220:. The structure of telomeres is highly conserve and is organized in multiple short tandem DNA repeats. Telomeres and DSBs have different functionality, such that telomeres prevent DNA repair activities. During telomeric DNA replication in the S/G2 and G1 phases of the cell cycle, the 3' lagging strand leaves a short overhang called a G-tail. Telomeric DNA ends at the 3' G tail end because the 3' lagging strand extends without its complementary 5' C leading strand. The G tail provide a major function to telomeric DNA such that the G tails control telomere homeostasis.
240:. Such process in S. Cerevisiae for example is negatively regulated by this activity. The MRX complex and the Ku complex bind simultaneously and independently to DSBs ends. In the presence of the telomere-associated proteins, MRX fails to bind to the DSB ends while the Ku complex binds to DSB ends. The bound Ku complex to the DSB ends protect the telomeres from nucleolytic degradation by
50:
342:
to the HR pathway. Cyclin-dependent protein kinase such as cdk1 serve as a positive regulator of the HR pathway. This positive regulator promotes 5′–3′ nucleolytic degradation of DNA ends. Along with cdk1, the MRX complex, B1 cyclin, and Spo11-induced DSBs serve as a positive regulators to the HR pathway.
332:
Resection ensures that DSBs are not repaired by NHEJ (which joins broken DNA ends together without ensuring that they match), but rather by methods based on homology (matching DNA sequences). Cyclin-dependent protein kinase such as cdk1 in yeast serves as a negative regulator of the NHEJ pathway. Any
297:
enzyme, Sgs1 enzyme, and the nucleases Exo1 and Dna2. Involvement of Sae2 Sar267 in DSB processing is highly conserved throughout eukaryotes, such that the Sae2 along with the MRX complex are involved in two major functions: single-strand annealing, and processing of hairpin DNA structures. Like all
341:
The presence of a ssDNA allows the broken end of the DNA to line up accurately with a matching sequence, so that it can be accurately repaired. For HR pathway to occur in the S and G2 phases of the cell cycle, availability of a sister chromatid is required. 5′–3′ resection automatically links a DSB
319:
The pathway of choice in DNA repair is highly regulated to guarantee that cells in the S/G2 and G1 phase use the appropriate mechanism. Regulators in both the NHEJ and HR pathway mediate the appropriate DNA repair response pathway. Furthermore, recent studies into DNA repair show that regulation of
310:
filament which can take part in the search for a matching region, allowing HR to take place. The 3' ssDNA coated by a RPA promotes the recruitment of Mec1. Mec1 further phosphorylates Sae2 along with cdk1. The resulting phosphorylation by Sae2 by Mec1 helps increase the effect of resection and this
142:
for activation, this event only happens in the G2 and S phases of the cell cycle during replication. DSBs that have not begun DNA end resection can be ligated by NHEJ pathway, but resection of a few nucleotides inhibits the NHEJ pathway and commits' DNA repair by the HR pathway. The NHEJ pathway is
264:
parts of the resection machinery. This process alleviates the inhibitory effect of the telomere-associated proteins, and allows Cdc13 (a binding protein on both the lagging strand, and leading strand) to cover telomeric DNA. The binding of cdc13 to DNA suppresses DNA damage checkpoint and allows
328:
DNA end resection is key in determining the correct pathway in NHEJ. For NHEJ pathway to occur, positive regulators such as the Ku and MRX complex mediate recruitment of other NHEJ-associated proteins such as Tel1, Lif1, Dnl4, and Nej1. Since NHEJ does not rely on end resection, NHEJ could only
273:
One of the important regulatory controls in mitotic cells is deciding which specific DSB repair pathway to take. Once a DSB is detected, the highly conserved complexes are recruited by the DNA ends. If the cell is in the G1 phase of the cell cycle, the complex Ku prevents resection to occur and
68:
recombination. Furthermore, the natural ends of the linear chromosomes resemble DSBs, and although DNA breaks can cause damage to the integrity of genomic DNA, the natural ends are packed into complex specialized DNA protective packages called
121:
Accurate repair of DSBs are essential in the upkeep of genome integrity. From the three mechanisms that exists to repair DSBs, NHEJ and HR repair mechanisms are the dominant pathways. Several highly conservative proteins trigger the
93:. Furthermore, DSBs can lead to genome rearrangements and instability. Cases where two complementary strands are linked at the point of the DSB have potential to be catastrophic, such that the cell will not be able to complete
126:
for detection of DSBs ensuing repair by either NHEJ or HR repair pathways. NHEJ mechanism functions in ligating two different DSBs with high fidelity, while HR relies on a homologous template to repair DSB ends.
