Genetics of experimental autoimmune encephalomyelitis in the mouse

Archivum Immunologiae et Therapiae Experimentalis

Volumn 52, Issue 5, 2004, Pages 316-325

  Andersson, Asa ^a,b   Karlsson, Jenny ^a  

^a LUND UNIVERSITY   (Sweden)

^b LUND UNIVERSITY   (Sweden)

Author keywords

Autoimmunity; EAE; Experimental models; Genetics; Multiple sclerosis

Indexed keywords

MYELIN PROTEIN;

ALLERGIC ENCEPHALOMYELITIS; ANIMAL EXPERIMENT; ANIMAL MODEL; GENE IDENTIFICATION; GENETIC PREDISPOSITION; GENETIC SUSCEPTIBILITY; GENETICS; HUMAN; IMMUNIZATION; MOUSE; MULTIPLE SCLEROSIS; NONHUMAN; QUANTITATIVE TRAIT LOCUS; REVIEW;

ANIMALS; ENCEPHALOMYELITIS, AUTOIMMUNE, EXPERIMENTAL; GENETIC PREDISPOSITION TO DISEASE; LINKAGE (GENETICS); MICE; QUANTITATIVE TRAIT LOCI;

EID: 10344255672     PISSN: 0004069X     EISSN: None     Source Type: Journal    
DOI: None     Document Type: Review

References (77)

