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To acquire iron (Fe), graminaceous plants secrete mugineic acid family phytosiderophores through the phytosiderophore efflux transporter TOM1 and take up Fe in the form of Fe(III)–phytosiderophore complexes.  Yellow stripe 1  ( ys1 ) and  ys3  are recessive mutants of maize ( Zea mays  L.) that show typical symptoms of Fe deficiency, i.e., interveinal chlorosis of the leaves. The  ys1  mutant is defective in the Fe(III)–phytosiderophore transporter YS1 and is therefore unable to take up Fe(III)–phytosiderophore complexes. While the  ys3  mutant has been shown to be defective in phytosiderophores release, the causative gene has not been identified. The present study was performed to characterize the expression profiles of the genes in  ys1  and  ys3  mutants to extend our understanding of Fe homeostasis in maize. Using quantitative real-time polymerase chain reaction, we assessed changes in the levels of gene expression in response to Fe deficiency of genes involved in Fe homeostasis, such as those related to phytosiderophore biosynthesis and Fe transport. As with other crops, these Fe deficiency-inducible genes were also upregulated in maize. In addition, these Fe deficiency-inducible genes were upregulated in both the  ys1  and  ys3  mutants, even under Fe-sufficient conditions. Indeed, the Fe concentrations in the roots of  ys1  and  ys3  plants were lower than that of wild-type controls. These results suggest that  ys1  and  ys3  are Fe-deficient during growth in the presence of Fe. In agreement with previous reports, the level of  YS1  expression decreased in the  ys1  mutant. Moreover, the expression level of a homolog of  TOM1  in maize decreased significantly in the  ys3  mutant. Unspliced introns of  ZmTOM1  were detected only in  ys3 , and not in  YS1YS3  or  ys1 , suggesting that  ZmTOM1  may be involved in the  ys3  phenotype.

Characterizing the Crucial Components of Iron Homeostasis in the Maize Mutants ys1 and ys3