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NifS and NifU (encoded by  nifS  and  nifU ) are generally dedicated to biogenesis of the nitrogenase Fe–S cluster in diazotrophs. However,  nifS  and  nifU  are not found in N 2 -fixing  Paenibacillus  strains, and the mechanisms involved in Fe–S cluster biosynthesis of nitrogenase is not clear. Here, we found that the genome of  Paenibacillus polymyxa  WLY78 contains the complete  sufCDSUB  operon, a partial  sufC2D2B2  operon, a  nifS -like gene, two  nifU -like genes ( nfuA -like and  yutI ), and two  iscS  genes. Deletion and complementation studies showed that the  sufC ,  sufD,  and  sufB  genes of the  sufCDSUB  operon, and  nifS -like and  yutI  genes were involved in the Fe–S cluster biosynthesis of nitrogenase. Heterologous complementation studies demonstrated that the  nifS -like gene of  P. polymyxa  WLY78 is interchangeable with  Klebsiella oxytoca nifS , but  P. polymyxa  WLY78 SufCDB cannot be functionally replaced by  K. oxytoca  NifU. In addition,  K. oxytoca nifU  and  Escherichia coli nfuA  are able to complement the  P. polymyxa  WLY78  yutI  mutant. Our findings thus indicate that the NifS-like and SufCDB proteins are the specific sulfur donor and the molecular scaffold, respectively, for the Fe–S cluster formation of nitrogenase in  P. polymyxa  WLY78. YutI can be an Fe–S cluster carrier involved in nitrogenase maturation in  P. polymyxa  WLY78.

Identification of Genes Involved in Fe–S Cluster Biosynthesis of Nitrogenase in Paenibacillus polymyxa WLY78