Higher order structural elements in ribosomal RNAs: pseudo-knots and the use of noncanonical pairs

The data base of prokaryotic small subunit ribosomal RNAs alone now numbers more than 400 sequences, while that for the large subunit rRNAs numbers more than 70 when eukaryotic, mitochondrial, and plastid sequences are also included. Comparisons among these rRNA sequences reveal a number of positions that covary in composition, suggestive of higher order structural elements; 5 such structures are reported for the small subunit rRNA and 15 for the large subunit rRNA. While some of these are properly (small) secondary structural elements, the majority would have to be classified as more complex "tertiary" interactions, which in some cases bring together diverse areas in the secondary structural diagram. A number of the covariances are not of the canonical type, indicating non-Watson-Crick interactions.