title: Chance, genetics, and the heterogeneity of disease and pathogenesis in systemic lupus erythematosus.
abstract: Systemic lupus erythematosus (SLE) is a remarkably complex and heterogeneous systemic autoimmune disease. Disease complexity within individuals and heterogeneity among individuals, even genetically identical individuals, is driven by stochastic execution of a complex inherited program. Genome-wide association studies (GWAS) have progressively improved understanding of which genes are most critical to the potential for SLE and provided illuminating insight about the immune mechanisms that are engaged in SLE. What initiates expression of the genetic program to cause SLE within an individual and how that program is initiated remains poorly understood. If we extrapolate from all of the different experimental mouse models for SLE, we can begin to appreciate why SLE is so heterogeneous and consequently why prediction of disease outcome is so difficult. In this review, we critically evaluate extrinsic versus intrinsic cellular functions in the clearance and elimination of cellular debris and how dysfunction in that system may promote autoimmunity to nuclear antigens. We also examine several mouse models genetically prone to SLE either because of natural inheritance or inheritance of induced mutations to illustrate how different immune mechanisms may initiate autoimmunity and affect disease pathogenesis. Finally, we describe the heterogeneity of disease manifestations in SLE and discuss the mechanisms of disease pathogenesis with emphasis on glomerulonephritis. Particular attention is given to discussion of how anti-DNA autoantibody initiates experimental lupus nephritis (LN) in mice.
authors.affiliation: Department of Microbiology, Immunology and Biochemistry, University of Tennessee Health Science Center, 858 Madison Avenue, Memphis, TN, 38163, USA, tmarion@uthsc.edu.
authors.forename: Tony N
authors.initials: TN
authors.lastname: Marion
authors.forename: Arnold E
authors.initials: AE
authors.lastname: Postlethwaite
mesh_terms: D000818:Animals; D015551:Autoimmunity; D004195:Disease Models, Animal; D006801:Humans; D007108:Immune Tolerance; D008180:Lupus Erythematosus, Systemic / Q000150:complications / Q000209:etiology* / Q000473:pathology*; D051379:Mice
references.citation: J Immunol. 2012 Jul 15;189(2):503-9
references.pmid: 22773662
references.citation: Br Med J. 1962 Apr 28;1(5286):1175-8
references.pmid: 13894877
references.citation: PLoS One. 2008 Mar 05;3(3):e1727
references.pmid: 18320046
references.citation: J Clin Invest. 2002 Dec;110(12):1823-30
references.pmid: 12488432
references.citation: Cytokine. 2013 May;62(2):244-52
references.pmid: 23561928
references.citation: J Immunol. 1996 Sep 1;157(5):2136-42
references.pmid: 8757338
references.citation: Transplantation. 2004 Aug 15;78(3):324-32
references.pmid: 15316358
references.citation: Semin Arthritis Rheum. 2010 Aug;40(1):32-52
references.pmid: 19880162
references.citation: Eur J Immunol. 2007 Jun;37(6):1691-701
references.pmid: 17523212
references.citation: PLoS Genet. 2007 Nov;3(11):e192
references.pmid: 17997607
references.citation: Scand J Immunol. 1998 Sep;48(3):223-32
references.pmid: 9743205
references.citation: J Dermatol Sci. 2013 Jul;71(1):22-8
references.pmid: 23664188
references.citation: Lupus. 2010 Jan;19(1):73-82
references.pmid: 19900981
references.citation: Am J Surg Pathol. 1990 Dec;14(12):1112-25
references.pmid: 2252102
references.citation: Neuroimmunomodulation. 2001;9(1):49-54
references.pmid: 11435752
references.citation: Autoimmun Rev. 2013 Jan;12(3):426-9
references.pmid: 22944298
references.citation: J Exp Med. 1957 Nov 1;106(5):649-60
references.pmid: 13475621
references.citation: Can Respir J. 2012 Jan-Feb;19(1):50-2
references.pmid: 22332137
references.citation: Nat Genet. 1997 Dec;17(4):393-8
references.pmid: 9398839
references.citation: J Exp Med. 1982 Sep 1;156(3):766-77
