The Catalytic Subunit of DNA-Dependent Protein Kinase Regulates Proliferation, Telomere Length, and Genomic Stability in Human Somatic Cells

REFERENCES

• Abdel-Rahman, W. M., K. Katsura, W. Rens, P. A. Gorman, D. Sheer, D. Bicknell, W. F. Bodmer, M. J. Arends, A. H. Wyllie, and P. A. Edwards. 2001. Spectral karyotyping suggests additional subsets of colorectal cancers characterized by pattern of chromosome rearrangement. Proc. Natl. Acad. Sci. USA 98:2538–2543.
   PubMedGoogle Scholar
• Allalunis-Turner, J., G. M. Barron, and R. S. Day III. 1997. Intact G2-phase checkpoint in cells of a human cell line lacking DNA-dependent protein kinase activity. Radiat. Res. 147:284–287.
   PubMed Web of Science ®Google Scholar
• Allalunis-Turner, M. J., G. M. Barron, R. S. Day III, K. D. Dobler, and R. Mirzayans. 1993. Isolation of two cell lines from a human malignant glioma specimen differing in sensitivity to radiation and chemotherapeutic drugs. Radiat. Res. 134:349–354.
   PubMedGoogle Scholar
• Anderson, C. W., and M. J. Allalunis-Turner. 2000. Human TP53 from the malignant glioma-derived cell lines M059J and M059K has a cancer-associated mutation in exon 8. Radiat. Res. 154:473–476.
   PubMed Web of Science ®Google Scholar
• Anderson, C. W., J. J. Dunn, P. I. Freimuth, A. M. Galloway, and M. J. Allalunis-Turner. 2001. Frameshift mutation in PRKDC, the gene for DNA-PKcs, in the DNA repair-defective, human, glioma-derived cell line M059J. Radiat. Res. 156:2–9.
   PubMed Web of Science ®Google Scholar
• Bailey, S. M., M. A. Brenneman, J. Halbrook, J. A. Nickoloff, R. L. Ullrich, and E. H. Goodwin. 2004. The kinase activity of DNA-PK is required to protect mammalian telomeres. DNA Repair 3:225–233.
   PubMedGoogle Scholar
• Bertuch, A. A., and V. Lundblad. 2003. The Ku heterodimer performs separable activities at double-strand breaks and chromosome termini. Mol. Cell. Biol. 23:8202–8215.
   PubMedGoogle Scholar
• Bogue, M. A., C. Jhappan, and D. B. Roth. 1998. Analysis of variable (diversity) joining recombination in DNA-dependent protein kinase (DNA-PK)-deficient mice reveals DNA-PK-independent pathways for both signal and coding joint formation. Proc. Natl. Acad. Sci. USA 95:15559–15564.
   PubMed Web of Science ®Google Scholar
• Bosma, G. C., R. P. Custer, and M. J. Bosma. 1983. A severe combined immunodeficiency mutation in the mouse. Nature 301:527–530.
   PubMed Web of Science ®Google Scholar
• Buck, D., L. Malivert, R. de Chasseval, A. Barraud, M. C. Fondaneche, O. Sanal, A. Plebani, J. L. Stephan, M. Hufnagel, F. le Deist, A. Fischer, A. Durandy, J. P. de Villartay, and P. Revy. 2006. Cernunnos, a novel nonhomologous end-joining factor, is mutated in human immunodeficiency with microcephaly. Cell 124:287–299.
   PubMed Web of Science ®Google Scholar
• Chamberlain, J. R., U. Schwarze, P. R. Wang, R. K. Hirata, K. D. Hankenson, J. M. Pace, R. A. Underwood, K. M. Song, M. Sussman, P. H. Byers, and D. W. Russell. 2004. Gene targeting in stem cells from individuals with osteogenesis imperfecta. Science 303:1198–1201.
   PubMed Web of Science ®Google Scholar
• Collis, S. J., T. L. DeWeese, P. A. Jeggo, and A. R. Parker. 2005. The life and death of DNA-PK. Oncogene 24:949–961.
   PubMed Web of Science ®Google Scholar
• Darroudi, F., and A. T. Natarajan. 1987. Cytological characterization of Chinese hamster ovary X-ray-sensitive mutant cells xrs 5 and xrs 6. I. Induction of chromosomal aberrations by X-irradiation and its modulation with 3-aminobenzamide and caffeine. Mutat. Res. 177:133–148.
