Abstract
Transcription presents challenges to genome stability both directly, by
altering genome topology and exposing single-stranded DNA to chemical
insults and nucleases, and indirectly by introducing potential obstacles
to the DNA replication machinery. Such obstacles include the RNA
polymerase holoenzyme itself, DNA bound regulatory factors,
G-quadruplexes and RNA::DNA hybrid structures known as R-loops. Here we
review the detrimental impacts of transcription on genome stability in
budding yeast, as well as the mitigating effects of
transcription-coupled DNA repair and of systems that maintain DNA
replication fork processivity and integrity. We conclude that the
impacts of transcription on DNA replication by the replisome must be
very mild with little direct mutagenic outcome under normal conditions.
However, transcription can significantly impair the fidelity of
replication fork rescue mechanisms, particularly Break Induced
Replication (BIR), which is used to restart collapsed replication forks
when other means fail. This leads to de novo mutations,
structural variation and extrachromosomal circular DNA formation that
contribute to genetic heterogeneity. By re-analysing published datasets,
we show that different classes of genes have different interactions with
the replisome, and that highly transcribed environment dependant genes
in S. cerevisiae tend to be located close to replication origins.
We have previously implicated interactions between replication origins
and BIR forks in adaptive transcription-induced copy number variation
events, which indicates that environment dependent genes are
preferentially located in genomic environments more prone to
instability, particularly under replication stress.