references.pmid: 7108443
references.citation: Curr Opin Immunol. 1998 Dec;10(6):718-25
references.pmid: 9914223
references.citation: Ann Rheum Dis. 1992 Feb;51(2):156-9
references.pmid: 1550395
references.citation: Clin Genet. 1985 Jun;27(6):535-42
references.pmid: 4017274
references.citation: Liver. 2000 Oct;20(5):366-73
references.pmid: 11092254
references.citation: J Exp Med. 1994 Dec 1;180(6):2341-6
references.pmid: 7964507
references.citation: Arthritis Rheum. 2008 Mar;58(3):843-53
references.pmid: 18311802
references.citation: J Immunol. 2014 Jun 1;192(11):5314-23
references.pmid: 24771850
references.citation: Prog Neurobiol. 2011 Nov;95(3):301-33
references.pmid: 21889967
references.citation: Biochim Biophys Acta. 2003 Nov 20;1639(3):169-76
references.pmid: 14636948
references.citation: J Immunol. 1993 Aug 1;151(3):1614-26
references.pmid: 8393048
references.citation: J Exp Med. 1985 Feb 1;161(2):378-91
references.pmid: 3919141
references.citation: Rheumatology (Oxford). 2001 Apr;40(4):424-31
references.pmid: 11312382
references.citation: Clin Immunol Immunopathol. 1995 Jul;76(1 Pt 1):59-67
references.pmid: 7606869
references.citation: Nat Rev Rheumatol. 2010 Dec;6(12):683-92
references.pmid: 21060334
references.citation: Nature. 1988 Jan 28;331(6154):356-8
references.pmid: 2893286
references.citation: J Leukoc Biol. 1986 Jan;39(1):49-62
references.pmid: 3455711
references.citation: Nature. 2001 May 10;411(6834):207-11
references.pmid: 11346799
references.citation: Nat Genet. 2001 Jan;27(1):20-1
references.pmid: 11137993
references.citation: Neurology. 2003 Jul 8;61(1):108-10
references.pmid: 12847168
references.citation: Am J Hum Genet. 2000 Dec;67(6):1460-9
references.pmid: 11078476
references.citation: Curr Opin Rheumatol. 1993 Sep;5(5):575-86
references.pmid: 8398608
references.citation: Arthritis Rheum. 2005 Dec 15;53(6):845-9
references.pmid: 16342090
references.citation: J Immunol. 1985 Dec;135(6):3850-6
references.pmid: 3877758
references.citation: Adv Immunol. 1989;44:93-151
references.pmid: 2646863
references.citation: Dermatol Ther. 2012 Mar-Apr;25(2):99-111
references.pmid: 22741931
references.citation: Am J Med. 1975 Feb;58(2):243-64
references.pmid: 1115070
references.citation: J Rheumatol. 2000 Jan;27(1):135-41
references.pmid: 10648029
references.citation: J Pathol. 2001 Aug;194(4):398-405
references.pmid: 11523046
references.citation: Proc Natl Acad Sci U S A. 2003 Mar 4;100(5):2610-5
references.pmid: 12604793
references.citation: Arthritis Rheum. 2009 Jul;60(7):2094-101
references.pmid: 19565516
references.citation: Arch Pathol Lab Med. 2001 Apr;125(4):475-83
references.pmid: 11260619
references.citation: J Clin Rheumatol. 2007 Jun;13(3):140-2
references.pmid: 17551380
references.citation: Arthritis Rheum. 2002 Oct;46(10):2665-72
references.pmid: 12384925
references.citation: Br J Rheumatol. 1995 Sep;34(9):866-72
references.pmid: 7582729
references.citation: Immunity. 1998 Apr;8(4):497-508
references.pmid: 9586639
references.citation: Arthritis Rheum. 1998 Jul;41(7):1241-50
references.pmid: 9663482
references.citation: Proc Natl Acad Sci U S A. 1996 Aug 6;93(16):8563-8
references.pmid: 8710910
references.citation: Ann Rheum Dis. 2001 Apr;60(4):372-9
references.pmid: 11247868
references.citation: Arthritis Rheum. 2009 Aug;60(8):2390-401
references.pmid: 19644848
references.citation: J Pediatr. 1983 Dec;103(6):889-91
references.pmid: 6644424
references.citation: Arthritis Rheum. 1992 Mar;35(3):311-8
references.pmid: 1536669
references.citation: Arthritis Rheum. 2009 Jun;60(6):1733-42
references.pmid: 19479824
references.citation: Semin Arthritis Rheum. 1993 Aug;23(1):8-15
references.pmid: 8235667
references.citation: Nature. 2000 Dec 7;408(6813):740-5
references.pmid: 11130078
references.citation: Am J Clin Pathol. 1986 May;85(5):552-6
references.pmid: 3706197
references.citation: Curr Opin Immunol. 2006 Jun;18(3):349-56
references.pmid: 16616472
