Genetics and regulation of nif and related genes in Klebsiella pneumoniae

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Abstract
Seventeen genes specifically required for nitrogen fixation are clustered on the chromosome of Klebsiella pneumoniae and form a complex regulon that is organized into eight transcriptional units. The nif promoters are representative of a new class of promoter, the members of which lack the consensus sequences normally found in prokaryotic promoters, nif gene transcription is positively controlled and requires: (1) the ntrA gene product, which replaces the rpoD -encoded sigma subunit of RNA polymerase to allow recognition of nif promoter sequences; and (2) the product of either the nitrogen regulation gene ntrC or the specific nif regulatory gene, nifA , which are both transcriptional activators. Most nif promoters require an upstream activator sequence (UAS) for nifA -mediated activation. The UAS acts independently of orientation and can function when placed 2 kilobases upstream from the transcription start site. Current evidence suggests that activation requires an interaction between proteins bound at the UAS and at the downstream nif promoter consensus, possibly via a loop in the DNA structure. Transcription of nif is modulated by the ntrB and nifL gene products. Both proteins can ‘sense’ environmental changes: ntrB prevents activation by ntrC in response to excess nitrogen whereas nifL prevents activation by nifA in response to fixed nitrogen and oxygen. The C-terminal end of ntrB shows clear homology at the amino acid level with a number of diverse control proteins involved in regulation or sensory transduction. Each member of this family interacts with another protein component showing homology to the N-terminal sequence of ntrC , but not to nifA . The significance of these protein homologies is discussed.