   PubMed Web of Science ®Google Scholar
• Ding, Q., L. Bramble, V. Yuzbasiyan-Gurkan, T. Bell, and K. Meek. 2002. DNA-PKcs mutations in dogs and horses: allele frequency and association with neoplasia. Gene 283:263–269.
   PubMedGoogle Scholar
• Espejel, S., S. Franco, A. Sgura, D. Gae, S. M. Bailey, G. E. Taccioli, and M. A. Blasco. 2002. Functional interaction between DNA-PKcs and telomerase in telomere length maintenance. EMBO J. 21:6275–6287.
   PubMed Web of Science ®Google Scholar
• Espejel, S., M. Martin, P. Klatt, J. Martin-Caballero, J. M. Flores, and M. A. Blasco. 2004. Shorter telomeres, accelerated ageing and increased lymphoma in DNA-PKcs-deficient mice. EMBO Rep. 5:503–509.
   PubMed Web of Science ®Google Scholar
• Fattah, F. J., N. F. Lichter, K. R. Fattah, S. Oh, and E. A. Hendrickson. 2008. Ku70, an essential gene, modulates the frequency of rAAV-mediated gene targeting in human somatic cells. Proc. Natl. Acad. Sci. USA 105:8703–8708.
   PubMed Web of Science ®Google Scholar
• Fattah, K. R., B. L. Ruis, and E. A. Hendrickson. 2008. Mutations to Ku reveal differences in human somatic cell lines. DNA Repair 7:762–774.
   PubMed Web of Science ®Google Scholar
• Ferreira, M. G., K. M. Miller, and J. P. Cooper. 2004. Indecent exposure: when telomeres become uncapped. Mol. Cell 13:7–18.
   PubMedGoogle Scholar
• Finnie, N. J., T. M. Gottlieb, T. Blunt, P. A. Jeggo, and S. P. Jackson. 1995. DNA-dependent protein kinase activity is absent in xrs-6 cells: implications for site-specific recombination and DNA double-strand break repair. Proc. Natl. Acad. Sci. USA 92:320–324.
   PubMed Web of Science ®Google Scholar
• Fujimori, A., R. Araki, R. Fukumura, T. Saito, M. Mori, K. Mita, K. Tatsumi, and M. Abe. 1997. The murine DNA-PKcs gene consists of 86 exons dispersed in more than 250 kb. Genomics 45:194–199.
   PubMedGoogle Scholar
• Fukumura, R., R. Araki, A. Fujimori, M. Mori, T. Saito, F. Watanabe, M. Sarashi, H. Itsukaichi, K. Eguchi-Kasai, K. Sato, K. Tatsumi, and M. Abe. 1998. Murine cell line SX9 bearing a mutation in the DNA-PKcs gene exhibits aberrant V(D)J recombination not only in the coding joint but also in the signal joint. J. Biol. Chem. 273:13058–13064.
   PubMedGoogle Scholar
• Gao, Y., J. Chaudhuri, C. Zhu, L. Davidson, D. T. Weaver, and F. W. Alt. 1998. A targeted DNA-PKcs-null mutation reveals DNA-PK-independent functions for Ku in V(D)J recombination. Immunity 9:367–376.
   PubMed Web of Science ®Google Scholar
• Gao, Y., Y. Sun, K. M. Frank, P. Dikkes, Y. Fujiwara, K. J. Seidl, J. M. Sekiguchi, G. A. Rathbun, W. Swat, J. Wang, R. T. Bronson, B. A. Malynn, M. Bryans, C. Zhu, J. Chaudhuri, L. Davidson, R. Ferrini, T. Stamato, S. H. Orkin, M. E. Greenberg, and F. W. Alt. 1998. A critical role for DNA end-joining proteins in both lymphogenesis and neurogenesis. Cell 95:891–902.
   PubMed Web of Science ®Google Scholar
• Gilley, D., H. Tanaka, M. P. Hande, A. Kurimasa, G. C. Li, M. Oshimura, and D. J. Chen. 2001. DNA-PKcs is critical for telomere capping. Proc. Natl. Acad. Sci. USA 98:15084–15088.
   PubMed Web of Science ®Google Scholar
• Goytisolo, F. A., E. Samper, S. Edmonson, G. E. Taccioli, and M. A. Blasco. 2001. The absence of the DNA-dependent protein kinase catalytic subunit in mice results in anaphase bridges and in increased telomeric fusions with normal telomere length and G-strand overhang. Mol. Cell. Biol. 21:3642–3651.
