Protein tyrosine phosphatase PTPN22 in human autoimmunity

Autoimmunity

Volumn 40, Issue 6, 2007, Pages 453-461

  Vang, Torkel ^a   Miletic, Ana V ^a   Bottini, Nunzio ^b   Mustelin, Tomas ^a,c  

^a BURNHAM INSTITUTE   (United States)

^b UNIVERSITY OF SOUTHERN CALIFORNIA   (United States)

^c AMGEN INC   (United States)

Author keywords

Graves' disease; Human autoimmunity; Protein tyrosine phosphatase; Rheumatoid arthritis; Type 1 diabetes

Indexed keywords

CBL PROTEIN; CYCLIC CITRULLINATED PEPTIDE ANTIBODY; CYTOTOXIC T LYMPHOCYTE ANTIGEN 4; GROWTH FACTOR RECEPTOR BOUND PROTEIN 2; HLA DR ANTIGEN; INTERLEUKIN 2; MAJOR HISTOCOMPATIBILITY ANTIGEN; PROTEIN KINASE FYN; PROTEIN KINASE LCK; PROTEIN KINASE ZAP 70; PROTEIN TYROSINE PHOSPHATASE; PROTEIN TYROSINE PHOSPHATASE N22; T LYMPHOCYTE RECEPTOR; UNCLASSIFIED DRUG;

AMINO ACID SEQUENCE; AMINO ACID SUBSTITUTION; ARTICLE; AUTOIMMUNE DISEASE; AUTOIMMUNITY; CELIAC DISEASE; DISEASE SEVERITY; EMPYEMA; ENTERITIS; ENZYME PHOSPHORYLATION; GENE MUTATION; GENETIC ASSOCIATION; GENETIC CODE; GENETIC PREDISPOSITION; GENETIC VARIABILITY; GRAVES DISEASE; HUMAN; IMMUNOPATHOGENESIS; INSULIN DEPENDENT DIABETES MELLITUS; JUVENILE RHEUMATOID ARTHRITIS; MULTIPLE SCLEROSIS; NONHUMAN; PROGNOSIS; PROTEIN FUNCTION; RHEUMATOID ARTHRITIS; SEQUENCE HOMOLOGY; SIGNAL TRANSDUCTION; SINGLE NUCLEOTIDE POLYMORPHISM; STREPTOCOCCUS INFECTION; SYSTEMIC LUPUS ERYTHEMATOSUS; T LYMPHOCYTE; VITILIGO;

ALLELES; AUTOIMMUNE DISEASES; AUTOIMMUNITY; GENETIC PREDISPOSITION TO DISEASE; HUMANS; POLYMORPHISM, SINGLE NUCLEOTIDE; PROTEIN TYROSINE PHOSPHATASE, NON-RECEPTOR TYPE 22; RECEPTORS, ANTIGEN, T-CELL;

EID: 34548323860     PISSN: 08916934     EISSN: 1607842X     Source Type: Journal    
DOI: 10.1080/08916930701464897     Document Type: Article

References (66)

