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In parallel, for pairs of ancestral tRNAs with complementary anticodons, we present updated evidence of concerted complementarity of the second bases in the acceptor stems.
Indeed, the number of inferred structures with the full complement of seven paired bases in the acceptor arm was only six in D. brevis and 11 in D. folliculorum.
Among the evidence in support of such ambiguity especially telling is the fact that the concerted complementarity of 2nd bases in the acceptor is observed for pairs of tRNAs with completely complementary anticodons, but is not observed for pairs in which only the central bases are complementary [ 22].
2) If the anticodon stem/loop and acceptor stem are homologous (in the true sense of having common ancestry instead of merely sequence or structural similarity), then the two codes a) shifted on the structure and b) became double stranded in the acceptor stem, whereby the proposed identity of bases in the acceptor stem with the anticodon is much more difficult to reconcile.
Worth noting are the pairs (underlined in Table 1) in which one "partner" has different second bases in the acceptor stem in different domains (for example, "G" in Bacteria and "C" in Archaea) and the other partner possesses the domain-specific complementary second bases ("C" in Bacteria and "G" in Archaea, respectively).
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In particular, TmcA mainly discriminates differences in bases in the anticodon stem, whereas TilS recognizes base pairs in the acceptor stem.
tRNA molecules are generally 75-87 nuclongides long and form clover-leaf shaped structures through base pairing in the acceptor stem; D-stem, TΨC stem and anticodon stem.
The homologous Andalucia tRNASer gga), for example, has a fully base-paired acceptor stem encoded in the mtDNA, and in other cases, nonstandard base pairs in the acceptor stem are overwhelmingly GċU or UċG, which in Seculamonas have been shown not to be edited (nor is an acceptor stem U × U mismatch in the mitochondrial tRNAHis of this organism) (Leigh and Lang 2004).
For lysidine formation in tRNAIle, we previously identified the tRNAIle-lysidine synthetase (tilS) that strictly discriminates tRNAIle from tRNAMet by recognizing two consecutive base pairs in the acceptor stem (Soma et al, 2003; Ikeuchi et al, 2005).
But the latter are not necessarily needed (or at least are not as much needed as the ribozymes) to provide aa-specific recognition of tRNAs in the bulged bases of the acceptor helix and/or in single-stranded anticodon loop.
To measure the extent of the dual complementarity (DC), we counted the number of tRNA pairs with complementary anticodons in which the second bases of the acceptor helix were also complementary, and divided it by the total number of pairs [ 22].
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