Accessibility control and machinery of immunoglobulin class switch recombination

Journal of Leukocyte Biology

Volumn 73, Issue 3, 2003, Pages 323-332

  Zhang, Ke ^a  

^a UNIVERSITY OF CALIFORNIA   (United States)

Author keywords

3 Ig enhancer; DNA repair; Germline transcription

Indexed keywords

CD40 ANTIGEN; CYTOKINE; DOUBLE STRANDED DNA; IMMUNOGLOBULIN A; IMMUNOGLOBULIN HEAVY CHAIN; IMMUNOGLOBULIN G;

B LYMPHOCYTE; CONFORMATIONAL TRANSITION; DNA CLEAVAGE; DNA CONFORMATION; DNA REPAIR; DNA STRAND BREAKAGE; ENHANCER REGION; GENE CONTROL; GENE FUNCTION; GENETIC TRANSCRIPTION; GERM LINE; IMMUNOGLOBULIN CLASS SWITCH RECOMBINATION; IMMUNOGLOBULIN PRODUCTION; NONHUMAN; PRIORITY JOURNAL; REGULATOR GENE; REVIEW; SIGNAL TRANSDUCTION; GENE EXPRESSION REGULATION; GENE REARRANGEMENT; GENETIC RECOMBINATION; GENETICS; HUMAN; IMMUNOGLOBULIN GENE;

GENE EXPRESSION REGULATION; HUMANS; IMMUNOGLOBULIN CLASS SWITCHING; IMMUNOGLOBULIN G; IMMUNOGLOBULIN SWITCH REGION; RECOMBINATION, GENETIC; TRANSCRIPTION, GENETIC;

EID: 0037686188     PISSN: 07415400     EISSN: None     Source Type: Journal    
DOI: 10.1189/jlb.0702339     Document Type: Review

References (114)

1. Wang, A. C., Wilson, S. K., Hooper, J. E., Fudenberg, H. H., Nisonoff, A. (1970) Evidence for control of synthesis of the variable regions of the heavy chains of immunoglobulin G and M by the same gene. Proc. Natl. Acad. Sci. USA 66, 337-341.
2. Cooper, M. D., Kearney, J. F., Lydyard, P. M., Grossi, C. E., Lawman, A. R. (1977) Studies of generation of B-cell diversity in mouse, man and chicken. Cold Spring Harb. Symp. Quant. Biol. 41, 139-145.
3. Pernis, B., Forni, L., Luzzati, A. L. (1977) Synthesis of multiple immunoglobulin classes by single lymphocytes. Cold Spring Harb. Symp. Quant. Biol. 41, 175-183.
4. Zhang, J., Alt, F. W., Honjo, T. (1995) Regulation of class switch recombination of the immunoglobulin heavy chain genes. In Immunoglobulin Genes, 2nd ed. (T. Honjo, F. W. Alt, eds.), Academic Press, London, 235-265.
5. Stavnezer, J. (2000) Molecular processes that regulate class switching. Curr. Top. Microbiol. Immunol. 245, 127-168.
6. Honjo, T., Kinoshita, K., Muramatsu, M. (2002) Molecular mechanism of class switch recombination: linkage with somatic hypermutation. Annu. Rev. Immunol. 20, 165-196.
7. Litinskiy, M. B., Nardelli, B., Hibert, D. M., He, B., Schaffer, A., Casali, P., Cerutti, A. (2002) DCs induce CD40-independent immunoglobulin class switching through BlyS and APRIL. Nat. Immunol. 3, 822-829.
8. Snapper, C. F., Marcu, K. B., Zelazowski, P. (1997) The immunoglobulin class switch: beyond "accessibility". Immunity 6, 217-223.
9. Fujieda, S., Zhang, K., Saxon, A. (1995) IL-4 plus CD40 monoclonal antibody induces human B cells γ subclass-specific isotype switch: switching to γ1, γ3 and γ4, but not γ2. J. Immunol. 155, 2318-2328.