1. Gottlieb, PA and Eisenbarth, GS (2002) In the molecular pathology of autoimmune diseases, p. 588. Taylor & Francis, New York
2. Hirschhorn, JN (2003) Genetic epidemiology of type 1 diabetes. Pediatr Diabetes, 4, pp. 87-100.
3. Maier, LM and Wicker, LS (2005) Genetic susceptibility to type 1 diabetes. Curr Opin Immunol, 17, pp. 601-608.
4. Roep, BO (2003) The role of T-cells in the pathogenesis of type 1 diabetes: From cause to cure. Diabetologia, 46, pp. 305-321.
5. Mustelin, T. and Altman, A. (2002) TCR signaling pathways and their relevance to autoimmunity. The molecular pathology of autoimmune diseases, pp. 127-141. Taylor & Francis, New York, NY
6. Mustelin, T. (2002) Keeping the T cell immune response in balance; role of protein tyrosine phosphatases in autoimmunity. Current directions on autoimmunity, 5, pp. 176-190. Karger Press
7. Nel, AE and Slaughter, N. (2002) T-cell activation through the antigen receptor. Part 2: Role of signaling cascades in T-cell differentiation, anergy, immune senescence, and development of immunotherapy. J Allergy Clin Immunol, 109, pp. 901-915.
8. Buchs, AE and Rapoport, MJ (2000) T cell signaling and autoimmune diabetes. J Pediatr Endocrinol Metab, 13, pp. 1549-1554.
9. Rapoport, MJ, Jaramillo, A., Zipris, D., Lazarus, AH, Serreze, DV, Leiter, EH, Cyopick, P., Danska, JS and Delovitch, TL (1993) Interleukin 4 reverses T cell proliferative unresponsiveness and prevents the onset of diabetes in nonobese diabetic mice. J Exp Med, 178, pp. 87-99.
10. Rapoport, MJ, Lazarus, AH, Jaramillo, A., Speck, E. and Delovitch, TL (1993) Thymic T cell anergy in autoimmune nonobese diabetic mice is mediated by deficient T cell receptor regulation of the pathway of p21ras activation. J Exp Med, 177, pp. 1221-1226.
11. Salojin, K., Zhang, J., Cameron, M., Gill, B., Arreaza, G., Ochi, A. and Delovitch, TL (1997) Impaired plasma membrane targeting of Grb2-murine son of sevenless (mSOS) complex and differential activation of the Fyn-T cell receptor (TCR)-zeta-Cbl pathway mediate T cell hyporesponsiveness in autoimmune nonobese diabetic mice. J Exp Med, 186, pp. 887-897.
12. De Maria, R., Todaro, M., Stassi, G., Blasi, F Di, Giordano, M., Galluzzo, A. and Giordano, C. (1994) Defective T cell receptor/CD3 complex signaling in human type 1 diabetes. Eur J Immunol, 24, pp. 999-1002.
13. Nervi, S., Atlan-Gepner, C., Fossat, C. and Vialettes, B. (1999) Constitutive impaired TCR/CD3-mediated activation of T cells in IDDM patients co-exist with normal co-stimulation pathways. J Autoimmun, 13, pp. 247-255.
14. Ueda, H., Howson, JM, Esposito, L., Heward, J., Snook, H., Chamberlain, G., Rainbow, DB, Hunter, KM, Smith, AN, Genova, G Di, Herr, MH, Dahlman, I., Payne, F., Smyth, D., Lowe, C., Twells, RC, Howlett, S., Healy, B., Nutland, S., Rance, HE, Everett, V., Smink, LJ, Lam, AC, Cordell, HJ, Walker, NM, Bordin, C., Hulme, J., Motzo, C., Cucca, F., Hess, JF, Metzker, ML, Rogers, J., Gregory, S., Allahabadia, A., Nithiyananthan, R., Tuomilehto-Wolf, E., Tuomilehto, J., Bingley, P., Gillespie, KM, Undlien, DE, Ronningen, KS, Guja, C., Ionescu-Tirgoviste, C., Savage, DA, Maxwell, AP, Carson, DJ, Patterson, CC, Franklyn, JA, Clayton, DG, Peterson, LB, Wicker, LS, Todd, JA and Gough, SC (2003) Association of the T-cell regulatory gene CTLA4 with susceptibility to autoimmune disease. Nature, 423, pp. 506-511.