1. Anderson A. C., Nicholson L. B., Legge K. L., Turchin V., Zaghouani H. and Kuchroo V. K. (2000): High frequency of autoreactive myelin proteolipid protein-specific T cells in the periphery of naive mice: mechanisms of selection of the self-reactive repertoire. J. Exp. Med., 191, 761-770.
2. Andersson E. C., Hansen B. E., Jacobsen H., Madsen L. S., Andersen C. B., Engberg J., Rothbard J. B., McDevitt G. S., Malmstrom V., Holmdahl R., Svejgaard A. and Fugger L. (1998): Definition of MHC and T cell receptor contacts in the HLA-DR4-restricted immunodominant epitope in type II collagen and characterization of collagen-induced arthritis in HLA-DR4 and human CD4 transgenic mice. Proc. Natl. Acad. Sci. USA, 95, 7574-7579.
3. Baker D., Rosenwasser O. A., O'Neill J. K. and Turk J. L. (1995): Genetic analysis of experimental allergic encephalomyelitis in mice. J. Immunol., 155, 4046-4051.
4. Barclay A. N., Wright G. J., Brooke G. and Brown M. H. (2002): CD200 and membrane protein interactions in the control of myeloid cells. Trends Immunol., 23, 285-290.
5. Bergsteinsdottir K., Yang H.-T., Pettersson U. and Holmdahl R. (2000): Evidence for common autoimmune disease genes controlling onset, severity, and chronicity based on experimental models for multiple sclerosis and rheumatoid arthritis. J. Immunol., 164, 1564-1568.
6. Blankenhorn E. P., Butterfield R. J., Rigby R., Cort L., Giambrone D., McDermott P., McEntee K., Solowski N., Meeker N. D., Zachary J. F., Doerge R. W. and Teuscher C. (2000): Genetic analysis of the influence of pertussis toxin on experimental allergic encephalomyelitis susceptibility: an environmental agent can override genetic checkpoints. J. Immunol., 164, 3420-3425.
7. Blom T., Franzen A., Heinegard D. and Holmdahl R. (2003): Comment on "The influence of the proinflammatory cytokine, osteopontin, on autoimmune demyelinating disease". Science, 299, 1845.
8. Boylan K. B., Takahashi N., Paty D. W., Sadownick A. D., Diamond M. and Hood L. E. (1990): DNA length polymorphism 5′ to the myelin basic protein gene is associated with multiple sclerosis. Ann. Neurol., 27, 291-297.
9. Butterfield R. J., Blankenhorn E. P., Roper R. J., Zachary J. F., Doerge R. W., Sudweeks J., Rose J. and Teuscher C. (1999): Genetic analysis of disease subtypes and sexual dimorphisms in mouse experimental allergic encephalomyelitis (EAE): relapsing/remitting and monophasic remitting/nonrelapsing EAE are immunogenetically distinct. J. Immunol., 162, 3096-3102.
10. Butterfield R. J., Blankenhorn E. P., Roper R. J., Zachary J. F., Doerge R. W. and Teuscher C. (2000): Identification of genetic loci controlling the characteristics and severity of brain and spinal cord lesions in experimental allergic encephalomyelitis. Am. J. Pathol., 157, 637-645.
11. Butterfield R. J., Sudweeks J. D., Blankenhorn E. P., Korngold R., Marini J. C., Todd J. A., Roper R. J. and Teuscher C. (1998): New genetic loci that control susceptibility and symptoms of experimental allergic encephalomyelitis in inbred mice. J. Immunol., 161, 1860-1867.
12. Chabas D., Baranzini S. E., Mitchell D., Bernard C. C., Rittling S. R., Denhardt D. T., Sobel R. A., Lock C., Karpuj M., Pedotti R., Heller R., Oksenberg J. R. and Steinman L. (2001): The influence of the proinflammatory cytokine, osteopontin, on autoimmune demyelinating disease. Science, 294, 1731-1735.
13. Compston A. and Coles A. (2002): Multiple sclerosis. Lancet, 359, 1221-1231.
14. Ebers G. C., Kukay K., Bulman D. E., Sadovnick A. D., Rice G., Anderson C., Armstrong H., Cousin K., Bell R. B., Hader W., Paty D. W., Hashimoto S., Oger J., Duquette P., Warren S., Gray T., O'Connor P., Nath A., Auty A., Metz L., Francis G., Paulseth J. E., Murray T. J., Pryse-Phillips W., Nelson R., Freedman M., Brunet D., Bouchard J.-P., Hinds D. and Risch N. (1996): A full genome screen in multiple sclerosis. Nat. Genet., 13, 472-476.
15. Encinas J. A., Lees M. B., Sobel R. A., Symonowics C., Weiner H. L., Seidman C. E., Seidman J. G. and Kuchroo V. J. (2001): Identification of genetic loci associated with paralysis, inflammation and weight loss in mouse experimental autoimmune encephalomyelitis. Int. Immunol., 13, 257-264.