10. Defrance, T., Vanbervliet, B., Briere, F., Durand, I., Rousset, F., Banchereau, J. (1992) Interleukin 10 and transforming growth factor beta cooperate to induce anti-CD40-activated naive human B cells to secrete immunoglobulin A. J. Exp. Med. 175, 671-682.
11. Punnonen, J., Aversa, G., Cocks, B. G., McKenzie, A. N., Menson, S., Zurawski, G., De Waal Malefyt, R., de Vries, J. E. (1993) Interleukin 13 induces interleukin 4-independent IgG4 and IgE synthesis and CD23 expression by human B cells. Proc. Natl. Acad. Sci. USA 90, 3730-3734.
12. van Vlasselaer, P., Punnonen, J., de Vries, J. E. (1992) Transforming growth factor-beta directs IgA switching in human B cells. J. Immunol. 148, 2062-2070.
13. Fujieda, S., Waschek, J. A., Zhang, K., Saxon, A. (1996) Vasoactive intestinal peptide induces Sα/Sμ switch circular DNA in human B cells. J. Clin. Investig. 98, 1527-1532.
14. Fayette, J., Dubois, B., Vandenabeele, S., Bridon, J. M., Vanbervliet, B., Durand, I., Banchereau, J., Caux, C., Briere, F. (1997) Human dendritic cells skew isotype switching of CD40-activated naive B cells towards IgA1 and IgA2. J. Exp. Med. 185, 1909-1918.
15. Stavnezer, J., Radcliffe, G., Lin, Y. C., Nietupski, J., Berggren, L., Sitia, R., Sevednson, E. (1988) Immunoglobulin heavy-chain switching may be directed by prior induction of transcripts from constant-region genes. Proc. Natl. Acad. Sci. USA 85, 7704-7708.
16. Jung, S., Rajewsky, K., Radbruch, A. (1993) Shutdown of class switch recombination by deletion of a switch region control element. Science 259, 984-987.
17. 2b promoter and exon. EMBO J. 12, 3529-3537.
18. Bottaro, A., Lansford, R., Xu, L., Zhang, J., Rothman, P., Alt, F. W, (1994) S region transcription per se promotes basal IgE class switch recombination but additional factors regulate the efficiency of the process. EMBO J. 13, 665-674.
19. Lorenz, M., Jung, S., Radbruch, A. (1995) Switch transcripts in immunoglobulin class switching. Science 267, 1825-1828.
20. Harriman, G. R., Bradley, A., Das, S., Rogers-Fani, P., Davis, A. C. (1996) IgA class switch in I exon-deficient mice: role of germline transcription in class switch recombination. J. Clin. Investig. 97, 477-485.
21. Kim, C. G., Epner, E. M., Forrester, W. C., Groudine, M. (1992) Inactivation of the human beta-globin gene by targeted insertion into the beta-globin locus control region. Genes Dev. 6, 928-938.
22. r cassettes inserted into the immunoglobulin heavy chain constant region locus. Proc. Natl. Acad. Sci. USA 96, 3000-3004.
23. Pinaud, E., Khamlichi, A., Morvan, C., Drouet, M., Nalesso, V., Bert, M., Cogne, M. (2001) Localization of the 3′ IgH locus elements that effect long-distance regulation of class switch recombination. Immunity 15, 187-199.
24. Kinoshita, K., Tashiro, J., Tomita, S., Lee, C. G., Honjo, T. (1998) Target specificity of immunoglobulin class switch recombination is not determined by nucleotide sequences of S regions. Immunity 9, 849-858.
25. Zhang, K., Zhang, L., Yamada, Y., Vu, M., Lee, A., Saxon, A. (2002) Efficiency of Iε promoter-directed switch recombination in GFP expression-based switch constructs works synergistically with other promoter and/or enhancer elements but is not tightly linked to the strength of transcription. Eur. J. Immunol. 32, 424-434.
