotein phosphatase and inhibition of the enzyme by zinc ion","volume":"10","author":"Zhuo S.","year":"1997","unstructured":"S. Zhuo, J. E. Dixon, Effects of sulfhydryl regents on the activity of lambda Ser\/Thr phosphoprotein phosphatase and inhibition of the enzyme by zinc ion. Protein Eng. 10, 1445\u20131452 (1997).","journal-title":"Protein Eng."},{"key":"e_1_3_4_29_2","doi-asserted-by":"crossref","first-page":"D828","DOI":"10.1093\/nar\/gkp810","article-title":"PhosPhAt: The Arabidopsis thaliana phosphorylation site database. An update","volume":"38","author":"Durek P.","year":"2010","unstructured":"P. Durek ., PhosPhAt: The Arabidopsis thaliana phosphorylation site database. An update. 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Plant. 163, 282\u2013296 (2018).","journal-title":"Physiol. Plant."},{"key":"e_1_3_4_48_2","doi-asserted-by":"crossref","first-page":"1433","DOI":"10.1111\/nph.15753","article-title":"Transcriptional integration of the responses to iron availability in Arabidopsis by the bHLH factor ILR3","volume":"223","author":"Tissot N.","year":"2019","unstructured":"N. Tissot ., Transcriptional integration of the responses to iron availability in Arabidopsis by the bHLH factor ILR3. New Phytol. 223, 1433\u20131446 (2019).","journal-title":"New Phytol."},{"key":"e_1_3_4_49_2","doi-asserted-by":"crossref","first-page":"893","DOI":"10.1104\/pp.112.208603","article-title":"Reciprocal interaction of the circadian clock with the iron homeostasis network in Arabidopsis","volume":"161","author":"Hong S.","year":"2013","unstructured":"S. Hong, S. A. Kim, M. L. Guerinot, C. R. McClung, Reciprocal interaction of the circadian clock with the iron homeostasis network in Arabidopsis. 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