   PubMed Web of Science ®Google Scholar
• Gravel, S., M. Larrivee, P. Labrecque, and R. J. Wellinger. 1998. Yeast Ku as a regulator of chromosomal DNA end structure. Science 280:741–744.
   PubMedGoogle Scholar
• Han, Z., T. H. Carter, W. H. Reeves, J. H. Wyche, and E. A. Hendrickson. 1996. DNA-dependent protein kinase is a target of a CPP32-like apoptotic protease. J. Biol. Chem. 271:25035–25040.
   PubMed Web of Science ®Google Scholar
• Han, Z., C. Johnston, W. H. Reeves, T. Carter, J. H. Wyche, and E. A. Hendrickson. 1996. Characterization of a Ku86 variant protein that results in altered DNA binding and diminished DNA-dependent protein kinase activity. J. Biol. Chem. 271:14098–14104.
   PubMedGoogle Scholar
• Hande, P., P. Slijepcevic, A. Silver, S. Bouffler, P. van Buul, P. Bryant, and P. Lansdorp. 1999. Elongated telomeres in scid mice. Genomics 56:221–223.
   PubMedGoogle Scholar
• Hefferin, M. L., and A. E. Tomkinson. 2005. Mechanism of DNA double-strand break repair by non-homologous end joining. DNA Repair 4:639–648.
   PubMed Web of Science ®Google Scholar
• Hendrickson, E. A. 2008. Gene targeting in human somatic cells, p. 509-525. In P. M. Conn (ed.), Source book of models for biomedical research. Humana Press, Inc., Totowa, NJ.
   Google Scholar
• Hendrickson, E. A., J. L. Huffman, and J. A. Tainer. 2006. Structural aspects of Ku and the DNA-dependent protein kinase complex, p. 629-684. In W. Seide, Y. W. Kow, and P. Doetsch (ed.), DNA damage recognition. Taylor and Francis Group, New York, NY.
   Google Scholar
• Jaco, I., P. Munoz, and M. A. Blasco. 2004. Role of human Ku86 in telomere length maintenance and telomere capping. Cancer Res. 64:7271–7278.
   PubMed Web of Science ®Google Scholar
• Jhappan, C., H. C. Morse III, R. D. Fleischmann, M. M. Gottesman, and G. Merlino. 1997. DNA-PKcs: a T-cell tumour suppressor encoded at the mouse scid locus. Nat. Genet. 17:483–486.
   PubMedGoogle Scholar
• Kim, J. S., H. Crooks, T. Dracheva, T. G. Nishanian, B. Singh, J. Jen, and T. Waldman. 2002. Oncogenic beta-catenin is required for bone morphogenetic protein 4 expression in human cancer cells. Cancer Res. 62:2744–2748.
   PubMedGoogle Scholar
• Kim, J. S., C. Lee, A. Foxworth, and T. Waldman. 2004. B-Raf is dispensable for K-Ras-mediated oncogenesis in human cancer cells. Cancer Res. 64:1932–1937.
   PubMedGoogle Scholar
• Kirchgessner, C. U., C. K. Patil, J. W. Evans, C. A. Cuomo, L. M. Fried, T. Carter, M. A. Oettinger, and J. M. Brown. 1995. DNA-dependent kinase (p350) as a candidate gene for the murine SCID defect. Science 267:1178–1183.
   PubMed Web of Science ®Google Scholar
• Kohli, M., C. Rago, C. Lengauer, K. W. Kinzler, and B. Vogelstein. 2004. Facile methods for generating human somatic cell gene knockouts using recombinant adeno-associated viruses. Nucleic Acids Res. 32:e3.
   PubMedGoogle Scholar
• Lee, S. E., R. A. Mitchell, A. Cheng, and E. A. Hendrickson. 1997. Evidence for DNA-PK-dependent and -independent DNA double-strand break repair pathways in mammalian cells as a function of the cell cycle. Mol. Cell. Biol. 17:1425–1433.
   PubMed Web of Science ®Google Scholar
• Lee, S. E., J. K. Moore, A. Holmes, K. Umezu, R. D. Kolodner, and J. E. Haber. 1998. Saccharomyces Ku70, Mre11/Rad50 and RPA proteins regulate adaptation to G2/M arrest after DNA damage. Cell 94:399–409.
