The requirement of de novo acetylation during DNA replication implies that both acetylated and non-acetylated forms of Smc3 are important throughout the cell cycle. The characterisation of the exact timing and pattern of Smc3 acetylation, by chromatin immunoprecipitation using a specific antibody against the acetylated protein, would be important to understand the role that each form plays. Additionally, it would also be interesting to understand the correlation between the timing of acetylation and the binding of replisome components in respect to the replication fork, to further elucidate the relationship between cohesin acetylation and replication fork progression. Eco1 is thought to associate with the DNA replication fork through its physical interaction with PCNA and this interaction is essential for cohesion establishment in S-phase (Lengronne et al., 2006; Moldovan et al., 2006). It would then be interesting to see whether this is also needed for Smc3 acetylation. Studies in vertebrates showed that although Smc3 acetylation occurs independently of DNA replication, functional acetylation occurs only in association with the replication machinery (Song et al., 2012). The recruitment of Eco1 to the replication fork by PCNA could therefore be the essential step to promote Smc3 acetylation and establish cohesion. However, I cannot exclude that additional factors at the replication fork may also be required to trigger Smc3 acetylation. Thus, studies to understand the timing of cohesin acetylation and its correlation with the binding of replisome components to DNA would provide us further insight about the proteins involved in the establishment of sister chromatid cohesion. If Smc3 acetylation indeed depends on the replication machinery, it would then be interesting to examine the consequences of a delay in S-phase in cohesin acetylation and subsequent establishment.
Several lines of evidence suggest that cohesion establishment is tightly coupled to DNA replication during S-phase. Further experiments involving factors that associate with the replication fork machinery and that contribute to cohesion establishment will enhance our understanding about the mechanism underlying sister chromatid cohesion. As Ctf4 and Chl1 act in an independent pathway of Eco1 it would be of interest to delineate this Eco1-independent pathway of cohesion establishment. A genetic approach could unveil the pathways targeted by these factors during cohesion establishment. The nature of mutations that modulate the requirement of Ctf4 and Chl1 for sister chromatid cohesion establishment would be expected to carry important information as to the mechanism involved.
In addition, it would also be important to determine the molecular functions of Ctf4 and Chl1 individually. Little is known about Chl1; it is a helicase involved in the establishment but not maintenance of sister chromatid cohesion. It would be interesting to check whether the helicase activity of Chl1 is required for cohesion establishment. Moreover, it is unknown when and where Chl1 is loaded onto the replication fork, thereby chromosome-wide binding analyses could pinpoint exactly when and where Chl1 is loaded onto the replication fork. In turn, Ctf4 is a component of elongating replication forks that interacts with several replication proteins, including the GINS complex and DNA polymerase alpha/primase. It would therefore be interesting to study which of the Ctf4-mediated replisome interactions contributes to sister chromatid cohesion establishment, possibly by mapping and disrupting each interaction individually by targeted point mutations. Exploring the function of the Eco1-independent cohesion establishment pathway is likely to hold important insight to understanding replication-coupled cohesion establishment. Understanding how the sister chromatids become linked immediately after replication represents a significant step forward in our understanding of the mechanisms involved in chromosome segregation.
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