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a caused by external environmental agents including uv radiation ionizing radiation and chemical toxins citation needed backbone damage edit ionizing radiation may produce highly reactive free radicals that can break the bonds in the dna double stranded breakages are especially damaging and hard to repair producing translocation and deletion of part of a chromosome alkylating agents like mustard gas diet tobacco smoke may also cause breakages in the dna backbone endogenous processes may also such as oxidative stress may also generate highly reactive oxygen species that can damage the dna citation needed crosslinking edit main article crosslinking of dna covalent bonds between the bases of nucleotides in dna be they in the same strand or opposing strands is referred to as crosslinking of dna crosslinking of dna may affect both the replication and the transcription of dna and it may be caused by exposure to a variety of agents some naturally occurring chemicals may also promote crosslinking such as psoralens after activation by uv radiation and nitrous acid interstrand cross linking between two strands causes more damage as it blocks replication and transcription and can cause chromosomal breakages and rearrangements some crosslinkers such as cyclophosphamide mitomycin c and cisplatin are used as anticancer chemotherapeutic because of their high degree of toxicity to proliferating cells citation needed dimerization edit main article dimer dimerization consists of the bonding of two monomers to form an oligomer such as the formation of pyrimidine dimers as a result of exposure to uv radiation which promotes the formation of a cyclobutyl ring between adjacent thymines in dna 37 these bulky bases would cause a distortion in the dna helixes and can interfere with dna replication and transcription in human skin cells thousands of dimers may be formed in a day due to normal exposure to sunlight dna polymerase η may help bypass these lesions in an error free manner 38 however individuals with defective dna repair function such as those with xeroderma pigmentosum are sensitive to sunlight and may be prone to skin cancer ethidium intercalated between two adenine thymine base pairs clinically whether a tumor has formed as a direct consequence of uv radiation is discernible via dna sequencing analysis for the characteristic context specific dimerization pattern that occurs due to excessive exposure to sunlight 39 intercalation between bases edit main article intercalation biochemistry the planar structure of chemicals such as ethidium bromide and proflavine allows them to insert between bases in dna this insert causes the dna s backbone to stretch and makes slippage in dna during replication more likely to occur since the bonding between the strands is made less stable by the stretching forward slippage will result in deletion mutation while reverse slippage will result in an insertion mutation also the intercalation into dna of anthracyclines such as daunorubicin and doxorubicin interferes with the functioning of the enzyme topoisomerase ii blocking replication as well as causing mitotic homologous recombination citation needed insertional mutagenesis edit main article insertional mutagenesis transposons and viruses or retrotransposons may insert dna sequences into coding regions or functional elements of a gene and result in inactivation of the gene 40 dna repair pathway edit dna damage happens frequently but dna damage does not always become a mutation only after the repair mechanism has failed or inaccurate allowing the damages to bypass then the mutation would happen and potentially lead to diseases like cancer citation needed base excision repair edit base excision repair corrects the small non helix distorting lesions of dna helix such as oxidative deaminated alkylation as well as basic single abase damages 30 dna glycosylase within the base excision repair mechanism recognizes those damages and cleaves the n glycosidic bond leave behind an abasic sites dna backbone would then be cut at that site by ap endonuclease after that dna polymerase fills the gap nucleotide excision repair edit nucleotide excision repair removes bulky lesion such as cpds clarification needed and 6 4 pp from uv radiation or damage from chemotherapeutic agents there are two major branches of nucleotide excision repair globular genome nucleotide excision repair and transcription coupled nucleotide excision repair globular genome ner the xpc rad23b and cetn2 protein complex scans for whole genome damage once the damage is found endonuclease such as xpf ercc1 and xpg will cut out the lesion from 5 to 3 and pol ε or xrcc1 lig3 will carry out the gap filling synthesis and ligation 30 mismatch repair edit mismatch repair removes base mismatch that have arisen during replication and the insertion deletion loop 41 humans employ the mutsα heterodimer msh2 msh6 to recognize the base mismatch 42 once the mismatch is found exo1 carries out the 5 directed mismatch excision which creates a gap later being filled by polδ rfc and hmgb 43 adaptive mutagenesis mechanisms edit main article adaptive mutation adaptive mutagenesis has been defined as mutagenesis mechanisms that enable an organism to adapt to an environmental stress since the variety of environmental stresses is very broad the mechanisms that enable it are also quite broad as far as research on the field has shown for instance in bacteria while modulation of the sos response and endogenous prophage dna synthesis has been shown to increase acinetobacter baumannii resistance to ciprofloxacin 16 resistance mechanisms are presumed to be linked to chromosomal mutation untransferable via horizontal gene transfer in some members of family enterobacteriaceae such as e coli salmonella spp klebsiella spp and enterobacter spp 44 chromosomal events specially gene amplification seem also to be relevant to this adaptive mutagenesis in bacteria 45 research in eukaryotic cells is much scarcer but chromosomal events seem also to be rather relevant while an ectopic intrachromosomal recombination has been reported to be involved in acquisition of resistance to 5 fluorocytosine in saccharomyces cerevisiae 17 genome duplications have been found to confer resistance in s cerevisiae to nutrient poor environments 21 46 47 laboratory applications edit main article mutagenesis molecular biology technique in the laboratory mutagenesis is a technique by which dna mutations are deliberately engineered to produce mutant genes proteins or strains of organisms various constituents of a gene such as its control elements and its gene product may be mutated so that the function of a gene or protein can be examined in detail the mutation may also produce mutant proteins with altered properties or enhanced or novel functions that may prove to be of use commercially mutant strains of organisms that have practical applications or allow the molecular basis of particular cell function to be investigated may also be produced citation needed early methods of mutagenesis produced entirely random mutations however modern methods of mutagenesis are capable of producing site specific mutations modern laboratory techniques used to generate these mutations include directed mutagenesis site directed mutagenesis pcr mutagenesis insertional mutagenesis signature tagged mutagenesis transposon mutagenesis sequence saturation mutagenesis see also edit carcinogenesis dna damage naturally occurring dna repair dysgenics mutagen mutation mutation breeding mutation rate transfection references edit beale g 1993 the discovery of mustard gas mutagenesis by auerbach and robson in 1941 genetics 134 2 393 399 doi 10 1093 genetics 134 2 393 pmc 1205483 pmid 8325476 kevin m gleason published 2017 03 07 hermann joseph muller s study of x rays as a mutagen 1926 1927 cite web cs1 maint numeric names authors list link genetics and genomics timeline 1927 hermann j muller 1890 1967 demonstrates 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