26. Muramatsu, M., Sankaranand, V. S., Anant, S., Sugai, M., Kinoshita, K., Davidson, N. O., Honjo, T. (1999) Specific expression of activation-induced cytidine deaminase (AID), a novel member of the RNA-editing deaminase family in germinal center B cells. J. Biol. Chem. 274, 18470-18476.
27. Muramatsu, M., Kinoshita, K., Fagarasan, S., Yamada, S., Shinkai, Y., Honjo, T. (2000) Class switch recombination and hypermutation require activation-induced cytidine deaminase (AID), a potential RNA editing enzyme. Cell 102, 553-563.
28. Chen, X. H., Patel, B. K., Wang, L. M., Frankel, M., Ellmore, N., Flavell, R. A., LaRochelle, W. J., Pierce, J. H. (1997) Jak1 expression is required for mediating interleukin-4-induced turosine phosphorylation of insulin receptor substrate and Stat6 signaling molecules. J. Biol. Chem. 272, 6556-6560.
29. Shi, M. J., Stavnezer, J. (1998) CBF (AML2) is induced by TGF-β1 to bind and activate the mouse germline Ig promoter. J. Immunol. 161, 6751-6760.
30. Hanai, J. I., Chen, L. F., Kanno, K., Ohtani-Fujita, N., Kim, W. Y., Guo, W. H., Imamura, T., Ishidou, Y., Fukuchi, M., Shi, M. J., Stavnezer, J., Kawabata, M., Miyazono, K., Ito, Y. (1999) Interaction and functional cooperation of PEBP2/CBF with Smads. Synergistic induction of the immunoglobulin germline C promoter. J. Biol. Chem. 274, 31577-31582.
31. Iciek, L. A., Delphin, S. A., Stavnezer, J. (1997) CD40 cross-linking induces Ig epsilon germline transcripts in B cells via activation of NF-kappaB: synergy with IL-4 induction. J. Immunol. 158, 4769-4719.
32. Shen, C. H., Stavnezer, J. (1998) Interaction of stat6 and NF-kappaB: direct association and synergistic activation of interleukin-4-induced transcription. Mol. Cell. Biol. 18, 3395-3404.
33. Shen, C. H., Stavnezer, J. (2001) Activation of the mouse Ig germline 2 promoter by IL-4 is dependent on AP-1 transcription factors. J. Immunol. 166, 411-423.
34. Yamada, T., Zhu, D., Saxon, A., Zhang, K. (2002) CD45 controls IL-4-mediated IgE class switch recombination in human B cells as JAK phosphatase. J. Biol. Chem. 277, 28830-28835.
35. Irie-Sasaki, J., Sasaki, T., Matsumoto, W., Opavsky, A., Cheng, M., Welstead, G., Griffiths, E., Krawczyk, C., Richardson, C. D., Aitken, K., Iscove, N., Koretzky, G., Johnson, P., Liu, P., Rothstein, D. M., Penninger, J. M. (2001) CD45 is a JAK phosphatase and negatively regulates cytokine receptor signaling. Nature 409, 349-354.
36. Lieberson, R., Giannini, S. L., Birshtein, B. K., Eckhardt, L. A. (1991) An enhancer at the 3′ end of the mouse immunoglobulin heavy chain locus. Nucleic Acids Res. 19, 933-937.
37. Mills, F. C., Harindranath, N., Mitchell, M., Max, E. E. (1997) Enhancer complexes located downstream of both human immunoglobulin Ca genes. J. Exp. Med. 186, 845-858.
38. Gregor, P. D., Morrison, S. L. (1986) Myeloma mutant with a novel 3′ flanking region: loss of normal sequence and insertion of repetitive elements leads to decreased transcription but normal processing of the alpha heavy-chain gene products. Mol. Cell. Biol. 6, 1903-1916.
