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Non- homologous end joining - Wikipedia. Non- homologous end joining (NHEJ) and homologous recombination (HR) in mammals during DNA double- strand break. Non- homologous end joining (NHEJ) is a pathway that repairs double- strand breaks in DNA. NHEJ is referred to as . These microhomologies are often present in single- stranded overhangs on the ends of double- strand breaks. When the overhangs are perfectly compatible, NHEJ usually repairs the break accurately.

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Inappropriate NHEJ can lead to translocations and telomere fusion, hallmarks of tumor cells. In this pathway, end resection reveals short microhomologies on either side of the break, which are then aligned to guide repair. Repair by MMEJ therefore leads to deletion of the DNA sequence between the microhomologies. In bacteria. NHEJ proteins have been identified in a number of bacteria, however, including Bacillus subtilis, Mycobacterium tuberculosis, and Mycobacterium smegmatis. Ku is a basket- shaped molecule that slides onto the DNA end and translocates inward. Ku may function as a docking site for other NHEJ proteins, and is known to interact with the DNA ligase IV complex and XLF.

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This step is not necessary if the ends are already compatible and have 3' hydroxyl and 5' phosphate termini. Little is known about the function of nucleases in NHEJ. Artemis is required for opening the hairpins that are formed on DNA ends during V(D)J recombination, a specific type of NHEJ, and may also participate in end trimming during general NHEJ. In this step, the 5' strand of the break is degraded by nucleases to create long 3' single- stranded tails. DSBs that have not been resected can be rejoined by NHEJ, but resection of even a few nucleotides strongly inhibits NHEJ and effectively commits the break to repair by recombination. This regulation is accomplished by the cyclin- dependent kinase. Cdk. 1 (Cdc. 28 in yeast), which is turned off in G1 and expressed in S and G2.

Cdk. 1 phosphorylates the nuclease Sae. Unlike typical cellular NHEJ, in which accurate repair is the most favorable outcome, error- prone repair in V(D)J recombination is beneficial in that it maximizes diversity in the coding sequence of these genes. Patients with mutations in NHEJ genes are unable to produce functional B cells and T cells and suffer from severe combined immunodeficiency (SCID).

At telomeres. Loss of capping proteins causes telomere shortening and inappropriate joining by NHEJ, producing dicentric chromosomes which are then pulled apart during mitosis. Paradoxically, some NHEJ proteins are involved in telomere capping. For example, Ku localizes to telomeres and its deletion leads to shortened telomeres. These syndromes share many features including cellular radiosensitivity, microcephaly and severe combined immunodeficiency (SCID) due to defective V(D)J recombination. Loss- of- function mutations in Artemis also cause SCID, but these patients do not show the neurological defects associated with LIG4 or XLF mutations. The difference in severity may be explained by the roles of the mutated proteins.

Artemis is a nuclease and is thought to be required only for repair of DSBs with damaged ends, whereas DNA Ligase IV and XLF are required for all NHEJ events. Many NHEJ genes have been knocked out in mice. Deletion of XRCC4 or LIG4 causes embryonic lethality in mice, indicating that NHEJ is essential for viability in mammals. In contrast, mice lacking Ku or DNA- PKcs are viable, probably because low levels of end joining can still occur in the absence of these components.

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Efficiency was higher in the skin, lung and kidney fibroblasts, and lower in heart fibroblasts and brain astrocytes. Furthermore, NHEJ efficiency declined with age.

The decline was 1. Reduced capability for NHEJ can lead to an increase in the number of unrepaired or faultily repaired DNA double- strand breaks that may then contribute to aging. Molecular and Cellular Biology. Evidence for a DNA polymerase beta (Pol.

Promotion of Dnl. DNA end- joining by the Rad. Mre. 11/Xrs. 2 and Hdf. Hdf. 2 complexes. PMID 1. 17. 41. 54. Zha S, Boboila C, Alt FW (August 2.

Hairpin opening and overhang processing by an Artemis/DNA- dependent protein kinase complex in nonhomologous end joining and V(D)J recombination. PMID 1. 19. 55. 43. Nick Mc. Elhinny SA, Ramsden DA (August 2. E.; Grawunder U.; Lieber M.

Cancer and aging as consequences of un- repaired DNA damage. In: New Research on DNA Damages (Editors: Honoka Kimura and Aoi Suzuki) Nova Science Publishers, Inc., New York, Chapter 1, pp.