16. Encinas J. A., Lees M. B., Sobel R. A., Symonowicz C., Greer J. M., Shovlin C. L., Weiner H. L., Seidman C. E., Seidman J. G. and Kuchroo V. K. (1996): Genetic analysis of susceptibility to experimental autoimmune encephalomyelitis in a cross between SJL/L and B10.S mice. J. Immunol., 157, 2186-2192.
17. Fritz R. B., Skeen M. J., Chou C. H., Garcia M. and Egorov I. K. (1985): Major histocompatibility complex-linked control of the murine immune response to myelin basic protein. J. Immunol., 134, 2328-2332.
18. Gebe J. A., Swanson E. and Kwok W. W. (2002): HLA class II peptide-binding and autoimmunity. Tissue Antigens, 59, 78-87.
19. Guerini F. R., Ferrante P., Losciale L., Caputo D., Lombardi M. L., Pirozzi G., Luongo V., Sudomoina M. A., Andreewski T. V., Alekseenkov A. D., Boiko A. N., Gusev E. I. and Favorova O. O. (2003): Myelin basic protein gene is associated with MS in DR4- and DR5-positive Italians and Russians. Neurology, 61, 520-526.
20. Haines J. L., Ter-Minassian M., Bazyk A., Gusella J. F., Kim D. J., Terwedow H., Pericak-Vance M. A., Rimmler J. B., Haynes C. S., Roses A. D., Lee A., Shaner B., Menold M., Seboun E., Fitoussi R. P., Gartioux C., Reyes C., Ribierre F., Gyapay G., Weissenbach J., Hauser S. L., Goodkin D. E., Lincoln R., Usuku K., Oksenberg J. R. et al. (1996): A complete genomic screen for multiple sclerosis underscores a role for the major histocompatability complex. The Multiple Sclerosis Genetics Group (see comments). Nat. Genet., 13, 469-471.
21. Haines J. L., Terwedow H. A., Burgess K., Pericak-Vance M. A., Rimmler J. B., Martin E. R., Oksenberg J. R., Lincoln R., Zhang D. Y., Banatao D. R., Gatto N., Goodkin D. E. and Hauser S. L. (1998): Linkage of the MHC to familial multiple sclerosis suggests genetic heterogeneity. The Multiple Sclerosis Genetics Group. Hum. Mol. Genet., 7, 1229-1234.
22. Haywood M. E., Hogarth M. B., Slingsby J. H., Rose S. J., Allen P. J., Thompson E. M., Maibaum M. A., Chandler P., Davies K. A., Simpson E., Walpor M. J. and Morley B. J. (2000): Identification of intervals on chromosomes 1, 3, and 13 linked to the development of lupus in BXSB mice. Arthritis Rheum., 43, 349-355.
23. Hemmer B., Archelos J. J. and Hartung H.-P. (2002): New concepts in the immunopathogenesis of multiple sclerosis. Nat. Rev. Neurosci., 3, 291-300.
24. Hockertz M. K., Paty D. W. and Beall S. S. (1998): Susceptibility to relapsing-progressive multiple sclerosis is associated with inheritance of genes linked to the variable region of the TcRβ locus: use of affected family-based controls. Am. J. Hum. Genet., 62, 373.
25. Ibrahim SM M. E., Bottcher T., Koczan D., Gold R., Rolfs A. and Thiesen H. J. (2001): Gene expression profiling of the nervous system in murine experimental autoimmune encephalomyelitis. Brain, 124, 1927-1938.
26. Jabs C., Greve B., Chang T. T., Sobel R. A., Sharpe A. H. and Kuchroo V. K. (2002): Genetic background determines the requirement for B7 costimulation in induction of autoimmunity. Eur. J. Immunol., 32, 2687-2697.
27. Jacobsen D. L., Gange S. J., Rose N. R. and Graham N. M. H. (1997): Epidemiology and estimated population burden of selected autoimmune diseases in the United States. Clin. Immunol. Immunopathol., 84, 223-243.
28. Jagodic M., Kornek B., Weissert R., Lassmann H., Olsson T., and Dahlman I. (2001): Congenic mapping confirms a locus on rat chromosome 10 conferring strong protection against myelin oligodendrocyte glycoprotein-induced experimental autoimmune encephalomyelitis. Immunogenetics, 53, 410-415.
29. Jansson L., Olsson T., Hojeberg B. and Holmdahl R. (1991): Chronic experimental autoimmune encephalomyelitis induced by the 89-101 myelin basic protein peptide in B10RIII (H-2r) mice. Eur. J. Immunol., 21, 693-699.
30. Jansson M., Panoutsakopoulou V., Baker J., Klein L. and Cantor H. (2002): Attenuated experimental autoimmune encephalomyelitis in eta-1/osteopontin- deficient mice. J. Immunol., 168, 2096-2099.