39. Giannini, S. L., Singh, M., Calvo, C. F., Ding, G., Birshtein, B. K. (1993) DNA regions flanking the mouse Ig 3′ enhancer are differentially methylated and DNase I hypersensitive during B cell differentiation. J. Immunol. 150, 1772-1780.
40. Sideras, P., Nilsson, L., Islam, K. B., Quintana, I. Z., Freihof, L., Rosen, A., Juliusson, G., Hammarström, L., Smith, C. L. (1992) Transcription of unrearranged Ig H chain genes in human B cell malignancies: biased expression of genes encoded within the first duplication unit of the Ig H chain locus. J. Immunol. 149, 244-252.
41. Tumas-Brundage, K. M., Vora, K. A., Manser, T. (1997) Evaluation of the role of the 3′ heavy chain enhancer [3′ E(hs1,2)] in Vh gene somatic hypermutation. Mol. Immunol. 34, 367-378.
42. Lieberson, R., Ong, J., Shi, X., Eckhardt, L. A. (1995) Transcription of an immunoglobulin γ2a gene ceases upon deletion of an enhancer region over 70 kilobases away. EMBO J. 14, 6229-6238.
43. Hu, Y., Pan, Q., Pardali, E., Mills, F. C., Bernstein, R. M., Max E. E., Sideras, P., Hammarstrom, L. (2000) Regulation of germline promoters by the two human Ig heavy chain 3′ α enhancers. J. Immunol. 164, 6380-6386.
44. Laurencikiene, J., Deveilaite, V., Severinson, E. (2001) HS1,2 enhancer regulation of germline ε and γ2b promoters in murine B lymphocytes: Evidence for specific promoter-enhancer interactions. J. Immunol. 167, 3257-3265.
45. Bulger, M., Groudine, R. (1999) Looping versus linking: toward a model for long-distance gene activation. Genes Dev. 13, 2465-2477.
46. Madisen, L., Groudine, M. (1994) Identification of a locus control region in the immunoglobulin heavy-chain locus that deregulates c-myc expression in plasmacytoma and Burkitt's lymphoma cells. Genes Dev. 8, 2212-2226.
47. Shi, X., Eckhardt, L. A. (2001) Deletional analyses reveal an essential role for the hs3b/hs/4 IgH 3′ α enhancer pair in an Ig-secreting but not an earlier-stage B cell line. Int. Immunol. 13, 1003-1012.
48. Chauveau, C., Jansson, E. A., Muller, S., Cogne, M., Pettersson, S. (1999) Ig heavy chain 3′ HS1-4 directs correct spatial position-independent expression of a linked transgene to B lineage cells. J. Immunol. 163, 4637-4641.
49. Singh, M., Birshtein, B. K. (1993) NF-HB (BSAP) is a repressor of the murine immunoglobulin heavy-chain 3 enhancer at early stages of B cell differentiation. Mol. Cell. Biol. 16, 3611-3622.
50. Neurath, M. F., Max, E. E., Strober, W. (1995) Pax5 (BSAP) regulates the murine immunoglobulin 3′α enhancer by suppressing binding of NF-P, a protein that controls heavy chain transcription. Proc. Natl. Acad. Sci. USA 92, 5336-5340.
51. Stevens, S., Ong, J., Kim, U., Eckhardt, A. L., Roeder, R. G. (2000) Role of OCA-B in 3′-IgH enhancer function. J. Immunol. 164, 5306-5312.
52. Michaelson, J. S., Singh, M., Snapper, C. M., Sha, W. C., Baltimore, D., Birshtein, B. K. (1996) Regulation of 3′ IgH enhancers by a common set of factors, including kappa B-binding proteins. J. Immunol. 156, 2828-2839.
53. Meyer, K. B., Skoberg, M., Margenfeld, C., Ireland, J., Petterson, S. (1996) Repression of the immunoglobulin heavy chain 3′α enhancer by helix-loop helix protein Id3 via a functionally important E47/E12 binding site: implications for developmental control of enhancer function. Eur. J. Immunol. 25, 1770-1777.