   PubMed Web of Science ®Google Scholar
• Lee, S. E., C. R. Pulaski, D. M. He, D. M. Benjamin, M. Voss, J. Um, and E. A. Hendrickson. 1995. Isolation of mammalian cell mutants that are X-ray sensitive, impaired in DNA double-strand break repair and defective for V(D)J recombination. Mutat. Res. 336:279–291.
   PubMedGoogle Scholar
• Lees-Miller, S. P., R. Godbout, D. W. Chan, M. Weinfeld, R. S. Day III, G. M. Barron, and J. Allalunis-Turner. 1995. Absence of p350 subunit of DNA-activated protein kinase from a radiosensitive human cell line. Science 267:1183–1185.
   PubMed Web of Science ®Google Scholar
• Li, G., C. Nelsen, and E. A. Hendrickson. 2002. Ku86 is essential in human somatic cells. Proc. Natl. Acad. Sci. USA 99:832–837.
   PubMed Web of Science ®Google Scholar
• Li, G. C., H. Ouyang, X. Li, H. Nagasawa, J. B. Little, D. J. Chen, C. C. Ling, Z. Fuks, and C. Cordon-Cardo. 1998. Ku70: a candidate tumor suppressor gene for murine T cell lymphoma. Mol. Cell 2:1–8.
   PubMed Web of Science ®Google Scholar
• Li, X., and W. D. Heyer. 2008. Homologous recombination in DNA repair and DNA damage tolerance. Cell Res. 18:99–113.
   PubMed Web of Science ®Google Scholar
• Lieber, M. R., Y. Ma, Y. Kefei, U. Pannicke, and K. Schwarz. 2006. The mechanism of vertebrate nonhomologous DNA end joining and its role in immune system gene rearrangements, p. 609-627. In W. Seide, Y. W. Kow, and P. Doetsch (ed.), DNA damage recognition. Taylor and Francis Group, New York, NY.
   Google Scholar
• Maser, R. S., and R. A. DePinho. 2002. Connecting chromosomes, crisis, and cancer. Science 297:565–569.
   PubMed Web of Science ®Google Scholar
• Masramon, L., M. Ribas, P. Cifuentes, R. Arribas, F. Garcia, J. Egozcue, M. A. Peinado, and R. Miro. 2000. Cytogenetic characterization of two colon cell lines by using conventional G-banding, comparative genomic hybridization, and whole chromosome painting. Cancer Genet. Cytogenet. 121:17–21.
   PubMedGoogle Scholar
• McElligott, R., and R. J. Wellinger. 1997. The terminal DNA structure of mammalian chromosomes. EMBO J. 16:3705–3714.
   PubMedGoogle Scholar
• McGuire, T. C., and M. J. Poppie. 1973. Hypogammaglobulinemia and thymic hypoplasia in horses: a primary combined immunodeficiency disorder. Infect. Immunol. 8:272–277.
   PubMed Web of Science ®Google Scholar
• Meek, K., S. Gupta, D. A. Ramsden, and S. P. Lees-Miller. 2004. The DNA-dependent protein kinase: the director at the end. Immunol. Rev. 200:132–141.
   PubMed Web of Science ®Google Scholar
• Meek, K., L. Kienker, C. Dallas, W. Wang, M. J. Dark, P. J. Venta, M. L. Huie, R. Hirschhorn, and T. Bell. 2001. SCID in Jack Russell terriers: a new animal model of DNA-PKcs deficiency. J. Immunol. 167:2142–2150.
   PubMedGoogle Scholar
• Montecucco, A., and G. Biamonti. 2007. Cellular response to etoposide treatment. Cancer Lett. 252:9–18.
   PubMed Web of Science ®Google Scholar
• Morozov, V. E., M. Falzon, C. W. Anderson, and E. L. Kuff. 1994. DNA-dependent protein kinase is activated by nicks and larger single-stranded gaps. J. Biol. Chem. 269:16684–16688.
   PubMed Web of Science ®Google Scholar
• Moshous, D., I. Callebaut, R. de Chasseval, B. Corneo, M. Cavazzana-Calvo, F. Le Deist, I. Tezcan, O. Sanal, Y. Bertrand, N. Philippe, A. Fischer, and J. P. de Villartay. 2001. Artemis, a novel DNA double-strand break repair/V(D)J recombination protein, is mutated in human severe combined immune deficiency. Cell 105:177–186.
