Abstract of the PDB Structure's related Publication:
The 23S rRNA nucleotide m(2)G2445 is highly conserved in bacteria, and in Escherichia coli this modification is added by the enzyme YcbY. With lengths of around 700 amino acids, YcbY orthologs are the largest rRNA methyltransferases identified in Gram-negative bacteria, and they appear to be fusions from two separate proteins found in Gram-positives. The crystal structures described here show that both the N- and C-terminal halves of E. coli YcbY have a methyltransferase active site and their folding patterns respectively resemble the Streptococcus mutans proteins Smu472 and Smu776. Mass spectrometric analyses of 23S rRNAs showed that the N-terminal region of YcbY and Smu472 are functionally equivalent and add the m(2)G2445 modification, while the C-terminal region of YcbY is responsible for the m(7)G2069 methylation on the opposite side of the same helix (H74). Smu776 does not target G2069, and this nucleotide remains unmodified in Gram-positive rRNAs. The E.coli YcbY enzyme is the first example of a methyltransferase catalyzing two mechanistically different types of RNA modification, and has been renamed as the Ribosomal large subunit methyltransferase, RlmKL. Our structural and functional data provide insights into how this bifunctional enzyme evolved.
RlmKL is a bifunctional protein (700 amino acids) that catalyzes the in vitro formation of two different methylated residues (m7G and m2G) within the same helix 74 in domain IV (Peptidyl Transferase Center) of 23S rRNA purified from the ycbY knock-out strain (positions 2069 and 2445 respectively). Assembled 50 S subunits are not a substrate for the RlmKL. It contains two Rossmann-fold methyltransferase domains. The N-terminal domain (RlmL domain) is a member of COG0116 and consists of a MTase domain and a RNA-binding THUMP domain. The C-terminal domain (RlmK domain) is a MTase and member of COG1092. AdoMet is the methyl group donor. RlmKL was shown to have an activity to unwind 23S rRNA helix 74 during substrate recognition and methylation. The proposed mechanism for the methylation of two target sites: first the 2069 m7G is introduced by RlmK domain, then the 2445 m2G by RlmL domain. The presence of RlmK domain facilitates methylation in position 2445 but the reactions are independent. P-loop of helix 80 is critical for substrate recognition. Phylogenetic analysis suggests that bifunctional RlmKL is only present in Gammaproteobacteria, and that the separated RlmK and RlmL are found in the genus Neisseria.