54. Kanda, K., Hu, H-M., Zhang, L., Grandchamps, J., Boxer, L. (2000) NF-κB activity is required for the deregulation of e-myc expression by the immunoglobulin heavy chain enhancer. J. Biol. Chem. 275, 32338-32346.
55. Zelazowski, P., Shen, Y., Snapper, C. M. (2000) NF-κB/p50 and NF-κB/c-Rel differentially regulate the activity of the 3′E-hs1,2 enhancer in normal murine B cells in an activation-dependent manner. Int. Immunol. 12, 1167-1172.
56. Arulampalam, V., Grant, P. A., Samuelsson, A., Lendahl, U., Pettersson, S. (1994) Lipopolysaccharide-dependent transactivation of the temporally regulated immunoglobulin heavy chain 3′ enhancer. Eur. J. Immunol. 24, 1671-1677.
57. Grant, P. A., Andersson, T., Neurath, M. F., Arulampalam, V., Bauch, A., Müller, R., Reth, M., Pettersson, S. (1996) A T cell controlled molecular pathway regulating the IgH locus: CD40-mediated activation of the IgH 3′ enhancer. EMBO J. 15, 6691-6700.
58. Cogne, M., Lansford, R., Bottaro, A., Zhang, J., Gorman, J., Young, F., Cheng, H. L., Alt, F. W. (1994) A class switch control region at the 3′ end of the immunoglobulin heavy chain locus. Cell 77, 737-747.
59. Groundine, M. (1997) Histone acetylation in chromatin structure and transcription. Nature 389, 349-352.
60. Struhl, K. (1998) Histone acetylation and transcriptional regulatory mechanisms. Genes Dev. 12, 599-606.
61. Madisen, L., Krumin, A., Hebbes, T., Groundine, M. (1998) The immunoglobulin heavy chain locus control region increases histone acetylation along linked c-myc gene. Mol. Cell. Biol. 18, 6281-6292.
62. McMurry, T. M., Krangel, M. S. (2000) A role for histone acetylation in the developmental regulation of V(D)J recombination. Science 287, 495-498.
63. Agata, Y., Katakai, T., Ye, S-K., Sugai, M., Gonda, H., Honjo, T., Koichi, I., Shimizu, A. (2001) Histone acetylation determines the developmentally regulated accessibility for T cell receptor g gene recombination. J. Exp. Med. 193, 873-879.
64. Drouin, E. E., Schrader, C. E., Stavnezer, J., Hansen, U. (2002) The ubiquitously expressed DNA-binding protein late SV40 factor binds Ig switch regions and represses class switching to IgA. J. Immunol. 168, 2847-2856.
65. Qiu, G., Stavnezer, J. (1998) Overexpression of BSAP/Pax-5 inhibits switching to IgA and enhances switching to IgE in the 1.29μ B cell line. J. Immunol. 161, 2906-2918.
66. 4 DNA binding by LR1 and its subunits, nucleolin and hnRNP D: a role for G-G pairing in immunoglobulin switch recombination. J. Biol. Chem. 274, 1066-1071.
67. Wuerffel, R., Jamieson, C. E., Morgan, L., Merkulov, G. V., Sen, R., Kenter, A. L. (1992) Switch recombination breakpoints are strictly correlated with DNA recognition motifs for immunoglobulin S3 DNA-binding proteins. J. Exp. Med. 176, 339-349.
68. Mizuta, T. R., Fukita, Y., Miyoshi, T., Shimizu, A., Honjo, T. (1993) Isolation of cDNA encoding a binding protein specific to 5′-phosphorylated single-stranded DNA with G-rich sequences. Nucleic Acids Res. 21, 1761-1766.
69. Lyon, C. J., Miranda, G. A., Piao, J. S., Aguilera, R. J. (1996) Characterization of an endonuclease activity which preferentially cleaves the G-rich immunoglobulin switch repeat sequences. Mol. Immunol. 33, 157-169.
