During the process of translation, the ribosome reads the mRNA in three nucleotide strings, known as codons, to create a protein. Once a codon is translated, the ribosome moves three nucleotides down the mRNA strand to translate the next codon. During this process, however, the ribosome can shift backwards, moving only two nucleotides instead of the usual three. This process, known as a -1 programmed ribosomal frameshift (-1PRF), often occurs when the ribosome is currently reading a slippery site: a 7-nucleotide string following the NNNWWWH template (where NNN is AAA, UUU, GGG, or CCC, WWW is AAA or UUU and H is A, U, or C). While the behavior of frameshifting is well documented, the mechanism behind it remains relatively unknown. To further explore -1PRFs, we can look at the mRNA codon directly following codons within the ribosome, known as the +1 codon. While not actively being translated by the ribosome, +1 codons have been found to alter the process of translation, such as slowing it down by hydrogen bonding to CAR, a surface adjacent to the tRNA anticodon composed of three residues in the ribosome structure.
This thesis looks further into the hydrogen bonds between +1 codons and CAR to understand if they have an influence on -1PRFs. To do this, multiple approaches were taken including molecular dynamics simulations and analysis of the S. cerevisiae genome. This analysis suggested a model in which positive out-of-frame hydrogen bonds between the mRNA and CAR (hydrogen bonds from the 1st and 2nd nucleotide of the +1 codon to the A and R of CAR, respectfully) create a pulling effect that can increase the probability of -1PRFs while all other hydrogen bonds stabilize the frame, decreasing the probability of -1PRFs. Analysis of the S. cerevisiae genome found that +1 codons that decrease -1PRF probability are underrepresented after slippery sites, indicating that slippery sites with higher frameshift probabilities have been selected for. Additional analysis showed that slippery sites precede both premature stop codons and long open reading frames after the frame shift (suggesting that alternative proteins are created after a -1PRF), further suggesting that slippery sites are used as a form of post-transcriptional modification.
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