31. Ji H., Gauguier D., Ohmura K., Gonzalez A., Duchatelle V., Danoy P., Garchon H. J., Degott C., Lathrop M., Benoist C. and Mathis D. (2001): Genetic influences on the end-stage effector phase of arthritis. J. Exp. Med., 194, 321-330.
32. Jirholt J., Lindqvist A. K., Karlsson J., Andersson Å. and Holmdahl R. (2002): Identification of susceptibility genes for experimental autoimmune encephalomyelitis that overcome the effect of protective alleles at the eae2 locus. Int. Immunol., 14, 79-85.
33. Johansson A. C., Sundler M., Kjellen P., Johannesson M., Cook A., Lindqvist A. K., Nakken B., Bolstad A. I., Jonsson R., Alarcon-Riquelme M. and Holmdahl R. (2001): Genetic control of collagen-induced arthritis in a cross with NOD and C57BL/10 mice is dependent on gene regions encoding complement factor 5 and FcγRIIb and is not associated with loci controlling diabetes. Eur. J. Immunol., 31, 1847-1856.
34. Karlsson J., Zhao X., Lonskaya I., Neptin M., Holmdahl R. and Andersson Å. (2003): Novel quantitative trait loci controlling development of experimental autoimmune encephalomyelitis and proportion of lymphocyte subpopulations. J. Immunol., 170, 1019-1026.
35. Keegan B. M. and Noseworthy J. H. (2002): Multiple sclerosis. Ann. Rev. Med., 53, 285-302.
36. Kornek B. and Laassmann H. (2003): Neuropathology of multiple sclerosis - new concepts. Brain Res. Bull., 61, 321-326.
37. Korngold R., Feldman A., Rorke L. B., Lublin F. D. and Doherty P. C. (1986): Acute experimental allergic encephalomyelitis in radiation bone marrow chimeras between high and low susceptible strains of mice. Immunogenetics, 24, 309-315.
38. Kuchroo V. K., Anderson A. C., Waldner H., Munder M., Bettelli E. and Nicholson B. (2002): T cell response in experimental autoimmune encephalomyelitis (EAE): role of self and cross-reactive antigens in shaping, tuning, and regulating the autopathogenic T-cell repertoire. Annu. Rev. Immunol., 20, 101-123.
39. Kuokkanen S., Sundvall M., Terwilliger J. D., Tienari P. J., Wikström J., Holmdahl R., Pettersson U. and Peltonen L. (1996): A putative vulnerability locus to multiple sclerosis maps to 5p14-p12 in a region syntenic to the murine locus Eae2. Nat. Genet., 13, 477-480.
40. Lang H. L. E., Jacobsen H., Ikemizu S., Andersson C., Harlos K., Madsen L., Hjorth P., Sondergaard L., Svejgaard A., Wucherpfennig K., Stuart D. I., Bell J. I., Jones E. Y. and Fugger L. (2002): A functional and structural basis for TCR cross-reactivity in multiple sclerosis. Nat. Immunol., 3, 940-943.
41. Lock C., Hermans G., Pedotti R., Brendolan A., Schadt E., Garren H., Langer-Gould A., Strober S., Cannella B., Allard J., Klonowski P., Austin A., Lad N., Kaminski N., Galli S. J., Oksenberg J. R., Raine C. S., Heller R. and Steinman L. (2002): Gene-microarray analysis of multiple sclerosis lesions yields new targets validated in autoimmune encephalomyelitis. Nat. Med., 8, 500-508.
42. Ma R. Z., Gao J., Meeker N. D., Fillmore P. D., Tung K. S., Watanabe T., Zachary J. F., Offner H., Blankenhorn E. P. and Teuscher C. (2002): Identification of Bphs, an autoimmune disease locus, as histamine receptor H1. Science, 297, 620-623.
43. Madsen L. S., Andersson E. C., Jansson L., Krogsgaard M., Andersen C. B., Engberg J., Strominger J. L., SVejgaard A., Hjorth J. P., Holmdahl R., Wucherpfennig K. W. and Fugger L. (1999): A humanized model for multiple sclerosis using HLA-DR2 and a human T-cell receptor. Nat. Genet., 23, 343-347.
44. Markovic-Plese S., Fukaura H., Zhang J., Al-Sabbagh A., Southwood S., Sette A., Kuchroo V. K. and Hafler D. A. (1995): T cell recognition of immunodominant and cryptic proteolipid protein epitopes in humans. J. Immunol., 155, 982-992.
45. Marrack P., Kappler J. and Kotzin B. L. (2001): Autoimmune disease: why and where it occurs. Nat. Med., 7, 899-905.
46. Marrosu M. G., Murru M. R., Costa G., Murru R., Muntoni F. and Cucca F. (1998): DRB1-DQA1-DQB1 loci and multiple sclerosis predisposition in the Sardinian population. Hum. Mol. Genet., 7, 1235-1237.