252:
In the late S/G2 phase of the cell cycle, the telomere-associated proteins RIF1, RIF2, and RAP2 exhibit their inhibitory effect by binding to telomeric DNA. In the Late S/G2 phase, the protein kinase
293:
Ser267. After phosphorylation occurs by Cdk1, MRX complex binds to dsDNA ends and generates short ssDNA that stretches in the 5' direction. The 5' ssDNA continues resection by the activity of the
196:-interacting protein (CtIP) needs to bind to the MRN complex so that the first phase of resection can begin, namely short-range end resection. After
73:
that prevent DNA repair activities. Telomeres and mitotic DSBs have different functionality, but both experience the same 5′–3′ degradation process.
777:"The isolation and partial characterization of age-correlated oligo-deoxyribo-ribonucleotides with covalently linked aspartyl-glutamyl polypeptides"
244:. This results in an inhibition of telomerase elongation at the DSB ends and prevents further telomere action at the G1 phase of the cell cycle.
1354:
1318:
147:
phase. In G1 phase there is no sister chromatids to repair DSBs via the HR pathway making the NHEJ pathway a critical repair mechanism.
1116:
396:
106:
39:
421:
Jimeno S, Mejías-Navarro F, Prados-Carvajal R, Huertas P (2019). "Controlling the balance between chromosome break repair pathways".
1223:
442:
229:
46:(ssDNA) allows the broken end of the DNA to line up accurately with a matching sequence, so that it can be accurately repaired.
282:
and the Nej1/XLF protein. This process results in error-prone religation of DSB ends at the G1 phase of the cell cycle.
386:
366:
102:
274:
triggers the NHEJ pathway factors. DSBs in the NHEJ pathway are ligated, a step in the NHEJ pathway that requires
286:
253:
167:
329:
happen in the G1 phase of the cell cycle. Both Ku and NHEJ-associated proteins prevent initiation of resection.
391:
110:
1206:
Pinto C, Anand R, Cejka P (2018). "Methods to Study DNA End
Resection II: Biochemical Reconstitution Assays".
97:
when it next divides, and will either die or, in rare cases, undergo chromosomal loss, duplications, and even
49:
261:
257:
197:
732:
Bjorksten J, Acharya PV, Ashman S, Wetlaufer DB (July 1971). "Gerogenic fractions in the tritiated rat".
64:
causing DNA end resection and repair activities to take place, but they are also normal intermediates in
361:
299:
123:
27:
150:
Before resection can take place, the break needs to be detected. In animals, this detection is done by
85:
is a kind of DNA damage in which both strands in the double helix are severed. DSBs only occur during
1263:
1017:"PARP2 Is the Predominant Poly(ADP-Ribose) Polymerase in Arabidopsis DNA Damage and Immune Responses"
57:
1208:
Mechanisms of DNA Recombination and Genome
Rearrangements: Methods to Study Homologous Recombination
1376:
333:
activity associated with the presence of cyclin dependent protein kinases inhibit the NHEJ pathway
43:
1068:"PARP1-dependent kinetics of recruitment of MRE11 and NBS1 proteins to multiple DNA damage sites"
757:
456:
1350:
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997:
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448:
438:
139:
1066:
Haince JF, McDonald D, Rodrigue A, Déry U, Masson JY, Hendzel MJ, Poirier GG (January 2008).
