The Rhizobium leguminosarum tonB gene is required for the uptake of siderophore and haem as sources of iron

In the N-2-fixing bacterium Rhizobium leguminosarum, mutations in a homologue of tonB (tonB(RI)) block the import of vicibactin and haem as iron sources in free-living bacteria. TonB(RI) mutants were normal for growth with ferric dicitrate and slightly reduced for growth with haemogloblin as sole iron sources. The deduced TonB(RI) product is larger than that of (for example) Escherichia coli, on account of an extended N-terminal domain. Transcription of tonBRI was enhanced in low-Fe growth conditions; this was not controlled by Fur, nor Rpol, an Fe-regulated extracytoplasmic sigma factor. Upstream of tonB(RI) and transcribed divergently is an operon, hmuPSTUV, whose products are homologous to ABC transporters involved in haem uptake in pathogenic bacteria. Expression of hmuPSTUV was enhanced in low-Fe conditions, and hmu mutants show slightly diminished growth on haem as sole Fe source, suggesting that there is more than one system for the uptake of this molecule. hmuPSTUV expression appears to be from three closely linked promoters. Downstream of hmuPSTUV, a gene that may encode an extracytoplasmic sigma factor was identified, but this gene, rpoZ, did not affect the transcription of tonB(RI) or hmuPSTUV. Mutations in tonBRI, hmu genes and rpoZ did not affect symbiotic N-2 fixation in peas.