references.citation: J Intern Med. 2013 Jun;273(6):544-54
references.pmid: 23464352
references.citation: Am Rev Respir Dis. 1979 Mar;119(3):471-503
references.pmid: 36018
references.citation: Neuroimage. 2007 Jan 15;34(2):694-701
references.pmid: 17112740
references.citation: Nat Immunol. 2008 Feb;9(2):166-75
references.pmid: 18157131
references.citation: Arthritis Rheum. 1999 Oct;42(10):2026-38
references.pmid: 10524673
references.citation: Autoimmunity. 2010 Mar;43(2):179-88
references.pmid: 19835488
references.citation: Am J Med. 1981 Apr;70(4):875-81
references.pmid: 7211922
references.citation: J Invest Dermatol. 1998 Jul;111(1):149-53
references.pmid: 9665402
references.citation: J Exp Med. 1992 Sep 1;176(3):761-79
references.pmid: 1512540
references.citation: Cell. 1996 May 3;85(3):303-6
references.pmid: 8616885
references.citation: Am Heart J. 1985 Dec;110(6):1257-65
references.pmid: 3907317
references.citation: Br J Dermatol. 2007 Oct;157(4):752-7
references.pmid: 17714558
references.citation: Proc Natl Acad Sci U S A. 1994 Apr 26;91(9):4062-6
references.pmid: 8171035
references.citation: Proc Natl Acad Sci U S A. 1994 Oct 11;91(21):10168-72
references.pmid: 7937857
references.citation: J Immunol. 2004 Sep 1;173(5):3437-45
references.pmid: 15322209
references.citation: Clin Chest Med. 2010 Sep;31(3):479-88
references.pmid: 20692540
references.citation: Arthritis Rheum. 2006 Aug;54(8):2550-7
references.pmid: 16868977
references.citation: Nat Genet. 1998 May;19(1):56-9
references.pmid: 9590289
references.citation: Brain Behav Immun. 2011 Feb;25(2):279-85
references.pmid: 20884345
references.citation: Arthritis Rheum. 1995 Sep;38(9):1260-70
references.pmid: 7575721
references.citation: J Immunol. 2004 Jun 1;172(11):6568-77
references.pmid: 15153471
references.citation: Rheumatology (Oxford). 2010 Dec;49(12):2243-54
references.pmid: 20823093
references.citation: Arthritis Rheum. 2011 Mar;63(3):722-32
references.pmid: 21360502
references.citation: Semin Nephrol. 1999 Jan;19(1):48-56
references.pmid: 9952280
references.citation: Curr Top Microbiol Immunol. 2007;316:359-86
references.pmid: 17969456
references.citation: Arthritis Rheum. 2006 Dec 15;55(6):900-4
references.pmid: 17139667
references.citation: J Clin Invest. 1989 Aug;84(2):627-34
references.pmid: 2760204
references.citation: J Rheumatol. 2006 Nov;33(11):2184-91
references.pmid: 17086605
references.citation: J Exp Med. 1977 May 1;145(5):1115-30
references.pmid: 323400
references.citation: J Immunol. 1987 May 1;138(9):2883-9
references.pmid: 3553329
references.citation: Arthritis Rheum. 2006 Jul;54(7):2198-210
references.pmid: 16804897
references.citation: Semin Arthritis Rheum. 2004 Jun;33(6):414-21
references.pmid: 15190526
references.citation: Ann Rheum Dis. 2001 Feb;60(2):106-11
references.pmid: 11156541
references.citation: J Neuroimmunol. 2010 Apr 15;221(1-2):46-52
references.pmid: 20207017
references.citation: Eur J Neurol. 2004 Jan;11(1):37-42
references.pmid: 14692886
references.citation: Br Med J (Clin Res Ed). 1983 Jan 22;286(6361):251-2
references.pmid: 6402058
references.citation: J Immunol. 1977 Dec;119(6):2157-62
references.pmid: 303258
references.citation: Nat Genet. 2007 Sep;39(9):1065-7
references.pmid: 17660818
references.citation: Eur J Immunol. 2000 Oct;30(10):2782-90
references.pmid: 11069058
references.citation: J Exp Med. 1978 Jun 1;147(6):1653-60
references.pmid: 681876
references.citation: Nature. 1963 Jan 12;197:197
references.pmid: 13953664
references.citation: Arthritis Rheum. 1979 Mar;22(3):226-35
references.pmid: 311204
references.citation: J Hepatol. 2005 Jun;42(6):850-9
references.pmid: 15885356
references.citation: Ann Intern Med. 1973 May;78(5):786
references.pmid: 4711795
references.citation: Intern Med. 2007;46(13):1033-7
references.pmid: 17603247
references.citation: J Rheumatol. 1997 Dec;24(12):2314-7
references.pmid: 9415634
references.citation: Tissue Antigens. 2014 Jun;83(6):401-8
references.pmid: 24697319