   PubMed Web of Science ®Google Scholar
• Munoz, P., M. Z. Zdzienicka, J. M. Blanchard, and J. Piette. 1998. Hypersensitivity of Ku-deficient cells toward the DNA topoisomerase II inhibitor ICRF-193 suggests a novel role for Ku antigen during the G2 and M phases of the cell cycle. Mol. Cell. Biol. 18:5797–5808.
   PubMedGoogle Scholar
• Myung, K., G. Ghosh, F. J. Fattah, G. Li, H. Kim, A. Dutia, E. Pak, S. Smith, and E. A. Hendrickson. 2004. Regulation of telomere length and suppression of genomic instability in human somatic cells by Ku86. Mol. Cell. Biol. 24:5050–5059.
   PubMed Web of Science ®Google Scholar
• Nussenzweig, A., C. Chen, V. da Costa Soares, M. Sanchez, K. Sokol, M. C. Nussenzweig, and G. C. Li. 1996. Requirement for Ku80 in growth and immunoglobulin V(D)J recombination. Nature 382:551–555.
   PubMed Web of Science ®Google Scholar
• O'Driscoll, M., K. M. Cerosaletti, P. M. Girard, Y. Dai, M. Stumm, B. Kysela, B. Hirsch, A. Gennery, S. E. Palmer, J. Seidel, R. A. Gatti, R. Varon, M. A. Oettinger, H. Neitzel, P. A. Jeggo, and P. Concannon. 2001. DNA ligase IV mutations identified in patients exhibiting developmental delay and immunodeficiency. Mol. Cell 8:1175–1185.
   PubMed Web of Science ®Google Scholar
• Ouyang, H., A. Nussenzweig, A. Kurimasa, V. C. Soares, X. Li, C. Cordon-Cardo, W. Li, N. Cheong, M. Nussenzweig, G. Iliakis, D. J. Chen, and G. C. Li. 1997. Ku70 is required for DNA repair but not for T cell antigen receptor gene recombination in vivo. J. Exp. Med. 186:921–929.
   PubMed Web of Science ®Google Scholar
• Peterson, S. R., A. Kurimasa, M. Oshimura, W. S. Dynan, E. M. Bradbury, and D. J. Chen. 1995. Loss of the 350kD catalytic subunit of the DNA dependent protein kinase in DNA double-strand break repair mutant mammalian cells. Proc. Natl. Acad. Sci. USA 92:3171–3174.
   PubMed Web of Science ®Google Scholar
• Priestley, A., H. J. Beamish, D. Gell, A. G. Amatucci, M. C. Muhlmann-Diaz, B. K. Singleton, G. C. Smith, T. Blunt, L. C. Schalkwyk, J. S. Bedford, S. P. Jackson, P. A. Jeggo, and G. E. Taccioli. 1998. Molecular and biochemical characterisation of DNA-dependent protein kinase-defective rodent mutant irs-20. Nucleic Acids Res. 26:1965–1973.
   PubMed Web of Science ®Google Scholar
• Rathmell, W. K., and G. Chu. 1994. A DNA end-binding factor involved in double-strand break repair and V(D)J recombination. Mol. Cell. Biol. 14:4741–4748.
   PubMed Web of Science ®Google Scholar
• Riballo, E., S. E. Critchlow, S.-H. Teo, A. J. Doherty, A. Priestley, B. Broughton, B. Kysela, H. Beamish, N. Plowman, C. F. Arlett, A. R. Lehmann, S. J. Jackson, and P. A. Jeggo. 1999. Identification of a defect in DNA ligase IV in a radiosensitive leukemia patient. Curr. Biol. 9:699–702.
   PubMed Web of Science ®Google Scholar
• Riha, K., and D. E. Shippen. 2003. Ku is required for telomeric C-rich strand maintenance but not for end-to-end chromosome fusions in Arabidopsis. Proc. Natl. Acad. Sci. USA 100:611–615.
   PubMedGoogle Scholar
• Sekiguchi, J. M., and D. O. Ferguson. 2006. DNA double-strand break repair: a relentless hunt uncovers new prey. Cell 124:260–262.
   PubMedGoogle Scholar
• Shima, N., A. Alcaraz, I. Liachko, T. R. Buske, C. A. Andrews, R. J. Munroe, S. A. Hartford, B. K. Tye, and J. C. Schimenti. 2007. A viable allele of Mcm4 causes chromosome instability and mammary adenocarcinomas in mice. Nat. Genet. 39:93–98.
