A physical mechanism that guarantees the accurate segregation of sister chromatids during mitosis arises from the ring shaped cohesin complex consisting of 4 subunits (SMC1A/B, SMC3, SCC1, and SA1/2 in humans). This complex encircles the two sister chromatids and resists the pulling force of microtubules.[8] The characteristic X-shape chromosomes are formed due to the centromeric cohesin protected by Shugoshin-PP2A complex.[9]
Kinetochore localization of Sgo1-PP2A is dependent upon phosphorylation on histone H2A of nucleosome, which is the important substrate of spindle checkpoint kinase BUB1.[10] Centromeric cohesin and H2A-pT120 specify two distinct pools of Sgo1-PP2A at inner centromeres and kinetochores respectively,[11] while the CDK1/cyclin B phosphorylation on Sgo1 is essential for Sgo1-PP2A to protect centromeric cohesin, not only for bringing PP2A to cohesin,[12] but also physically shield out the negative regulator WAPAL from cohesin.[13]
^"Human PubMed Reference:". National Center for Biotechnology Information, U.S. National Library of Medicine.
^"Mouse PubMed Reference:". National Center for Biotechnology Information, U.S. National Library of Medicine.
^"UniProt". www.uniprot.org. Retrieved 6 November 2022.
^Scanlan MJ, Gout I, Gordon CM, Williamson B, Stockert E, Gure AO, Jäger D, Chen YT, Mackay A, O'Hare MJ, Old LJ (Mar 2001). "Humoral immunity to human breast cancer: antigen definition and quantitative analysis of mRNA expression". Cancer Immunity. 1: 4. PMID12747765.
Wang X, Dai W (Oct 2005). "Shugoshin, a guardian for sister chromatid segregation". Experimental Cell Research. 310 (1): 1–9. doi:10.1016/j.yexcr.2005.07.018. PMID16112668.
Fu G, Hua S, Ward T, Ding X, Yang Y, Guo Z, Yao X (Jun 2007). "D-box is required for the degradation of human Shugoshin and chromosome alignment". Biochemical and Biophysical Research Communications. 357 (3): 672–8. doi:10.1016/j.bbrc.2007.03.204. PMID17448445.