70. Li, M. J., Peakaman, M. C., Golub, E. I., Reddy, G, Ward, D. C., Radding, C. M., Maizels, N. (1996) Rad51 expression and localization in B cells carrying out class switching recombination. Proc. Natl. Acad. Sci. USA 93, 10222-10226.
71. Borggrefe, T., Wabl, M., Akhmedov, A. T., Jessberger, R. (1998) A B-cell-specific DNA recombination complex. J. Biol. Chem. 273, 17025-17035.
72. Kataota, T., Miyata, T., Honjo, T. (1981) Repetitive sequences in class switch recombination regions of immunoglobulin heavy chain genes. Cell 23, 357-368.
73. Reaban, M. E., Griffin, J. A. (1990) Induction of RNA-stabilized DNA conformers by transcription of an immunoglobulin switch region. Nature 348, 342-344.
74. Daniels, G. A., Lieber, M. R. (1995) RNA: DNA complex formation upon transcription of immunoglobulin switch regions: implications for the mechanism and regulation of class switch recombination. Nucleic Acids Res, 23, 5006-5011.
75. Tian, M., Alt, F. W. (2000) Transcription-induced cleavage of immunoglobulin switch regions by nucleotide excision repair nucleases in vitro. J. Biol. Chem. 275, 24163-24172.
76. , Lee, C. G., Kinoshita, K., Arudchandran, A., Cerritelli, S. M., Crouch, R. J., Honjo, T. (2001) Quantitative regulation of class switch recombination by S region transcription. J. Exp. Med. 194, 365-374.
77. Sen, D., Gilbert, W. (1988) Formation of parallel four-stranded complexes by guanine-rich motifs in DNA and its implications for meiosis. Nature 334, 364-366.
78. Tashiro, J., Kinoshita, K., Honjo, T. (2001) Palindromic but not G-rich sequences are targets of class switch recombination. Int. Immunol. 13, 495-505.
79. Luby, T. M., Schrader, C. E., Stavnezer, J., Selsing, E. (2001) The μ switch region tandem repeats are important, but not required, for antibody class switch recombination. J. Exp. Med. 193, 159-168.
80. Revy, P., Muto, T., Levy, Y., Geissmann, F., Plebani, A., Sanal, O., Catalan, N., Forveille, M., Dufourcq-Labelouse, R., Gennery A., Tezcan, I., Ersoy, F., Kayserili, H., Ugazio, A. G., Brousse, N., Muramatsu, M., Notarangelo, L. D., Kinoshita, K., Honjo, T., Fisher, A., Durandy, A. (2000) Activation-induced cytidine deaminase (AID) deficiency causes the autosomal recessive form of the Hyper-IgM syndrome (HIGM2). Cell 102, 565-575.
81. Martin, A., Bardwell, P. D., Woo, J. G., Fan, M., Shulman, M. J., Scharff, M. D. (2002) Activation-induced cytidine deaminase turns on somatic hypermutation in hybridomas. Nature 415, 802-806.
82. Arakawa, H., Hauschild, J., Buerstedde, J. M. (2002) Requirement of the activation-induced deaminase (AID) gene for immunoglobulin gene conversion. Science 295, 1301-1306.
83. Okazaki, I., Kinoshita, K., Muramatsu, M., Yoshikawa, K., Honjo, T. (2002) The AID enzyme induces class switch recombination in fibroblast. Nature 416, 340-345.
84. Peterson-Mahrt, S. K., Harris, R. S., Neuberger, M. S. (2002) AID mutates E. coli suggesting a DNA deamination mechanism for antibody diversification. Nature 418, 99-103.
85. Papavasiliou, F. N., Schatz, D. G. (2002) The activation-induced deaminase functions in a postcleavage step of the somatic hypermutation process. J. Exp. Med. 195, 1193-1198.
86. Bross, L., Muramatsu, M., Kinoshita, K., Honjo, T., Jacobs, H. (2002) DNA double-strand breaks: prior to but not sufficient in targeting hypermutation. J. Exp. Med. 195, 1187-1192.
