In spite of considerable divergence in the centromere DNA sequence, basic architecture of a KT is evolutionarily conserved from yeast to humans. However, the identification

of a large number of KT proteins paved the way of understanding conserved and diverged regulatory steps that lead to the formation of a multiprotein KT super-complex on the centromere DNA in different organisms. Because it is a daunting task to summarize the entire spectrum of information in a minireview, we focus here on the recent understanding in the process selleck chemicals llc of KT assembly in three yeasts: Saccharomyces cerevisiae, Schizosaccharomyces pombe and Candida albicans. Studies in these unicellular organisms suggest that although the basic process of KT assembly remains DAPT the same, the dependence of a conserved protein for its KT localization may vary in these organisms. The precise transmission of the genetic information from one generation to the next during the mitotic cell cycle is extremely important for a eukaryotic organism. This process involves faithful duplication of the whole genome during S phase followed by segregation of the duplicated genome with high fidelity during mitosis. The molecular mechanisms that ensure equal distribution of duplicated chromosomes in mitosis require proper assembly of a large multiprotein complex at the centromere (CEN),

known as the kinetochore (KT). The primary function Florfenicol of a KT is to attach the chromosome to the dynamic plus ends of spindle microtubules (MTs), a crucial step in segregation of chromosomes. KTs are also associated with the formation of heterochromatin at the centromeric/pericentric regions and maintenance of cohesion between sister chromatids till anaphase onset (Cleveland et al., 2003; Cheeseman & Desai, 2008). Additionally, a KT is involved in the recruitment of the spindle assembly checkpoint machinery that monitors the KT-MT attachment and initiates signals to prevent cell cycle progression if an error persists. Once all the chromosomes are bi-orientated, separation of two sister chromatids marks the onset of anaphase. Any defect

in the KT structure can disrupt KT–MT interaction that may result in an unequal distribution of chromosomes leading to aneuploidy. In metazoan cells, the nuclear envelope breaks down during mitosis that allows KT–MT interaction to facilitate bi-oriented chromosomes to arrange on a plane known as the metaphase plate (Nasmyth, 2001; Guttinger et al., 2009). In contrast, the nuclear envelope never breaks down in budding yeasts and thus cells undergo closed mitosis without formation of a metaphase plate (Straight et al., 1997; Sazer, 2005; De Souza & Osmani, 2007). Existence of a metaphase plate is unlikely in Schizosaccharomyces pombe and Candida albicans as well. Interestingly, a semi-open mitosis has been reported recently in fission yeast Schizosaccharomyces japonicus (Aoki et al., 2011; Yam et al., 2011).