1281:
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1038:
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987:
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502:
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170:
86:
1108:
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Casari E, Rinaldi C, Marsella A, Gnugnoli M, Colombo CV, Bonetti D, Longhese MP (2019).
1286:
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1016:
992:
967:
831:
806:
745:
709:
684:
627:
602:
507:
482:
216:
Linear chromosomes are packed into complex specialized DNA protective packages called
1370:
927:"DNA joint dependence of pol X family polymerase action in nonhomologous end joining"
320:
DNA end resection is governed by the activity of cdk1 in the cell replication cycle.
307:
241:
761:
460:
290:
201:
1150:"Processing of DNA Double-Strand Breaks by the MRX Complex in a Chromatin Context"
1033:
700:
434:
351:
279:
237:
205:
186:
163:
143:
involved throughout the cell cycle, but it is critical to DNA repair during the
1256:
Proceedings of the
National Academy of Sciences of the United States of America
1250:
Xue C, Wang W, Crickard JB, Moevus CJ, Kwon Y, Sung P, Greene EC (March 2019).
1215:
1345:
Poon RY (2016-01-01). "Mitotic
Catastrophe". In Bradshaw RA, Stahl PD (eds.).
498:
376:
371:
356:
275:
90:
82:
61:
35:
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resection to occur while allowing for telomerase elongation at the DSB ends.
1276:
256:(cyclin-dependent) promotes telomeric resection. This control is exerted by
155:
1295:
1233:
1185:
1093:
1084:
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1052:
1001:
952:
943:
926:
881:
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855:
840:
718:
636:
618:
516:
452:
788:
776:
753:
1252:"Regulatory control of Sgs1 and Dna2 during eukaryotic DNA end resection"
968:"The multifaceted roles of PARP1 in DNA repair and chromatin remodelling"
381:
294:
217:
144:
135:
98:
70:
983:
854:
Liang L, Deng L, Chen Y, Li GC, Shao C, Tischfield JA (September 2005).
130:
DNA end resection in the HR pathway only occurs at two specific phases:
895:
Watson JD, Baker TA, Bell SP, Gann A, Levine M, Losick R, eds. (2004).
204:, probably about 300 base pairs from the end, and then acts as a 3'→5'
131:
94:
65:
42:
to produce a 3' single-stranded sequence. The presence of a section of
822:
200:
CtIP binds, the Mre11 subunit is able to cut the 5'-terminated strand
53:
Mechanism of regulation of 5' resection of
Mitotic and Telomeric DSBs.
1113:
Atlas of
Genetics and Cytogenetics in Oncology and Haematology - NBS1
182:
174:
285:
If the cells are in S/G2 phase, mitotic DSBs are controlled through
228:
In the G1 phase of the cell cycle, the telomere-associated proteins
899:(5th ed.). Pearson Benjamin Cummings; CSHL Press. Ch. 9, 10.
303:
233:
193:
178:
159:
151:
601:
Longhese MP, Bonetti D, Manfrini N, Clerici M (September 2010).
185:
in mammals, or Xrs2 in yeast, where this complex is called the
31:
298:
ssDNA in the nucleus, the resected region is first coated by
807:"DNA resection in eukaryotes: deciding how to fix the break"
113:
HR. Of these, only NHEJ does not rely on DNA end resection.
1210:. Methods in Enzymology. Vol. 600. pp. 67–106.
162:
seems to play this role. PARP binding then recruits the
311:
in turn leads to the DNA damage checkpoint activation.
925:
Daley JM, Laan RL, Suresh A, Wilson TE (August 2005).
423:
Advances in
Protein Chemistry and Structural Biology
856:"Modulation of DNA end joining by nuclear proteins"
685:"DNA end resection: many nucleases make light work"
1015:Song J, Keppler BD, Wise RR, Bent AF (May 2015).
603:"Mechanisms and regulation of DNA end resection"
236:bind to telomeric DNA and prevent access to the
966:Ray Chaudhuri A, Nussenzweig A (October 2017).
