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Iron (Fe) deficiency in plants limits crop growth and productivity. Molecular mechanisms that plants use to sense and respond to Fe deficiency by coordinated expression of Fe-uptake genes are not fully understood. The C940-fe chlorotic melon ( Cucumis melo ) mutant known as  fefe  is unable to upregulate Fe-uptake genes, however, the  FeFe  gene had not been identified. In this study, we used two F 2  mapping populations to map and identify the  FeFe  gene as  bHLH38 , a homolog of subgroup Ib  bHLH  genes from  Arabidopsis thaliana  that are involved in transcriptional regulation of Fe-uptake genes in partnership with the  FIT  gene. A Ty1-copia type retrotransposon insertion of 5.056 kb within  bHLH38  is responsible for the defect in  bHLH38  in  fefe , based on sequencing and expression analysis. This retrotransposon insertion results in multiple non-functional transcripts expected to result in an altered and truncated protein sequence. Hairy root transformation of  fefe  plants using wild-type  bHLH38  resulted in functional complementation of the chlorotic  fefe  phenotype. Using a yeast-2-hybrid assay, the transcription factor Fit interacted with the wild-type bHLH38 protein, but did not interact with the  fefe  bHLH38 protein, suggesting that heterodimer formation of Fit/bHLH38 to regulate Fe-uptake genes does not occur in  fefe  roots. The second subgroup Ib  bHLH  gene in the melon genome is not functionally redundant to  bHLH38 , in contrast to Arabidopsis where four subgroup Ib  bHLH  genes are functionally redundant. Whereas the Arabidopsis  bHLH  transcript levels are upregulated by Fe deficiency, melon  bHLH38  was not regulated at the transcript level. Thus, the  fefe  mutant may provide a platform for studying  bHLH38  genes and proteins from other plant species.

Mapping and Characterization of the fefe Gene That Controls Iron Uptake in Melon (Cucumis melo L.)