87. Chung, I. K., Mehta, V. B., Spitzner, J. R., Muller, M. T. (1992) Eukaryotic topoisomerase II cleavage of parallel stranded DNA tetraplexes. Nucleic Acids Res. 20, 1973-1977.
88. Shanmugam, A., Shi, M., Yauch, L., Stavnezer, J., Kenter, A. L. (2000) Evidence for class-specific factors in immunoglobulin isotype switching. J. Exp. Med. 191, 1365-1380.
89. Ma, L., Wortis, H. H., Kenter, A. L. (2002) Two new isotype-specific switching activities detected for Ig class switching. J. Immunol. 168, 2835-2846.
90. Zhang, K., Mills, F. C., Saxon, A. (1994) Switch circles from IL-4 directed ε class switching from human B lymphocytes: evidence for direct, sequential and multiple step sequential switch from μ to ε immunoglobulin heavy chain gene. J. Immunol. 152, 3427-3435.
91. Chen, X., Kinoshita, K., Honjo, T. (2001) Variable deletion and duplication at recombination junction ends: implication for staggered double-strand cleavage in class switch recombination. Proc. Natl. Acad. Sci. USA 98, 13860-13865.
92. Difilippantonio, M. J., Zhu, J., Chen, H. T., Meffre, E., Nussenzweig, M. C., Max, E. E., Ried, T., Nussenzweig, A. (2000) DNA repair protein Ku80 suppresses chromosomal aberrations and malignant transformation. Nature 404, 510-514.
93. Nick McElhinny, S. A., Snowden, C. M., McCarville, J., Ramsden, D. A. (2000) Ku recruits the XRCC4-ligase IV complex to DNA ends. Mol. Cell. Biol. 20, 2996-3003.
94. Manis, J. P., Gu, Y., Lansford, R., Sonoda, E., Ferrini, R., Davidson, L., Rajewsky, K., Alt, F. W. (1998) Ku 70 is required for late B cell development and immunoglobulin heavy chain class switching. J. Exp. Med. 187, 2081-2089.
95. Casellas, R., Nussenzweig, A., Wuerffel, R., Pelanada, R., Reichlin, A., Suh, H., Qin, X. F., Besmer, E., Kenter, A., Rajewsky, A., K. (1998) Ku 80 is required for immunoglobulin isotype switching. EMBO J. 17, 2404-2411.
96. Rolink, A., Melchers, F., Anderson, J. (1996) The SCID but not the RAG-2 gene products is required for Sμ,-Sε heavy chain class switching. Immunity 5, 319-330.
97. Manis, J. P., Dudley, D., Kaylor, L., Alt, F. W. (2002) IgH class switch recombination to IgG1 in DNA-PKcs-deficient B cells. Immunity 16, 607-617.
98. Zelazowski, P., Max, E. E., Kehry, M. R., Snapper, C. M. (1997) Regulation of Ku expression in normal murine B cells by stimuli that promoter switch recombination. J. Immunol. 159, 2559-2564.
99. Morio, T., Hanissian, S. H., Bacharier, L. B., Teraoka, H., Nonoyama, S., Seki, M., Kondo, J., Nakano, H., Lee, S-K., Geha, R. S., Yata, J. I. (1999) Ku in the cytoplasm association with CD40 in human B cells and translocates into the nucleus following incubation with IL-4 and anti-CD40 mAb. Immunity 11, 339-348.
100. Moshous, D., Callebaut, I., de Chasseval, R., Corneo, B., Cavazzana-Calvo, M., Le Deist, F., Tezcan, I., Sanal, O., Bertrand, Y., Philippe, N., Fischer, A., de Villarty, J. P. (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.
101. Ma, Y., Pannicke, U., Schwarz, K., Lieber, M. R. (2002) Hairpin opening and overhang processing by an artemis/DNA-dependent protein kinase complex in nonhomologous end joining and V(D)J recombination. Cell 108, 781-794.