references.citation: Genes Immun. 2011 Jun;12(4):270-9
references.pmid: 21270825
references.citation: Lab Invest. 2013 Jun;93(6):626-38
references.pmid: 23628898
references.citation: J Immunol. 2008 Feb 1;180(3):1938-47
references.pmid: 18209092
references.citation: Medicine (Baltimore). 2006 Mar;85(2):95-104
references.pmid: 16609348
references.citation: Clin Exp Rheumatol. 2005 Sep-Oct;23(5 Suppl 39):S120-32
references.pmid: 16273796
references.citation: Pol Arch Med Wewn. 2008 Jan-Feb;118(1-2):57-63
references.pmid: 18405174
references.citation: Immunity. 2012 Dec 14;37(6):1104-15
references.pmid: 23123062
references.citation: Immunity. 2006 Sep;25(3):383-92
references.pmid: 16979570
references.citation: Immunity. 1995 Nov;3(5):549-60
references.pmid: 7584145
references.citation: Curr Opin Rheumatol. 2013 Sep;25(5):561-8
references.pmid: 23917156
references.citation: Eur J Immunol. 1994 Mar;24(3):677-83
references.pmid: 7510240
references.citation: Clin Immunol. 2005 Nov;117(2):104-11
references.pmid: 16214080
references.citation: Clin Rheumatol. 2014 Jan;33(1):105-10
references.pmid: 24096639
references.citation: Arthritis Rheum. 2011 Jun;63(6):1689-97
references.pmid: 21425124
references.citation: Lupus. 2000;9(8):573-83
references.pmid: 11035431
references.citation: J Invest Dermatol. 2011 Jan;131(1):133-40
references.pmid: 20720564
references.citation: Brain Behav Immun. 2008 Aug;22(6):949-59
references.pmid: 18411022
references.citation: J Exp Med. 1987 May 1;165(5):1237-51
references.pmid: 3494806
references.citation: Ann N Y Acad Sci. 1997 Aug 14;823:256-62
references.pmid: 9292052
references.citation: Arthritis Rheum. 2007 Apr;56(4):1242-50
references.pmid: 17393453
references.citation: Arthritis Res Ther. 2009 Oct 14;11(5):R154
references.pmid: 19828047
references.citation: J Leukoc Biol. 2012 Sep;92(3):577-91
references.pmid: 22753952
references.citation: Case Rep Pulmonol. 2012;2012:836017
references.pmid: 22934226
references.citation: Medicine (Baltimore). 1954 Dec;33(4):291-437
references.pmid: 13223169
references.citation: J Immunol. 2003 Dec 1;171(11):5778-86
references.pmid: 14634086
references.citation: Arthritis Rheum. 2004 Feb;50(2):640-9
references.pmid: 14872509
references.citation: Hum Mol Genet. 2006 Nov 1;15(21):3195-205
references.pmid: 17000707
references.citation: Cell. 2008 Aug 22;134(4):569-71
references.pmid: 18724930
references.citation: Medicine (Baltimore). 1997 May;76(3):192-202
references.pmid: 9193454
references.citation: Autoimmun Rev. 2007 Jan;6(3):155-61
references.pmid: 17289551
references.citation: Nat Rev Rheumatol. 2011 Apr;7(4):225-33
references.pmid: 21386795
references.citation: J Clin Invest. 1994 Aug;94(2):568-77
references.pmid: 8040312
references.citation: Arthritis Rheum. 2012 Aug;64(8):2677-86
references.pmid: 22553077
references.citation: Clin Exp Immunol. 2009 Aug;157(2):209-15
references.pmid: 19604260
references.citation: J Biol Chem. 1999 Mar 5;274(10):6130-7
references.pmid: 10037696
references.citation: Ann Rheum Dis. 2007 Dec;66(12):1661-8
references.pmid: 17504842
references.citation: Proc Natl Acad Sci U S A. 1976 May;73(5):1646-50
references.pmid: 1064037
references.citation: J Immunol. 2006 Jun 15;176(12):7143-7
references.pmid: 16751355
references.citation: Lupus. 2006;15(9):600-5
references.pmid: 17080916
references.citation: Am J Cardiol. 1990 May 1;65(16):1145-8
references.pmid: 2330902
references.citation: N Engl J Med. 2008 Feb 28;358(9):929-39
references.pmid: 18305268
references.citation: Eur J Immunol. 1982 Feb;12(2):152-9
references.pmid: 6978818
references.citation: Scand J Immunol. 2010 May;71(5):317-28
references.pmid: 20500682
references.citation: Immunity. 2012 Apr 20;36(4):503-14
references.pmid: 22520844
references.citation: J Biomed Biotechnol. 2011;2011:271694
references.pmid: 21403825
references.citation: N Engl J Med. 1996 Nov 7;335(19):1424-30
references.pmid: 8875919
