Anaphase lag

mosaicism and development as well as the progression and heterogeneity of cancers.^[3]

microtubules originating from both spindle poles of the dividing cell. The merotelic attachments can occur in two ways: centrosome spindle attachments from both poles on the same chromatid kinetochore^[5] or the formation of a third centrosome whose microtubule spindles attach to a chromatid kinetochore.^[6] Because the chromatid is being pulled in two opposing directions or away from the correct centriole, it cannot migrate to the mass of segregated chromatids at either pole. If the migration is significantly delayed the reformation of nuclei will begin to occur without a full complement of chromosomes. This nuclear envelope formation is also seen for the lone lagging sister chromatid, forming a micronucleus. The micronucleus has the capacity to persist in the daughter cell but with abnormal replication and maintenance machinery. This allows for the accumulation of mutations, increasing the potential for future miss-segregation events.^[2] In total these events cause problematic aneuploid cells with increased genomic instability. This has important implications in the development and persistence of cancers as well as debilitating developmental diseases.^[7]

genomic instability, referring to the increased frequency in sequence mutation, chromosome rearrangement, and aneuploidy.^[8] The instability allows a cancerous growth to increasingly diverge from normal cell growth and division, with the potential to gain new traits such as angiogenesis, immune system evasion, and loss of cell cycle checkpoint genes. Aneuploidy is a drastic divergence from the normal karyotype, as such the potential heterogeneity within these cells makes diagnosis and treatment increasingly difficult.^[7]

knock-outs showed increased frequency of lagging anaphase chromatids; subsequent gene correction in human glioblastoma cell lines reduced the occurrence of this genomic instability.^[11]

genomic instability in cancer prognosis/treatment continues, identifying the causes and consequences of mechanisms such as anaphase lag will be critical to understanding how cancer develops as well as developing better multi-target therapies.^{[citation needed}