102. Schradera, C. E., Edelmannb, W., Kucherlapatic, R., Stavnezer, J. (1999) Reduced isotype switching in splenic B cells from mice deficient in mismatch repair enzymes. J. Exp. Med. 190, 323-330.
103. Ehrenstein, M. R., Neuberger, M. S. (1999) Deficiency in Msh2 affects the efficiency and local sequence specificity of immunoglobulin class-switch recombination: parallels with somatic hypennutation. EMBO J. 18, 3484-3490.
104. Ehrenstein, M. R., Rada, C., Jones, A. M., Milstein, C., Neuberger, M. S. (2001) Switch junction sequences in PMS2-deficient mice reveal a microhoniology-mediated mechanism of Ig class switch recombination. Proc. Natl. Acad. Sci. USA 98, 14553-14558.
105. Schrader, C. E., Vardo, J., Stavnezer, J. (2002) Role for mismatch repair proteins Msh2, Mlh1, and Pms2 in immunoglobulin class switching shown by sequence analysis of recombination junctions. J. Exp. Med. 195, 367-373.
106. Wilson, T. E., Lieber, M. R. (1999) Efficient processing of DNA ends during yeast nonhomologous end joining. Evidence for a DNA polymerase beta (Pol4)-dependent pathway. J. Biol. Chem. 274, 23599-23609.
107. Dunick, W., Hertz, G. Z., Scappino, L., Gritzmacher, C. (1993) DNA sequences at immunoglobulin switch region recombination sites. Nucleic Acids Res. 21, 365-376.
108. Zan, H., Komori, A., Li, Z., Cerutti, A., Schaffer, A., Flajnik, M. F., Diaz, M., Casali, P. (2001) The translesion DNA polymerase zeta plays a major role in Ig and bcl-6 somatic hypermutation. Immunity 14, 643-653.
109. Grawunder, U., Zimmer, D., Fugmann, S., Schwarz, K., Lieber, M. R. (1998) DNA ligase IV is essential for V(D)J recombination and DNA double-strand break repair in human precursor lymphocytes. Mol. Cell 2, 477-484.
110. Grawunder, U., Wilm, M., Wu, X., Kulesza, P., Wilson, T. E., Mann, M., Lieber, M. R. (1997) Activity of DNA ligase IV stimulated by complex formation with XRCC4 protein in mammalian cells. Nature 388, 492-495.
111. O'Driscoll, M., Cerosaletti, K. M., Girard, P. M., Dai, Y., Stumm, M., Kysela, B., Hirsch, B., Gennery, A., Palmer, S. E., Seidel, J., Gatti, R. A., Varon, R., Oettinger, M. A., Neitzel, H., Jeggo, P. A., Concannon, P. (2001) DNA ligase IV mutations identified in patients exhibiting developmental delay and immunodeficiency. Mol. Cell 8, 1175-1185.
112. Petersen, S., Casellas, R., Reina-San-Martin, B., Chen, H. T., Difilippantonio, M. J., Wilson, P. C., Hanitsch, L., Celeste, A., Muramatsu, M., Pilch, D. R., Redon, C., Ried, T., Bonner, W. M., Honjo, T., Nussenzweig, M. C., Nussenzweig, A. (2001) AID is required to initiate Nbs1/gamma-H2AX focus formation and mutations at sites of class switching. Nature 414, 660-665.
113. Harfst, E., Cooper, S., Neubauer, S., Distel, L., Grawunder, U. (2000) Normal V(D)J recombination in cells from patients with Nijmegen breakage syndrome. Mol. Immunol. 37, 915-929.
114. Pan, Q., Petit-Frere, C., Lahdesmaki, A., Gregorek, H., Chrzanowska, K. H., Hammarstrom, L. (2002) Alternative end joining during switch recombination in patients with Ataxia-Telangiectasia. Eur. J. Immunol. 32, 1300-1308.