references.citation: Lupus. 2003;12(12):919-28
references.pmid: 14714912
references.citation: Arthritis Rheum. 2009 Apr;60(4):1076-84
references.pmid: 19333917
references.citation: Nat Genet. 2008 Feb;40(2):204-10
references.pmid: 18204446
references.citation: Arthritis Res Ther. 2006;8(1):R4
references.pmid: 16356190
references.citation: J Neuroimmunol. 2006 Dec;181(1-2):150-6
references.pmid: 17064784
references.citation: J Immunol. 1985 Feb;134(2):880-4
references.pmid: 3880792
references.citation: J Clin Lab Anal. 2008;22(4):271-7
references.pmid: 18634076
references.citation: J Immunol. 1996 Feb 1;156(3):1151-6
references.pmid: 8557992
references.citation: Nature. 2005 May 26;435(7041):452-8
references.pmid: 15917799
references.citation: Arthritis Rheum. 1980 Apr;23(4):392-403
references.pmid: 7370058
references.citation: J Biol Chem. 2002 Jul 26;277(30):27535-44
references.pmid: 11983693
references.citation: Adv Immunol. 2000;76:227-324
references.pmid: 11079100
references.citation: Mol Med. 2007 Jan-Feb;13(1-2):59-68
references.pmid: 17515957
references.citation: J Cell Biol. 1979 Apr;81(1):137-53
references.pmid: 90048
references.citation: Eur J Immunol. 1998 Sep;28(9):2928-35
references.pmid: 9754580
references.citation: Eur J Immunol. 1989 Jan;19(1):137-43
references.pmid: 2784103
references.citation: PLoS One. 2010 Jun 15;5(6):e11124
references.pmid: 20559547
references.citation: Clin Exp Dermatol. 2011 Jul;36(5):512-20
references.pmid: 21631571
references.citation: PLoS One. 2012;7(11):e49715
references.pmid: 23166752
references.citation: Cell. 1994 Feb 11;76(3):519-29
references.pmid: 8313472
references.citation: Nat Genet. 2001 Jan;27(1):18-20
references.pmid: 11137992
references.citation: J Exp Med. 2005 Jul 18;202(2):321-31
references.pmid: 16027240
references.citation: Chest. 1987 Jun;91(6):891-7
references.pmid: 3556057
references.citation: J Immunol. 1997 Oct 15;159(8):3838-48
references.pmid: 9378971
references.citation: Ann Rheum Dis. 2012 Jan;71(1):136-42
references.pmid: 21978998
references.citation: Arthritis Res. 2002;4 Suppl 3:S279-93
references.pmid: 12110148
references.citation: Arch Intern Med. 1981 Feb;141(2):201-3
references.pmid: 6970022
references.citation: Arthritis Rheum. 2007 Oct;56(10):3420-32
references.pmid: 17907141
references.citation: J Exp Med. 1981 Apr 1;153(4):897-909
references.pmid: 6972993
references.citation: Medicine (Baltimore). 1971 Mar;50(2):85-95
references.pmid: 4109481
references.citation: Immunity. 2006 Sep;25(3):417-28
references.pmid: 16973389
references.citation: Proc Natl Acad Sci U S A. 1998 Aug 4;95(16):9465-70
references.pmid: 9689103
references.citation: Ann Neurol. 2005 May;57(5):664-78
references.pmid: 15852401
references.citation: J Exp Med. 1994 Oct 1;180(4):1295-306
references.pmid: 7931063
references.citation: J Immunol. 2006 Nov 15;177(10):7287-95
references.pmid: 17082647
references.citation: Am J Pathol. 2006 Jun;168(6):1779-92
references.pmid: 16723695
references.citation: Curr Opin Pediatr. 1998 Dec;10(6):600-6
references.pmid: 9848020
references.citation: Expert Rev Clin Immunol. 2009 Jul;5(4):391-403
references.pmid: 20477036
references.citation: Eur J Intern Med. 2010 Jun;21(3):202-7
references.pmid: 20493423
references.citation: Arthritis Rheumatol. 2014 Mar;66(3):694-706
references.pmid: 24574230
references.citation: Clin Dermatol. 2004 Mar-Apr;22(2):148-56
references.pmid: 15234016
references.citation: Kidney Int. 2012 Jul;82(2):184-92
references.pmid: 22297676
references.citation: Semin Arthritis Rheum. 2004 Oct;34(2):501-37
references.pmid: 15505768
references.citation: Thromb Haemost. 1990 Apr 12;63(2):169-73
references.pmid: 2114042
references.citation: Mol Pharmacol. 2007 May;71(5):1427-34
references.pmid: 17322004
references.citation: Proc Natl Acad Sci U S A. 1996 Mar 19;93(6):2264-71
references.pmid: 8637861
references.citation: J Exp Med. 1957 Aug 1;106(2):191-202
references.pmid: 13449231
