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In
genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinian friar wor ...
, a centimorgan (abbreviated cM) or map unit (m.u.) is a unit for measuring
genetic linkage Genetic linkage is the tendency of DNA sequences that are close together on a chromosome to be inherited together during the meiosis phase of sexual reproduction. Two genetic markers that are physically near to each other are unlikely to be sep ...
. It is defined as the distance between
chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins ar ...
positions (also termed loci or markers) for which the expected average number of intervening
chromosomal crossover Chromosomal crossover, or crossing over, is the exchange of genetic material during sexual reproduction between two homologous chromosomes' non-sister chromatids that results in recombinant chromosomes. It is one of the final phases of geneti ...
s in a single generation is 0.01. It is often used to infer distance along a chromosome. However, it is not a true physical distance.


Relation to physical distance

The number of
base pair A base pair (bp) is a fundamental unit of double-stranded nucleic acids consisting of two nucleobases bound to each other by hydrogen bonds. They form the building blocks of the DNA double helix and contribute to the folded structure of both DN ...
s to which it corresponds varies widely across the
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ...
(different regions of a chromosome have different propensities towards crossover) and it also depends on whether the
meiosis Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately r ...
in which the crossing-over takes place is a part of
oogenesis Oogenesis, ovogenesis, or oögenesis is the differentiation of the ovum (egg cell) into a cell competent to further develop when fertilized. It is developed from the primary oocyte by maturation. Oogenesis is initiated in the embryonic stage. O ...
(formation of female gametes) or
spermatogenesis Spermatogenesis is the process by which haploid spermatozoa develop from germ cells in the seminiferous tubules of the testis. This process starts with the mitotic division of the stem cells located close to the basement membrane of the tubu ...
(formation of male gametes). One centimorgan corresponds to about 1 million base pairs in humans on average. The relationship is only rough, as the physical chromosomal distance corresponding to one centimorgan varies from place to place in the genome, and also varies between men and women since recombination during gamete formation in females is significantly more frequent than in males. Kong et al. calculated that the female genome is 4460 cM long, while the male genome is only 2590 cM long. ''
Plasmodium falciparum ''Plasmodium falciparum'' is a unicellular protozoan parasite of humans, and the deadliest species of '' Plasmodium'' that causes malaria in humans. The parasite is transmitted through the bite of a female '' Anopheles'' mosquito and causes the ...
'' has an average recombination distance of ~15 kb per centimorgan: markers separated by 15 kb of DNA (15,000 nucleotides) have an expected rate of chromosomal crossovers of 0.01 per generation. Note that non- syntenic genes (genes residing on different chromosomes) are inherently unlinked, and cM distances are not applicable to them.


Relation to the probability of recombination

Because
genetic recombination Genetic recombination (also known as genetic reshuffling) is the exchange of genetic material between different organisms which leads to production of offspring with combinations of traits that differ from those found in either parent. In eukaryo ...
between two markers is detected only if there are an odd number of chromosomal crossovers between the two markers, the distance in centimorgans does not correspond exactly to the probability of genetic recombination. Assuming
J. B. S. Haldane John Burdon Sanderson Haldane (; 5 November 18921 December 1964), nicknamed "Jack" or "JBS", was a British-Indian scientist who worked in physiology, genetics, evolutionary biology, and mathematics. With innovative use of statistics in biolog ...
's map function, in which the number of chromosomal crossovers is distributed according to a
Poisson distribution In probability theory and statistics, the Poisson distribution is a discrete probability distribution that expresses the probability of a given number of events occurring in a fixed interval of time or space if these events occur with a known c ...
, a genetic distance of ''d'' centimorgans will lead to an odd number of chromosomal crossovers, and hence a detectable genetic recombination, with probability :\ P(\text, \textd\text) = \sum_^ \ P(2k + 1 \text, \textd\text) : = \sum_^ e^ \frac = e^ \sinh(d/100) = \frac\,, where sinh is the
hyperbolic sine In mathematics, hyperbolic functions are analogues of the ordinary trigonometric functions, but defined using the hyperbola rather than the circle. Just as the points form a circle with a unit radius, the points form the right half of the ...
function. The probability of recombination is approximately ''d''/100 for small values of ''d'' and approaches 50% as ''d'' goes to infinity. The formula can be inverted, giving the distance in centimorgans as a function of the recombination probability: :d=50 \ln\left(\right)\,.


Etymology

The centimorgan was named in honor of geneticist
Thomas Hunt Morgan Thomas Hunt Morgan (September 25, 1866 – December 4, 1945) was an American evolutionary biologist, geneticist, embryologist, and science author who won the Nobel Prize in Physiology or Medicine in 1933 for discoveries elucidating the role th ...
by
J. B. S. Haldane John Burdon Sanderson Haldane (; 5 November 18921 December 1964), nicknamed "Jack" or "JBS", was a British-Indian scientist who worked in physiology, genetics, evolutionary biology, and mathematics. With innovative use of statistics in biolog ...
. However, its parent unit, the morgan, is rarely used today.


See also

*
Mutation rate In genetics, the mutation rate is the frequency of new mutations in a single gene or organism over time. Mutation rates are not constant and are not limited to a single type of mutation; there are many different types of mutations. Mutation rates ...


References


Further reading

* {{refend Units of measurement Genetics