At the termini of linear chromosomes, nucleotide sequences known as telomeres repeat tandemly. Their primary function is to safeguard chromosome ends from degradation and end-to-end fusion, thereby ensuring genome stability . The number of repeats in human telomere sequences varies between cells and even between chromosomes, and they are composed of C1-3A/TG1-3 and TTAGGG/CCCTAA repeats . After DNA replication is finished, the terminal primer is degraded, which leads to the loss of a small segment of DNA at the 5' end of the lagging strand . Additionally, the linear template's average length of approximately 250 nucleotides cannot be replicated at the end . The "end replication problem" is named after the progressive shortening of telomeres that results from the incomplete replication of DNA polymerase at the chromosome end with each successive cell division . DNA double-strand breaks, which are triggered by short or dysfunctional telomeres, are identified as the cause of cell replicative senescence .