references.citation: Bull NYU Hosp Jt Dis. 2010;68(4):251-6
references.pmid: 21162701
references.citation: J Immunol. 1989 May 1;142(9):3076-82
references.pmid: 2785132
references.citation: J Clin Rheumatol. 2006 Feb;12(1):37-40
references.pmid: 16484880
references.citation: Arthritis Care Res (Hoboken). 2012 Jun;64(6):797-808
references.pmid: 22556106
references.citation: Arthritis Rheum. 2010 Mar;62(3):845-54
references.pmid: 20131231
references.citation: Arthritis Rheum. 1999 Sep;42(9):1785-96
references.pmid: 10513791
references.citation: Can Med Assoc J. 1974 Dec 21;111(12):1331-3
references.pmid: 4140754
references.citation: J Clin Invest. 1987 Sep;80(3):691-7
references.pmid: 3624484
references.citation: Immunity. 1994 Jun;1(3):219-29
references.pmid: 7889410
references.citation: Semin Arthritis Rheum. 2012 Oct;42(2):179-85
references.pmid: 22595642
references.citation: Semin Immunopathol. 2013 Jul;35(4):465-80
references.pmid: 23595413
references.citation: J Autoimmun. 2010 May;34(3):J247-57
references.pmid: 20018479
references.citation: Clin Exp Immunol. 2004 Oct;138(1):171-8
references.pmid: 15373921
references.citation: Int Immunol. 2000 Nov;12(11):1569-78
references.pmid: 11058577
references.citation: Arthritis Rheum. 1997 Sep;40(9):1725
references.pmid: 9324032
references.citation: Autoimmunity. 2004 Mar;37(2):85-93
references.pmid: 15293878
references.citation: Angiology. 1985 Jul;36(7):431-41
references.pmid: 4025948
references.citation: Lupus. 1996 Jun;5(3):196-205
references.pmid: 8803890
references.citation: J Clin Invest. 1997 Dec 15;100(12):3019-26
references.pmid: 9399948
references.citation: J Immunol. 2006 Dec 1;177(11):8258-65
references.pmid: 17114504
references.citation: Arthritis Rheum. 1992 Aug;35(8):953-9
references.pmid: 1642661
references.citation: Br Med J. 1979 May 12;1(6173):1235-8
references.pmid: 455010
references.citation: FASEB J. 2010 Jun;24(6):1682-8
references.pmid: 20065106
references.citation: Immunity. 2000 Aug;13(2):277-85
references.pmid: 10981970
references.citation: Immunity. 2005 Jul;23(1):41-51
references.pmid: 16039578
references.citation: N Engl J Med. 2008 Feb 28;358(9):900-9
references.pmid: 18204098
references.citation: Clin Immunol Immunopathol. 1983 Jun;27(3):387-402
references.pmid: 6191905
references.citation: J Immunol. 1987 Nov 15;139(10):3287-94
references.pmid: 3500215
references.citation: Lupus. 2005;14(10):827-31
references.pmid: 16302678
references.citation: J Immunol. 1997 May 15;158(10):4525-8
references.pmid: 9144462
references.citation: Lupus. 2000;9(7):498-503
references.pmid: 11035414
references.citation: Arthritis Rheum. 1982 Nov;25(11):1271-7
references.pmid: 7138600
references.citation: Nat Genet. 1997 Dec;17(4):399-403
references.pmid: 9398840
references.citation: Immunol Rev. 2010 Sep;237(1):72-89
references.pmid: 20727030
references.citation: Curr Opin Rheumatol. 2008 Sep;20(5):589-94
references.pmid: 18698183
references.citation: Clin Exp Immunol. 1992 Oct;90(1):56-62
references.pmid: 1395101
references.citation: Genet Epidemiol. 2011 Jul;35(5):310-7
references.pmid: 21374718
references.citation: Nature. 2002 Apr 11;416(6881):603-7
references.pmid: 11948342
references.citation: Nat Genet. 2000 Jun;25(2):177-81
references.pmid: 10835632
references.citation: J Exp Med. 1967 Oct 1;126(4):607-24
references.pmid: 4168098
references.citation: PLoS Genet. 2011 Mar;7(3):e1001323
references.pmid: 21408207
references.citation: Rheumatology (Oxford). 2007 Mar;46(3):467-72
references.pmid: 16901949
references.citation: J Exp Med. 1999 Apr 19;189(8):1307-13
references.pmid: 10209047
references.citation: J Exp Med. 1978 Nov 1;148(5):1198-215
references.pmid: 309911
references.citation: PLoS Genet. 2010 Feb 12;6(2):e1000841
references.pmid: 20169177
references.citation: Genes Immun. 2009 Jun;10(4):285-96
references.pmid: 19404300
references.citation: Lupus. 2010 Oct;19(12):1374-83
references.pmid: 20947545
references.citation: Sci Transl Med. 2012 Oct 24;4(157):157ra141