1107:Uhrhammer N, Bay JO, Gatti RA (October 2002).
1349:. Waltham: Academic Press. pp. 399–403.
302:(RPA) complex, but RPA is then replaced with
8:
683:Mimitou EP, Symington LS (September 2009).
734:Journal of the American Geriatrics Society
1285:
1275:
1175:
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1042:
1032:
991:
942:
871:
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811:Nature Structural & Molecular Biology
781:Johns Hopkins Medical Journal. Supplement
708:
626:
506:
487:Genomics, Proteomics & Bioinformatics
483:"DNA End Resection: Facts and Mechanisms"
289:activity and involves phosphorylation of
101:. Three mechanisms exist to repair DSBs:
34:(dsDNA) is modified by cutting away some
208:to strip away the end of the 5' strand.
48:
408:
166:to the breakage site. This is a highly
972:Nature Reviews. Molecular Cell Biology
1340:
1338:
1311:New research directions in DNA repair
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1201:
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60:(DSBs) can occur at any phase of the
26:, is a biochemical process where the
7:
1109:"NBN (Nijmegen breakage syndrome 1)"
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1072:The Journal of Biological Chemistry
931:The Journal of Biological Chemistry
860:The Journal of Biological Chemistry
1154:Frontiers in Molecular Biosciences
746:10.1111/j.1532-5415.1971.tb02577.x
397:Microhomology-mediated end joining
138:phases. Since HR pathway requires
107:microhomology-mediated end joining
14:
154:; similar systems exist in other
248:Telomeres in the late S/G2 phase
30:of a section of double-stranded
1119:from the original on 2006-09-29
1:
897:Molecular Biology of the Gene
1347:Encyclopedia of Cell Biology
1034:10.1371/journal.pgen.1005200
701:10.1016/j.dnarep.2009.04.017
481:Liu T, Huang J (June 2016).
435:10.1016/bs.apcsb.2018.10.004
278:activity of Dnl4-Lif1/XRCC4
192:Before resection can start,
1393:
1216:10.1016/bs.mie.2017.11.009
805:Huertas P (January 2010).
367:Non-homologous end joining
212:Resection of telomere DSBs
103:non-homologous end joining
499:10.1016/j.gpb.2016.05.002
269:Resection of mitotic DSBs
1167:10.3389/fmolb.2019.00043
392:Homologous Recombination
258:cyclin-dependent kinases
111:homologous recombination
1277:10.1073/pnas.1819276116
1085:10.1074/jbc.M706734200
944:10.1074/jbc.M505277200
873:10.1074/jbc.M503776200
619:10.1038/emboj.2010.165
54:
300:Replication protein A
224:Telomeres in G1 phase
124:DNA Damage Checkpoint
52:
1309:Chen C, ed. (2013).
429:. Elsevier: 95–134.
357:Double-strand breaks
58:Double-strand breaks
1313:. Croatia: InTech.
1268:2019PNAS..116.6091X
984:10.1038/nrm.2017.53
937:(32): 29030–29037.
866:(36): 31442–31449.
775:Acharya PV (1972).
337:Positive regulators
202:endonucleolytically
181:and NBS1 (known as
83:double-strand break
44:single-stranded DNA
16:Biochemical process
55:
1356:978-0-12-394796-3
1320:978-953-51-1114-6
1262:(13): 6091–6100.
823:10.1038/nsmb.1710
613:(17): 2864–2874.
140:sister chromatids
24:5′–3′ degradation
20:DNA end resection
1384:
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1087:
1078:(2): 1197–1208.
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607:The EMBO Journal
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1027:(5): e1005200.
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978:(10): 610–621.
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87:DNA replication
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783:(1): 254–260.
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740:(7): 561–574.
724:
695:(9): 983–995.
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522:
493:(3): 126–130.
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198:phosphorylated
173:consisting of
118:
115:
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22:, also called
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10:
9:
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4:
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2:
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262:phosphorylate
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158:: in plants,
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1121:. Retrieved
1112:
1102:
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1024:
1020:
1010:
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817:(1): 11–16.
