The kinetochore forms in eukaryotes and assembles on the centromere and links the chromosome to microtubule polymers from the mitotic spindle during mitosis and meiosis. The kinetochore contains two regions: an inner kinetochore, which is tightly associated with the centromere DNA; and an outer kinetochore, which interacts with microtubules. "Monocentric" organisms, including vertebrates, fungi, and most plants, have a single centromeric region on each chromosome which assembles one kinetochore. "Holocentric" organisms, such as nematodes, assemble a kinetochore along the entire length of a chromosome. During mitosis, which occurs after chromosomes are duplicated during S phase, two identical "sister" chromatids are held together each with its own kinetochore which face in opposing directions and attach to opposite poles of the mitotic spindle. Following the transition from metaphase to anaphase, the sister chromatids separate from each other, and the individual kinetochores on each chromatid drive their movement to the spindle poles that will define the two new daughter cells. Thus, the kinetochore is essential for the chromosome segregation that is classically associated with mitosis and meiosis.
Even the simplest kinetochores consist of more than 45 different proteins. Many of these proteins are conserved throughout eukaryote species, including a specialized histone H3 variant (called CENP-A or CenH3) which helps the kinetochore associate with DNA. Other proteins in the kinetochore attach it to the microtubules of the mitotic spindle. There are also motor proteins, including both dynein and kinesin, which generate forces that move chromosomes during mitosis. Other proteins, such as MAD2 monitor the microtubule attachment as well as the tension between sister kinetochores and activate the spindle checkpoint to arrest the cell cycle when either of these is absent.