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PhoU, a conserved protein that has been proposed to coordinate phosphate import, is a negative regulator of drug tolerance in most bacteria. In  Staphylococcus epidermidis , the role of PhoU in biofilm formation and drug tolerance has not yet been investigated. Two PhoU homologs in the genome of  S. epidermidis  have been identified by the presence of the conserved motif E(D)XXXD of PhoU. We separately constructed Δ phoU1  and Δ phoU2  mutants of  S. epidermidis  strain 1457. The Δ phoU2  mutant displayed growth retardation, a weakened biofilm formation capacity, a higher sensitivity to H 2 O 2 , and reduced tolerance to multiple antibiotics. However, deletion of  phoU1  had no effect on those. We compared the transcriptome profiles of the Δ phoU2  and Δ phoU1  mutants with that of the parent strain. In the Δ phoU2  mutant, expression of genes related to inorganic phosphate uptake was significantly upregulated ( pst  operon) and the levels of intracellular inorganic polyphosphate (polyP) were increased. In the Δ phoU2  mutant, expression of enzymes in the pentose phosphate pathway (PPP) was downregulated and less NADP (NADPH) was detected, consistent with the high sensitivity to H 2 O 2  and the growth retardation of the Δ phoU2  mutant. The upregulated expression of ATP synthase was consistent with the high intracellular ATP content in the Δ phoU2  mutant, which may have been related to the lower drug tolerance of the Δ phoU2  mutant. This study demonstrates that PhoU2, but not PhoU1, in  S. epidermidis  regulates bacterial growth, biofilm formation, oxidative stress, and drug tolerance in association with alterations to inorganic phosphate metabolism, the pentose phosphate pathway, galactose metabolism, the tricarboxylic acid (TCA) or citric cycle, glycolysis and gluconeogenesis, and respiratory reactions.   IMPORTANCE  PhoU is widely conserved throughout the bacterial kingdom and plays an important role in response to stress and metabolic maintenance. In our study, two PhoU homologs were found in  S. epidermidis . The function of  phoU2 , but not  phoU1 , in  S. epidermidis  is related to growth, drug tolerance, the oxidative stress response, polyP levels, and ATP accumulation. In addition,  phoU2  regulates biofilm formation. Hence,  phoU2  is a regulator of both drug tolerance and biofilm formation, which are two bacterial properties that present major challenges to the clinical treatment of infections. Analysis of differential gene expression revealed that  phoU2  is involved in fundamental metabolic processes, such as the PPP pathway. These findings indicate that  phoU2  is a crucial regulator in  S. epidermidis .

journal of bacteriology

PhoU2 but Not PhoU1 as an Important Regulator of Biofilm Formation and Tolerance to Multiple Stresses by Participating in Various Fundamental Metabolic Processes in Staphylococcus epidermidis