   PubMed Web of Science ®Google Scholar
• Shin, E. K., L. E. Perryman, and K. Meek. 1997. A kinase-negative mutation of DNA-PKcs in equine SCID results in defective coding and signal joint formation. J. Immunol. 158:3565–3569.
   PubMed Web of Science ®Google Scholar
• Shirasawa, S., M. Furuse, N. Yokoyama, and T. Sasazuki. 1993. Altered growth of human colon cancer cell lines disrupted at activated Ki-ras. Science 260:85–88.
   PubMed Web of Science ®Google Scholar
• Soulas-Sprauel, P., P. Rivera-Munoz, L. Malivert, G. Le Guyader, V. Abramowski, P. Revy, and J. P. de Villartay. 2007. V(D)J and immunoglobulin class switch recombinations: a paradigm to study the regulation of DNA end-joining. Oncogene 26:7780–7791.
   PubMed Web of Science ®Google Scholar
• Taccioli, G. E., A. G. Amatucci, H. J. Beamish, D. Gell, X. H. Xiang, M. I. Torres Arzayus, A. Priestley, S. P. Jackson, A. Marshak Rothstein, P. A. Jeggo, and V. L. Herrera. 1998. Targeted disruption of the catalytic subunit of the DNA-PK gene in mice confers severe combined immunodeficiency and radiosensitivity. Immunity 9:355–366.
   PubMed Web of Science ®Google Scholar
• Takai, H., R. C. Wang, K. K. Takai, H. Yang, and T. de Lange. 2007. Tel2 regulates the stability of PI3K-related protein kinases. Cell 131:1248–1259.
   PubMed Web of Science ®Google Scholar
• Tsuchida, R., T. Yamada, M. Takagi, A. Shimada, C. Ishioka, Y. Katsuki, T. Igarashi, L. Chessa, D. Delia, H. Teraoka, and S. Mizutani. 2002. Detection of ATM gene mutation in human glioma cell line M059J by a rapid frameshift/stop codon assay in yeast. Radiat. Res. 158:195–201.
   PubMedGoogle Scholar
• Uegaki, K., N. Adachi, S. So, S. Iiizumi, and H. Koyama. 2006. Heterozygous inactivation of human Ku70/Ku86 heterodimer does not affect cell growth, double-strand break repair, or genome integrity. DNA Repair 5:303–311.
   PubMed Web of Science ®Google Scholar
• van Steensel, B., A. Smogorzewska, and T. de Lange. 1998. TRF2 protects human telomeres from end-to-end fusions. Cell 92:401–413.
   PubMed Web of Science ®Google Scholar
• van Veelen, L., J. Wesoly, and R. Kanaar. 2006. Biochemical and cellular aspects of homologous recombination, p. 581-607. In W. Seide, Y. W. Kow, and P. Doetsch (ed.), DNA damage recognition. Taylor and Francis Group, New York, NY.
   Google Scholar
• Vasileva, A., and R. Jessberger. 2005. Precise hit: adeno-associated virus in gene targeting. Nat. Rev. Microbiol. 3:837–847.
   PubMed Web of Science ®Google Scholar
• Wiler, R., R. Leber, B. B. Moore, L. F. VanDyk, L. E. Perryman, and K. Meek. 1995. Equine severe combined immunodeficiency: a defect in V(D)J recombination and DNA-dependent protein kinase activity. Proc. Natl. Acad. Sci. USA 92:11485–11489.
   PubMed Web of Science ®Google Scholar
• Woods, T., W. Wang, E. Convery, A. Errami, M. Z. Zdzienicka, and K. Meek. 2002. A single amino acid substitution in DNA-PKcs explains the novel phenotype of the CHO mutant, XR-C2. Nucleic Acids Res. 30:5120–5128.
   PubMedGoogle Scholar
• Zdzienicka, M. Z. 1999. Mammalian X-ray-sensitive mutants which are defective in non-homologous (illegitimate) DNA double-strand break repair. Biochimie 81:107–116.
   PubMedGoogle Scholar
• Zhu, C., M. A. Bogue, D. S. Lim, P. Hasty, and D. B. Roth. 1996. Ku86-deficient mice exhibit severe combined immunodeficiency and defective processing of V(D)J recombination intermediates. Cell 86:379–389.
   PubMedGoogle Scholar