references.pmid: 23100627
references.citation: PLoS One. 2013;8(2):e57082
references.pmid: 23451151
references.citation: Br J Rheumatol. 1996 Mar;35(3):248-54
references.pmid: 8620300
references.citation: Adv Immunol. 1968;9:215-66
references.pmid: 4883742
references.citation: J Rheumatol. 2007 Oct;34(10):2010-7
references.pmid: 17896804
references.citation: Nat Rev Immunol. 2005 May;5(5):375-86
references.pmid: 15864272
references.citation: Chest. 2000 Oct;118(4):1083-90
references.pmid: 11035681
references.citation: J Exp Med. 2009 Mar 16;206(3):561-76
references.pmid: 19221396
references.citation: Rheumatology (Oxford). 2011 Sep;50(9):1578-85
references.pmid: 21208977
references.citation: Immunity. 2001 Sep;15(3):397-408
references.pmid: 11567630
references.citation: Am J Med. 1980 Dec;69(6):849-58
references.pmid: 7004181
references.citation: Am J Pathol. 1965 Oct;47(4):677-94
references.pmid: 5837733
references.citation: Genome Res. 2010 Feb;20(2):170-9
references.pmid: 20028698
references.citation: Proc Natl Acad Sci U S A. 2006 Dec 26;103(52):19854-9
references.pmid: 17170137
references.citation: J Neurosci Res. 2001 Apr 1;64(1):26-33
references.pmid: 11276048
references.citation: Proc Natl Acad Sci U S A. 2009 Dec 8;106(49):20865-70
references.pmid: 19923437
references.citation: J Biol Chem. 1999 Jul 9;274(28):19655-60
references.pmid: 10391904
references.citation: Int Immunol. 2001 Dec;13(12):1461-9
references.pmid: 11717187
references.citation: Circulation. 1990 Aug;82(2):369-75
references.pmid: 2372888
references.citation: J Immunol. 2005 Dec 15;175(12):8312-9
references.pmid: 16339572
references.citation: Stroke. 1985 May-Jun;16(3):512-4
references.pmid: 4002268
references.citation: Am J Med. 1979 Feb;66(2):210-5
references.pmid: 218447
references.citation: J Exp Med. 1990 Feb 1;171(2):503-17
references.pmid: 2303783
references.citation: Immunogenetics. 1992;35(3):214-6
references.pmid: 1537612
references.citation: Nat Rev Genet. 2009 May;10(5):285-90
references.pmid: 19337289
references.citation: Arthritis Rheum. 2003 Oct;48(10):2881-7
references.pmid: 14558094
references.citation: J Immunol. 2008 Apr 1;180(7):5101-8
references.pmid: 18354236
references.citation: Ann Rheum Dis. 2000 Mar;59(3):217-22
references.pmid: 10700431
references.citation: Arthritis Rheum. 2005 Nov;52(11):3639-45
references.pmid: 16255056
references.citation: J Immunol. 2005 Mar 1;174(5):3041-50
references.pmid: 15728518
references.citation: Arthritis Rheum. 2008 Jan;58(1):15-25
references.pmid: 18163481
references.citation: Nat Rev Rheumatol. 2012 Feb 14;8(4):214-23
references.pmid: 22331061
references.citation: Immunity. 1997 Jul;7(1):69-81
references.pmid: 9252121
references.citation: Science. 1998 Feb 13;279(5353):1052-4
references.pmid: 9461440
references.citation: J Exp Med. 1987 Sep 1;166(3):798-803
references.pmid: 3114409
references.citation: J Rheumatol. 2009 Feb;36(2):416-21
references.pmid: 19208567
references.citation: Genes Immun. 2004 Sep;5(6):484-92
references.pmid: 15295621
references.citation: Nature. 2005 Jul 7;436(7047):138-41
references.pmid: 16001076
references.citation: Immunity. 2004 Aug;21(2):179-88
references.pmid: 15308099
references.citation: J Exp Med. 1990 Jan 1;171(1):265-92
references.pmid: 2104919
references.citation: J Am Chem Soc. 2009 Oct 21;131(41):15005-13
references.pmid: 19778017
references.citation: Proc Natl Acad Sci U S A. 2006 Jan 17;103(3):678-83
references.pmid: 16407105
references.citation: J Biomed Biotechnol. 2012;2012:798924
references.pmid: 22654485
references.citation: J Exp Med. 1976 Aug 1;144(2):428-43
references.pmid: 8578
references.citation: Nature. 1997 Sep 18;389(6648):251-60
references.pmid: 9305837
references.citation: Kidney Int. 2006 May;69(10):1846-51
references.pmid: 16598205
references.citation: Arthritis Rheum. 2004 Mar;50(3):858-65
references.pmid: 15022328
references.citation: Curr Opin Rheumatol. 2003 Sep;15(5):548-56
