Soil bacteria that also form mutualistic symbioses in plants encounter two major levels of selection. One occurs during adaptation to and survival in soil, and the other occurs in concert with host plant speciation and adaptation. Actinobacteria from the genus Frankia are facultative symbionts that form N 2 -fixing root nodules on diverse and globally distributed angiosperms in the "actinorhizal" symbioses. Three closely related clades of Frankia sp. strains are recognized; members of each clade infect a subset of plants from among eight angiosperm families. We sequenced the genomes from three strains; their sizes varied from 5.43 Mbp for a narrow host range strain (Frankia sp. strain HFPCcI3) to 7.50 Mbp for a medium host range strain (Frankia alni strain ACN14a) to 9.04 Mbp for a broad host range strain (Frankia sp. strain EAN1pec.) This size divergence is the largest yet reported for such closely related soil bacteria (97.8%-98
The Frankia genome contains two truncated hemoglobin genes (hboN and hboO) whose functions remain to be determined. Nitric oxide (NO) generated by the addition of 400 M SNAP (S-nitroso-N-acetylpenicillamine) caused a 10-fold increase in hboN gene expression but had no effect on hboO expression. The addition of the NO scavenger, carboxy-PT10, reduced the effect of SNAP. hboO gene expression increased under low-oxygen conditions, while hboN expression was unaffected. These results suggest that HboN may function in protection from nitrosative stress and that HboO may act as an oxygen transport molecule for increased respiration in hypoxic environments.
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