47. Matejuk A., Hopke C., Dwyer J., Subramanian S., Jones R. E., Bourdette D. N., Vandenbark A. A. and Offner H. (2003): CNS gene expression pattern associated with spontaneous experimental autoimmune encephalomyclitis. J. Neurosci. Res., 73, 667-678.
48. Meeker N. D., Hickey W. F., Korngold R., Hansen W. K., Sudweeks J. D., Wardell B. B., Griffith J. S. and Teuscher C. (1995): Multiple loci govern the bone marrow-derived immunoregulatory mechanism controlling dominant resistance to autoimmune orchitis. Proc. Natl. Acad. Sci. USA, 92, 5684-5688.
49. Mitchell D. A., Pickering M. C., Warren J., Fossati-Jimack L., Cortes-Hernandez J., Cook H. T., Botto M. and Walport M. J. (2002): C1q deficiency and autoimmunity: the effects of genetic background on disease expression. J. Immunol., 168, 2538-2543.
50. Morel L., Blenman K. R., Croker B. P. and Wakeland E. K. (2001): The major murine systemic lupus erythematosus susceptibility locus, Sle1, is a cluster of functionally related genes. Proc. Natl. Acad. Sci. USA, 98, 1787-1792.
51. Morel L., Mohan C., Yu Y., Schiffenbauer J., Rudofsky U. H., Tian N., Longmate J. A. and Wakeland E. K. (1999): Multiplex inheritance of component phenotypes in a murine model of lupus. Mamm. Genome, 10, 176-181.
52. Murata K., Nose M., Ndhlovu L. C., Sato T., Sugamura K. and Ishii N. (2002): Constitutive OX40/OX40 ligand interaction induces autoimmune-like diseases. J. Immunol., 169, 4628-4636.
53. Olofsson P., Holmberg J., Tordsson J., Lu S., Akerstrom B. and Holmdahl R. (2003): Positional identification of Ncf1 as a gene that regulates arthritis severity in rats. Nat. Genet., 33, 25-32.
54. Otto J. M., Cs-Szabo G., Gallagher J., Velins S., Mikecz K., Buzas E. I., Enders J. T., Li Y., Olsen B. R. and Glant T. T. (1999): Identification of multiple loci linked to inflammation and autoantibody production by a genome scan of a murine model of rheumatoid arthritis. Arthritis Rheum., 42, 2524-2531.
55. Pedotti R., Devoss J. J., Youssef S., Mitchell D., Wedemeyer J., Madanat R., Garren H., Fontoura P., Tsai M., Galli S. J., Sobel R. A. and Steinman L. (2003): Multiple elements of the allergic arm of the immune response modulate autoimmune demyelination. Proc. Natt. Acad. Sci. USA, 100, 1867-1872.
56. Pihlaja H., Rantamaki T., Wikstrom J., Sumelahti M. L., Laaksonen M., Ilonen J., Ruutiainen J., Pirttila T., Elovaara I., Reunanen M., Kuokkanen S., Peltonen L., Koivisto K. and Tienari P. J. (2003): Linkage disequilibrium between the MBP tetranucleotide repeat and multiple sclerosis is restricted to a geographically defined subpopulation in Finland. Genes Immun., 4, 138-146.
57. Pugliatti M., Sotgiu S. and Rosati G. (2002): The worldwide prevalence of multiple sclerosis. Clin. Neurol. Neurosurg., 104, 182-191.
58. Roth M.-P., Viratelle C., Dolbois L., Delverdier M., Borot N., Pelletier L., Druet P., Clanet M. and Coppin H, (1999): A genome-wide search identifies two susceptibility loci for experimental autoimmune encephalomyelitis on rat chromosomes 4 and 10. J. Immunol., 162, 1917-1922.
59. Saarela J., Schoenberg Fejzo M., Chen D., Finnila S., Parkkonen M., Kuokkanen S., Sobel E., Tienari P. J., Sumelahti M. L., Wikstrom J., Elovaara I., Koivisto K., Pirttila T., Reunanen M., Palotie A. and Peltonen L. (2002): Fine mapping of a multiple sclerosis locus to 2.5 Mb on chromosome 17q22-q24. Hum. Mol. Genet., 11, 2257-2267.
60. Sawcer S., Jones H. B., Feakes R., Gray J., Smaldon N., Chataway J., Robertson N., Clayton D., Goodfellow P. N. and Compston A. (1996): A genome screen in multiple sclerosis reveals susceptibility loci on chromosome 6p21 and 17q22 (see comments). Nat. Genet., 13, 464-468.
61. Steinman L. (1999): Assessment of animal models for MS and demyelinating disease in the design of rational therapy. Neuron, 24, 511-514.
62. Steinman L. (2001): Multiple sclerosis: a two-stage disease. Nat. Immunol., 2, 762-764.
63. Steinman L. (2003): Optic neuritis, a variant of experimental encephalomyelitis, a durable model for all seasons, now in its seventieth year. J. Exp. Med., 197, 1065-1071.
