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Spx is a global regulator of genes that are induced by disulfide stress in  Bacillus subtilis . The regulon that it governs is comprised of over 120 genes based on microarray analysis, although it is not known how many of these are under direct Spx control. Most of the Spx-regulated genes (SRGs) are of unknown function, but many encode products that are conserved in low %GC Gram-positive bacteria. Using a gene-disruption library of  B. subtilis  genomic mutations, the SRGs were screened for phenotypes related to Spx-controlled activities, such as poor growth in minimal medium and sensitivity to methyglyoxal, but nearly all of the SRG mutations showed little if any phenotype. To uncover SRG function, the mutations were rescreened in an  spx  mutant background to determine which mutant SRG allele would enhance the  spx  mutant phenotype. One of the SRGs,  ytpQ  was the site of a mutation that, when combined with an  spx  null mutation, elevated the severity of the Spx mutant phenotype, as shown by reduced growth in a minimal medium and by hypersensitivity to methyglyoxal. The  ytpQ  mutant showed elevated oxidative protein damage when exposed to methylglyoxal, and reduced growth rate in liquid culture. Proteomic and transcriptomic data indicated that the  ytpQ  mutation caused the derepression of the Fur and PerR regulons of  B. subtilis . Our study suggests that the  ytpQ  gene, encoding a conserved DUF1444 protein, functions directly or indirectly in iron homeostasis. The  ytpQ  mutant phenotype mimics that of a  fur  mutation, suggesting a condition of low cellular iron. In vitro transcription analysis indicated that Spx stimulates transcription from the  ytpPQR  operon within which the  ytpQ  gene resides. The work uncovers a link between Spx and control of iron homeostasis.

Phenotype Enhancement Screen of a Regulatory spx Mutant Unveils a Role for the ytpQ Gene in the Control of Iron Homeostasis