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733:
727:
692:
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610:
606:
490:
486:
426:
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340:
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324:NHEJ pathway
318:
284:
272:
251:
232:, RIF2, and
227:
215:
191:
149:
129:
120:
109:(MMEJ), and
80:
56:
23:
19:
18:
352:Exonuclease
280:heterodimer
238:MRX complex
206:exonuclease
187:MRX complex
164:MRN complex
36:nucleotides
1377:DNA repair
1123:2008-02-12
689:DNA Repair
403:References
377:Cell cycle
372:Nucleotide
362:Blunt ends
315:Regulators
306:to form a
276:DNA ligase
156:eukaryotes
91:cell cycle
77:Background
62:cell cycle
1329:957280914
905:936762772
218:telomeres
168:conserved
117:Mechanism
99:mutations
71:telomeres
38:from the
28:blunt end
1371:Category
1296:30850524
1234:29458776
1186:31231660
1117:Archived
1094:18025084
1053:25950582
1002:28676700
953:15964833
882:16012167
841:20051983
762:33154242
719:19473888
637:20647996
517:27240470
461:73459973
453:30798939
382:Telomere
346:See also
295:helicase
260:, which
105:(NHEJ),
1287:6442620
1264:Bibcode
1177:6567933
1044:4423837
993:6591728
832:2850169
789:5055816
754:5106728
710:2760233
628:2944052
508:4936662
171:complex
95:mitosis
89:of the
66:mitosis
1353:
1327:
1317:
1294:
1284:
1232:
1222:
1184:
1174:
1160:: 43.
1092:
1051:
1041:
1000:
990:
951:
903:
880:
839:
829:
787:
760:
752:
717:
707:
635:
625:
515:
505:
459:
451:
441:
183:Nibrin
40:5' end
758:S2CID
457:S2CID
304:RAD51
194:CtBP1
179:Rad50
175:Mre11
160:PARP2
152:PARP1
1351:ISBN
1325:OCLC
1315:ISBN
1292:PMID
1230:PMID
1220:ISBN
1182:PMID
1090:PMID
1049:PMID
998:PMID
949:PMID
901:OCLC
878:PMID
837:PMID
785:PMID
750:PMID
715:PMID
633:PMID
513:PMID
449:PMID
439:ISBN
387:NHEJ
291:Sae2
287:Cdk1
254:CDK1
242:exo1
234:RAP2
230:RIF1
134:and
1282:PMC
1272:doi
1260:116
1212:doi
1172:PMC
1162:doi
1080:doi
1076:283
1039:PMC
1029:doi
988:PMC
980:doi
939:doi
935:280
868:doi
864:280
827:PMC
819:doi
742:doi
705:PMC
697:doi
623:PMC
615:doi
503:PMC
495:doi
431:doi
427:115
189:).
32:DNA
1373::
1337:^
1323:.
1290:.
1280:.
1270:.
1258:.
1254:.
1242:^
1228:.
1218:.
1194:^
1180:.
1170:.
1156:.
1152:.
1132:^
1115:.
1111:.
1088:.
1074:.
1070:.
1047:.
1037:.
1025:11
1023:.
1019:.
996:.
986:.
976:18
974:.
970:.
947:.
933:.
929:.
913:^
876:.
862:.
858:.
835:.
825:.
815:17
813:.
809:.
797:^
779:.
756:.
748:.
738:19
736:.
713:.
703:.
691:.
687:.
645:^
631:.
621:.
611:29
609:.
605:.
525:^
511:.
501:.
491:14
489:.
485:.
469:^
455:.
447:.
437:.
425:.
411:^
177:,
145:G1
136:G2
81:A
1359:.
1331:.
1298:.
1274::
1266::
1236:.
1214::
1188:.
1164::
1158:6
1126:.
1096:.
1082::
1055:.
1031::
1004:.
982::
955:.
941::
907:.
884:.
870::
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821::
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764:.
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721:.
699::
693:8
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132:S
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