64. Sudweeks J. D., Todd J. A., Blankenhorn E. P., Wardell B. B., Woodward S. R., Meeker N. D., Estes S. S. and Teuscher C. (1993): Locus controlling Bordetella pertussis-induced histamine sensitization (Bphs), an autoimmune disease-susceptibility gene, maps distal to T-cell receptor β-chain gene on mouse chromosome 6. Proc. Natl. Acad. Sci. USA., 90, 3700-3704.
65. Sundvall M., Jirholt J., Yang H. T., Jansson L., Engstrom A., Pettersson U. and Holmdahl R. (1995): Identification of murine loci associated with susceptibility to chronic experimental autoimmune encephalomyelitis. Nat. Genet., 10, 313-317.
66. Teuscher C., Butterfield R. J., Ma R. Z., Zachary J. F., Doerge R. W. and Blankenhorn E. P. (1999): Sequence polymorphism in the chemokines Scya1 (TCA-3), Scya2 (monocyte chemoattractant protein (MCP)-1), and Scya12 (NCP-5) are candidates for eae7, a locus controlling susceptibility to monophasic remitting/nonrelapsing experimental allergic encephalomyelitis. J. Immunol., 163, 2262-2266.
67. Tienari P. J., Wikström J., Sajantila A., Palo J. and Peltonen L. (1992): Genetic susceptibility to multiple sclerosis linked to myelin basic protein gene. Lancet, 340, 987-991.
68. Todd J. A., Aitman T. J., Cornall R. J., Ghosh S., Hall J. R., Hearne C. M., Knight A. M., Love J. M., Mcaleer M. A., Prins J. B., Rodrigues N., Lathrop M., Pressey A., Delarato N. H., Peterson L. B. and Wicker L. S. (1991): Genetic analysis of autoimmune type 1 diabetes mellitus in mice. Nature, 351, 542-547.
69. Tokuhiro S., Yamada R., Chang X., Suzuki A., Kochi Y., Sawada T., Suzuki M., Nagasaki M., Ohtsuki M., Ono M., Furukawa H., Nagashima M., Yoshino S., Mabuchi A., Sekine A., Saito S., Takahashi A., Tsunoda T., Nakamura Y. and Yamamoto K. (2003): An intronic SNP in a RUNX1 binding site of SLC22A4, encoding an organic cation transporter, is associated with rheumatoid arthritis. Nat. Genet., 35, 341-348.
70. Wakeland E., Morel L., Achey K., Yui M. and Longmate J. (1997): Speed congenics: a classic technique in the fast lane (relatively speaking). Immunol. Today, 18, 472-477.
71. Wardell B. B., Michael S. D., Tung K. S. K., Todd J. A., Blankenhorn E. P., McEntee K., Sudweeks J., Hansen W. K., Meeker N. D., Griffith J. S., Livingstone K. D. and Teuscher C. (1995): Aod1, the immunoregulatory locus controlling abrogation of tolerance in neonatal thymectomy-induced autoimmune ovarian dysgenesis, maps to mouse chromosome 16. Proc. Natl. Acad. Sci. USA, 92, 4758-4762.
72. Weis J. J., McCracken B. A., Ma Y., Fairbairn D., Roper R. J., Morrison T. B., Weis J. H., Zachary J. F., Doerge R. W. and Teuscher C. (1999): Identification of quantitative trait loci governing arthritis severity and humoral responses in the murine model of Lyme disease. J. Immunol., 162, 948-956.
73. Whitham R. H., Bourdette D. N., Hashim G. A., Herndon R. M., Ilg R. C., Vandenbark A. A. and Offner H. (1991): Lymphocytes from SJL/J mice immunized with spinal cord respond selectively to a peptide of proteolipid protein and transfer relapsing demyelinating experimental autoimmune encephalomyelitis. J. Immunol., 146, 101-107.
74. Wucherpfennig K. W., Sette A., Southwood S., Oseroff C., Matsui M., Strominger J. L. and Hafler D. A. (1994): Structural requirements for binding of an immunodominant myelin basic protein peptide to DR2 isotypes and for its recognition by human T cell clones. J. Exp. Med., 179, 279-290.
75. Wucherpfennig K. W., Zhang J., Witek C., Matsui M., Modabber Y., Ota K. and Hafler D. A. (1994): Clonal expansion and persistence of human T cells specific for an immunodominant myelin basic protein peptide. J. Immunol., 152, 5581-5592.
76. Yokoi N., Komeda K., Wang H. Y., Yano H., Kitada K., Saitoh Y., Seino Y., Yasuda K., Serikawa T. and Seino S. (2002): Cblb is a major susceptibility gene for rat type 1 diabetes mellitus. Nat. Genet., 31, 391-394.
77. Kono D. H., Burlingame R. W., Owens D. G., Kuramochi A., Balderas R. S., Balomenos D. and Theofilopoulos A. N. (1994): Lupus susceptibility loci in New Zealand mice. Proc. Natl. Acad. Sci. USA, 91, 10168-10172.