15. Bottini, N., Musumeci, L., Alonso, A., Rahmouni, S., Nika, K., Rostamkhani, M., MacMurray, J., Pellecchia, M., Eisenbarth, GS, Comings, D. and Mustelin, T. (2004) A functional variant of lymphoid tyrosine phosphatase is associated with type I diabetes. Nat Genet, 36, pp. 337-338.
16. Smyth, D., Cooper, JD, Collins, JE, Heward, JM, Franklyn, JA, Howson, JMM, Vella, A., Nutland, S., Rance, HE, Maier, L., Barratt, BJ, Guja, C., Ionescu-Târgovite, C., Savage, DA, Dunger, DB, Widmer, B., Strachan, DP, Ring, SM, Walker, N., Clayton, DG, Twells, RCJ, Gough, SCL and Todd, JA (2004) Replication of an association between the lymphoid tyrosine phosphatase locus (LYP/PTPN22) with type 1 diabetes, and evidence for its role as a general autoimmunity locus. Diabetes, 53, pp. 3020-3023.
17. Ladner, MB, Bottini, N., Valdes, AM and Noble, JA (2005) Association of the single nucleotide polymorphism C1858T of the PTPN22 gene with type 1 diabetes. Hum Immunol, 66, pp. 60-64.
18. Tessier, MC, Hudson, TJ and Polychronakos, C. (2005) Confirmation of the association of the R620W polymorphism in the protein tyrosine phosphatase PTPN22 with type 1 diabetes in a family based study. J Med Genet, 42, pp. 266-270.
19. Zheng, W. and She, JX (2005) Genetic association between a lymphoid tyrosine phosphatase (PTPN22) and type 1 diabetes. Diabetes, 54, pp. 906-908.
20. Begovich, AB, Carlton, VE, Honigberg, LA, Schrodi, SJ, Chokkalingam, AP, Alexander, HC, Ardlie, KG, Huang, Q., Smith, AM, Spoerke, JM, Conn, MT, Chang, M., Chang, SY, Saiki, RK, Catanese, JJ, Leong, DU, Garcia, VE, McAllister, LB, Jeffery, DA, Lee, AT, Batliwalla, F., Remmers, E., Criswell, LA, Seldin, MF, Kastner, DL, Amos, CI, Sninsky, JJ and Gregersen, PK (2004) A Missense single-nucleotide polymorphism in a gene encoding a protein tyrosine phosphatase (PTPN22) is associated with rheumatoid arthritis. Am J Hum Genet, 75, pp. 330-337.
21. van Oene, M., Wintle, RF, Liu, X., Yazdanpanah, M., Newman, B., Kwan, A., Johnson, B., Owen, J., Greer, W., Mosher, D., Maksymowych, W., Keystone, E., Rubin, LA, Amos, CI and Siminovitch, KA (2005) Association of the lymphoid tyrosine phosphatase R620W variant with rheumatoid arthritis, but not Crohn's disease, in Canadian populations. Arthritis Rheum, 52, pp. 1993-1998.
22. Hinks, A., Barton, A., John, S., Bruce, I., Hawkins, C., Griffiths, CE, Donn, R., Thomson, W., Silman, A. and Worthington, J. (2005) Association between the PTPN22 gene and rheumatoid arthritis and juvenile idiopathic arthritis in a UK population: Further support that PTPN22 is an autoimmunity gene. Arthritis Rheum, 52, pp. 1694-1699.
23. Simkins, HM, Merriman, ME, Highton, J., Chapman, PT, O'Donnell, JL, Jones, PB, Gow, PJ, McLean, L., Pokorny, V., Harrison, AA and Merriman, TR (2005) Association of the PTPN22 locus with rheumatoid arthritis in a New Zealand Caucasian cohort. Arthritis Rheum, 52, pp. 2222-2225.
24. Lee, AT, Liew, A., Bombardier, C., Weisman, M., Massarotti, EM, Kent, J., Wolfe, F., Begovich, AB and Gregersen, PK (2005) The PTPN22 R620W polymorphism associates with RF positive rheumatoid arthritis in a dose-dependent manner but not with HLA-SE status. Genes Immun, 6, pp. 129-133.
25. Orozco, G., Sanchez, E., Gonzalez-Gay, MA, Lopez-Nevot, MA, Torres, B., Caliz, R., Ortego-Centeno, N., Jimenez-Alonso, J., Pascual-Salcedo, D., Balsa, A., de Pablo, R., Nunez-Roldan, A., Gonzalez-Escribano, MF and Martin, J. (2005) Association of a functional singlenucleotide polymorphism of PTPN22, encoding lymphoid protein phosphatase, with rheumatoid arthritis and systemic lupus erythematosus. Arthritis Rheum, 52, pp. 219-224.
