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FIT (FER\u2010LIKE IRON DEFICIENCY\u2010INDUCED TRANSCRIPTION FACTOR) and four bHLH Ib members (bHLH38, bHLH39, bHLH100 and bHLH101) are the key regulators of Fe homeostasis. Here, we reveal that FIT and bHLH Ib control the up\u2010regulation of Cu\u2010uptake genes (<jats:italic>COPT2<\/jats:italic>, <jats:italic>FRO4<\/jats:italic> and <jats:italic>FRO5<\/jats:italic>) by Fe deficiency, and Cu is required for improving plant growth under Fe\u2010deficiency conditions. The induction of Cu\u2010uptake gene expression and the elevation of Cu concentration are inhibited in the <jats:italic>fit\u20102<\/jats:italic> or <jats:italic>bhlh4x<\/jats:italic> (the quadruple mutant of four bHLH Ib genes) under Fe\u2010deficiency conditions. The dual overexpression of both <jats:italic>bHLH38<\/jats:italic> (or <jats:italic>bHLH39<\/jats:italic>) and <jats:italic>FIT<\/jats:italic> activates the expression of <jats:italic>COPT2<\/jats:italic>, <jats:italic>FRO4<\/jats:italic> and <jats:italic>FRO5<\/jats:italic> and increases Cu accumulation. Furthermore, bHLH Ib proteins directly bind to the promoters of <jats:italic>COPT2, FRO4<\/jats:italic> and <jats:italic>FRO5<\/jats:italic>. Either Cu supplement or overexpression of <jats:italic>COPT2<\/jats:italic> or <jats:italic>FRO4<\/jats:italic> improves the growth of <jats:italic>fit\u20102<\/jats:italic> under Fe\u2010deficiency conditions. Moreover, the induction of <jats:italic>COPT2, FRO4<\/jats:italic> and <jats:italic>FRO5<\/jats:italic> by Fe deficiency is independent of SPL7, a central regulator of Cu\u2010deficiency responses. This work through the link between bHLH Ib\/FIT and <jats:italic>COPT2\/FRO4\/FRO5<\/jats:italic> under Fe\u2010deficiency conditions establishes a new relationship between Cu and Fe homeostasis.<\/jats:p>","DOI":"10.1111\/pce.14000","type":"journal-article","created":{"date-parts":[[2021,1,19]],"date-time":"2021-01-19T18:22:18Z","timestamp":1611080538000},"page":"1679-1691","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":78,"title":["<scp>FIT<\/scp> and <scp>bHLH<\/scp> Ib transcription factors modulate iron and copper crosstalk in Arabidopsis"],"prefix":"10.1111","volume":"44","author":[{"given":"Yuerong","family":"Cai","sequence":"first","affiliation":[{"name":"CAS Key Laboratory of Tropical Plant Resources and Sustainable Use, Xishuangbanna Tropical Botanical Garden, The Innovative Academy of Seed Design Chinese Academy of Sciences  Kunming China"},{"name":"Center of Economic Botany, Core Botanical Gardens Chinese Academy 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