references.pmid: 12960479
references.citation: Adv Nephrol Necker Hosp. 1976;6:47-61
references.pmid: 139082
references.citation: J Exp Med. 1999 May 17;189(10):1639-48
references.pmid: 10330443
references.citation: Immunol Rev. 2012 Sep;249(1):158-75
references.pmid: 22889221
references.citation: Clin Immunol. 2013 Jan;146(1):1-9
references.pmid: 23159786
references.citation: J Clin Invest. 1967 Apr;46(4):569-79
references.pmid: 4164260
references.citation: Curr Dir Autoimmun. 2003;6:123-53
references.pmid: 12408050
references.citation: Ann Rheum Dis. 2012 Dec;71(12):2028-34
references.pmid: 22586164
references.citation: J Womens Health (Larchmt). 2004 Jul-Aug;13(6):713-8
references.pmid: 15333286
references.citation: J Immunol. 1999 Apr 15;162(8):4406-16
references.pmid: 10201976
references.citation: Nature. 1964 Sep 5;203:1080-1
references.pmid: 14223091
references.citation: Kidney Int. 1994 Dec;46(6):1627-34
references.pmid: 7700021
references.citation: J Rheumatol. 1985 Oct;12(5):919-23
references.pmid: 3878879
references.citation: Cell. 1995 Oct 20;83(2):301-11
references.pmid: 7585947
references.citation: Arthritis Rheum. 2012 Feb;64(2):485-92
references.pmid: 21952918
references.citation: Arq Bras Cardiol. 2000 Nov;75(5):446-8
references.pmid: 11080755
references.citation: Rheumatology (Oxford). 2007 Jul;46(7):1052-6
references.pmid: 17500073
references.citation: Immunobiology. 2002 Sep;205(4-5):395-406
references.pmid: 12396002
references.citation: Kidney Int. 2007 Apr;71(7):664-72
references.pmid: 17332738
references.citation: Diabetes. 2012 May;61(5):1281-90
references.pmid: 22403299
references.citation: Proc Natl Acad Sci U S A. 2007 Apr 24;104(17):7193-8
references.pmid: 17409188
references.citation: PLoS One. 2012;7(12):e51090
references.pmid: 23236436
references.citation: Annu Rev Immunol. 2007;25:619-47
references.pmid: 17201681
references.citation: Nephrol Dial Transplant. 2008 Jan;23(1):385-6
references.pmid: 17933839
references.citation: Semin Arthritis Rheum. 2014 Jun;43(6):759-66
references.pmid: 24332116
references.citation: Am J Med. 1981 Nov;71(5):791-8
references.pmid: 7304651
references.citation: Medicine (Baltimore). 1992 Jul;71(4):216-23
references.pmid: 1518395
references.citation: Ann Rheum Dis. 2008 Jan;67(1):11-18
references.pmid: 17502354
references.citation: Arthritis Rheum. 2002 Jan;46(1):191-201
references.pmid: 11817590
references.citation: Lupus. 1997;6(2):181-92
references.pmid: 9061667
references.citation: Lab Invest. 1993 Apr;68(4):419-26
references.pmid: 8479150
references.citation: Rev Bras Reumatol. 2010 Sep-Oct;50(5):539-51
references.pmid: 21125190
references.citation: Nat Genet. 2000 Jun;25(2):135-6
references.pmid: 10835621
references.citation: Arthritis Rheum. 2005 Apr;52(4):1138-47
references.pmid: 15818688
references.citation: Nephrol Dial Transplant. 2006 Nov;21(11):3062-73
references.pmid: 16954173
references.citation: Nat Clin Pract Rheumatol. 2009 Feb;5(2):106-14
references.pmid: 19182817
references.citation: Nat Genet. 2008 Jan;40(1):83-9
references.pmid: 18059267
references.citation: Br J Dermatol. 2012 May;166(5):971-5
references.pmid: 22242767
references.citation: Am J Med. 1975 Oct;59(4):533-52
references.pmid: 1101680
references.citation: Semin Immunol. 2007 Feb;19(1):11-23
references.pmid: 17276080
references.citation: Curr Opin Immunol. 2012 Oct;24(5):530-7
references.pmid: 22889593
references.citation: Arthritis Rheum. 2011 Aug;63(8):2407-15
references.pmid: 21506093
references.citation: Medicine (Baltimore). 1975 Sep;54(5):397-409
references.pmid: 125838
references.citation: Arthritis Rheum. 1997 Oct;40(10):1810-6
references.pmid: 9336415
references.citation: Arthritis Care Res (Hoboken). 2012 Dec;64(12):1928-33
references.pmid: 22933330
references.citation: Autoimmun Rev. 2009 Sep;9(1):45-8
references.pmid: 19285156
references.citation: Curr Opin Rheumatol. 2004 Sep;16(5):527-33
references.pmid: 15314489