26. Gregersen, PK and Batliwalla, F. (2005) PTPN22 and rheumatoid arthritis: Gratifying replication. Arthritis Rheum, 52, pp. 1952-1955.
27. Viken, MK, Amundsen, SS, Kvien, TK, Boberg, KM, Gilboe, IM, Lilleby, V., Sollid, LM, Forre, OT, Thorsby, E., Smerdel, A. and Lie, BA (2005) Association analysis of the 1858C > T polymorphism in the PTPN22 gene in juvenile idiopathic arthritis and other autoimmune diseases. Genes Immun, 6, pp. 271-273.
28. Hinks, A., Barton, A., John, S., Bruce, I., Hawkins, C., Griffiths, CE, Donn, R., Thomson, W., Silman, A. and Worthington, J. (2005) Association between the PTPN22 gene and rheumatoid arthritis and juvenile idiopathic arthritis in a UK population: Further support that PTPN22 is an autoimmunity gene. Arthritis Rheum, 52, pp. 1694-1699.
29. Kyogoku, C., Langefeld, CD, Ortmann, WA, Lee, A., Selby, S., Carlton, VEH, Chang, M., Ramos, P., Baechler, EC, Batliwalla, FM, Novitzke, J., Williams, AH, Gillett, C., Rodine, P., Graham, RR, Ardlie, KG, Gaffney, PM, Moser, KL, Petri, M., Begovich, AB, Gregersen, PK and Behrens, TW (2004) Genetic association of the R620W polymorphism of protein tyrosine phosphatase PTPN22 with human SLE. Am J Hum Genet, 75, pp. 504-507.
30. Cantor, RM, Graham, DS, Lingren, CM, Farwell, L., Jager, PL, Bottini, N., Grossman, JM, Wallace, DJ, Hahn, BH, Julkunen, H., Hebert, LA, Rovin, BH, Birmingham, DJ, Rioux, JD, Yu, CY, Kere, J., Vyse, TJ and Tsao, BP (2005) Association analysis of the R620W polymorphism of protein tyrosine phosphatase PTPN22 in systemic lupus erythematosus families: Increased t allele frequency in systemic lupus erythematosus patients with autoimmune thyroid disease. Arthritis Rheum, 52, pp. 2396-2402.
31. Reddy, MV, Johansson, M., Sturfelt, G., Jonsen, A., Gunnarsson, I., Svenungsson, E., Rantapaa-Dahlqvist, S. and Alarcon-Riquelme, ME (2005) The R620W C/T polymorphism of the gene PTPN22 is associated with SLE independently of the association of PDCD1. Genes Immun, 6, pp. 658-662.
32. Behrens, TW, Graham, RR, Kyogoku, C., Baechler, EC, Ramos, PS, Gillett, C., Bauer, J., Ortmann, WA, Hippen, KL, Peterson, E., Langefeld, CD, Moser, KL, Gaffney, PM and Gregersen, PK (2005) Progress towards understanding the genetic pathogenesis of systemic lupus erythematosus. Novartis Found Symp, 267, pp. 145-160.
33. Velaga, MR, Wilson, V., Jennings, CE, Owen, CJ, Herington, S., Donaldson, PT, Ball, SG, James, RA, Quinton, R., Perros, P. and Pearce, SH (2004) The codon 620 tryptophan allele of the lymphoid tyrosine phosphatase (LYP) gene is a major determinant of Graves' disease. J Clin Endocrinol Metab, 89, pp. 5862-5865.
34. Skorka, A., Bednarczuk, T., Bar-Andziak, E., Nauman, J. and Ploski, R. (2005) Lymphoid tyrosine phosphatase (PTPN22/LYP) variant and Graves' disease in a polish population: Association and gene dose-dependent correlation with age of onset. Clin Endocrinol (Oxf), 62, pp. 679-682.
35. Canton, I., Akhtar, S., Gavalas, NG, Gawkrodger, DJ, Blomhoff, A., Watson, PF, Weetman, AP and Kemp, EH (2005) A single-nucleotide polymorphism in the gene encoding lymphoid protein tyrosine phosphatase (PTPN22) confers susceptibility to generalised vitiligo. Genes Immun, 6, pp. 584-587.
36. Begovich, AB, Caillier, SJ, Alexander, HC, Penko, JM, Hauser, SL, Barcellos, LF and Oksenberg, JR (2005) The R620W polymorphism of the protein tyrosine phosphatase PTPN22 is not associated with multiple sclerosis. Am J Hum Genet, 76, pp. 184-187.
