ciceri (Figure  1, Figure  2) It is likely that an exchange betw

ciceri (Figure  1, Figure  2). It is likely that an exchange between M. loti and a common

ancestor of S. meliloti, S. medicae and S. this website fredii NGR234 occurred. M. loti is located in the same clade as the Brucella and O. anthropi in the species tree (Figure  2). Despite this, M. loti contains many of the genes corresponding to the adonitol and L-arabitol type loci of other species that cluster close to the base of the species tree such as Bradyrhizobium spp. (Figure  2). The presence of these factors in addition to the chimeric composition of the M. loti locus leads us to hypothesise that an ancestor of M. loti may have contained both an erythritol locus like that of the Brucella as well as a polyol type locus like that seen in the Bradyrhizobia, A. cryptum and V. eiseniae. The lalA, rbtB, rbtC suboperon appears to be the key component of the polyol locus in the Bradyrhizobium type loci (Figure  1). Among the Epigenetics inhibitor 19 loci identified, these three genes can be linked into a suboperon, embedded within the main locus (eg. R. litoralis) or split among two transcriptional units (see A. cryptum or V. eiseniae). As well, the gene module (or suboperon) eryR, tpiB- rpiB is presumably

found in all erythritol utilizing bacteria. The acquisition of this module along with the lalA, rbtB and rbtC suboperon may have allowed for the evolution of the more complex S. meliloti type locus (see Figure  2). The absence of fucA in S. fredii NGR234 and M. loti appears to be an example of the loss of an “ORFan” gene event having occurred. The gene is Temsirolimus order still present in S. meliloti however it has been shown that it is not necessary for the catabolism

of erythritol, adonitol, or L-arabitol [15]. It is likely that it was lost during the divergence of M. loti and S. fredii NGR234 from their common ancestors to S. meliloti. If this is true, it may be reasonable to assume that fucA may eventually also be lost from the S. meliloti erythritol locus. In S. meliloti, erythritol uptake Cytidine deaminase has been shown to be carried out by the proteins encoded by mptABCDE[15, 16], whereas in R. leguminosarum growth using erythritol is dependent upon the eryEFG[20]. Although both transporters appear to carry out the same function, the phylogenetic analysis clearly shows that they have distinct ancestors and may be best classified as analogues rather than orthologues (Figure  3). In addition, it has been shown that MptABCDE is also capable of transporting adonitol and L-arabitol [15]. We note that these polyols appear to have stereo-chemical identity over three carbons and that EryA of S. meliloti can also use adonitol and L-arabitol as substrates [15]. It is unknown whether EryA from R. leguminosarum has the ability to interact with these substrates. The three distinct groups of loci we have identified probably correspond to the metabolic potential of these regions to utilize polyols. The locus of S.

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