Micronucleus is the name given to the small nucleus that forms whenever a chromosome or a fragment of a chromosome is not incorporated into one of the daughter nuclei during cell division. It usually is a sign of genotoxic events and chromosomal instability. Micronuclei are commonly seen in cancerous cells and may indicate genomic damage events that can increase the risk of developmental or degenerative diseases. Micronuclei form during anaphase from lagging acentric chromosome or chromatid fragments caused by incorrectly repaired or unrepaired DNA breaks or by nondisjunction of chromosomes. This incorrect segregation of chromosomes may result from hypomethylation of repeat sequences present in pericentromeric DNA, irregularities in kinetochore proteins or their assembly, dysfunctional spindle apparatus, or flawed anaphase checkpoint genes. Micronuclei can contribute to genome instability by promoting a catastrophic mutational event called chromothripsis. Many micronucleus assays have been developed to test for the presence of these structures and determine their frequency in cells exposed to certain chemicals or subjected to stressful conditions. The term micronucleus may also refer to the smaller nucleus in ciliate protozoans, such as the Paramecium. In mitosis it divides by fission, and in conjugation a pair of gamete micronuclei undergo reciprocal fusion to form a zygote nucleus, which gives rise to the macronuclei and micronuclei of the individuals of the next cycle of fission. Discovery Micronuclei in newly formed red blood cells in humans are known as Howell-Jolly bodies because these structures were first identified and described in erythrocytes by hematologists William Howell and Justin Jolly. These structures were later found to be associated with deficiencies in vitamins such as folate and B12. The relationship between formation of micronuclei and exposure to environmental factors was first reported in root tip cells exposed to ionizing radiation. Micronucleus induction by a chemical was first reported in Ehrlich ascites tumor cells treated with colchicine. Formation Micronuclei primarily result from acentric chromosome fragments or lagging whole chromosomes that are not included in the daughter nuclei produced by mitosis because they fail to correctly attach to the spindle during the segregation of chromosomes in anaphase. These full chromosomes or chromatid fragments are eventually enclosed by a nuclear membranes and are structurally similar to conventional nuclei, albeit smaller in size. This small nucleus is referred to as a micronucleus. The formation of micronuclei can only be observed in cells undergoing nuclear division and can be clearly seen using cytochalasin B