37. Carlton, VE, Chokkalingam, AP, Schrodi, SJ, Brandon, R., Alexander, HC, Chang, M., Catanese, JJ, Leong, DU, Ardlie, KG, Kastner, DL, Seldin, MF, Criswell, LA, Gregersen, PK, Beasley, E., Thomson, G., Amos, CI and Begovich, AB (2005) PTPN22 genetic variation: Evidence for multiple variants associated with rheumatoid arthritis. Am J Hum Genet, 77, pp. 567-581.
38. Huffmeier, U., Steffens, M., Burkhardt, H., Lascorz, J., Schurmeier-Horst, F., Stander, M., Kelsch, R., Baumann, C., Kuster, W., Mossner, R., Reich, K., Wienker, TF, Traupe, H. and Reis, A. (2005) Evidence for susceptibility determinant(s) to psoriasis vulgaris in or near PTPN22 in German patients. J Med Genet
39. Concannon, P., Erlich, HA, Julier, C., Morahan, G., Nerup, J., Pociot, F., Todd, JA and Rich, SS (2005) Type 1 diabetes genetics consortium. Type 1 diabetes: Evidence for susceptibility loci from four genome-wide linkage scans in 1,435 multiplex families. Diabetes, 54, pp. 2995-3001.
40. Plenge, RM, Padyukov, L., Remmers, EF, Purcell, S., Lee, AT, Karlson, EW, Wolfe, F., Kastner, DL, Alfredsson, L., Altshuler, D., Gregersen, PK, Klareskog, L. and Rioux, JD (2005) Replication of putative candidate-gene associations with rheumatoid arthritis in>4,000 samples from North America and Sweden: Association of susceptibility with PTPN22, CTLA4, and PADI4. Am J Hum Genet, 77, pp. 1044-1060.
41. Cohen, S., Dadi, H., Shaoul, E., Sharfe, N. and Roifman, CM (1999) Cloning and characterization of a lymphoid-specific, inducible human protein tyrosine phosphatase, Lyp. Blood, 93, pp. 2013-2024.
42. Matthews, RJ, Bowne, DB, Flores, E. and Thomas, ML (1992) Characterization of hematopoietic intracellular protein tyrosine phosphatases: Description of a phosphatase containing an SH2 domain and another enriched in proline-, glutamic acid-, serine-, and threonine-rich sequences. Mol Cell Biol, 12, pp. 2396-2405.
43. Cloutier, JF and Veillette, A. (1996) Association of inhibitory tyrosine protein kinase p50csk with protein tyrosine phosphatase PEP in T cells and other hemopoietic cells. EMBO J, 15, pp. 4909-4918.
44. Gregorieff, A., Cloutier, JF and Veillette, A. (1998) Sequence requirements for association of protein-tyrosine phosphatase PEP with the Src homology 3 domain of inhibitory tyrosine protein kinase p50(Csk). J Biol Chem, 273, pp. 13217-13222.
45. Nada, S., Okada, M., MacAuley, A., Cooper, JA and Nakagawa, H. (1991) Cloning of a complementary DNA for a protein-tyrosine kinase that specifically phosphorylates a negative regulatory site of p60c-src. Nature, 351, pp. 69-72.
46. Bergman, M., Mustelin, T., Oetken, C., Partanen, J., Flint, NA, Amrein, KE, Autero, M., Burn, P. and Alitalo, K. (1992) The human p50csk tyrosine kinase phosphorylates p56lck at Tyr-505 and down-regulates its catalytic activity. EMBO J, 11, pp. 2919-2924.
47. Chow, LML, Fournel, M., Davidson, D. and Veillette, A. (1993) Negative regulation of T-cell receptor signalling by the tyrosine kinase p50csk. Nature, 365, pp. 156-160.
48. Gjörloff-Wingren, A., Saxena, M., Williams, S., Hammi, D. and Mustelin, T. (1999) Characterization of TCR-induced receptor-proximal signaling events negatively regulated by the protein tyrosine phosphatase PEP. Eur J Immunol, 29, pp. 3845-3854.
49. Cloutier, JF and Veillette, A. (1999) Cooperative inhibition of T-cell antigen receptor signaling by a complex between a kinase and a phosphatase. J Exp Med, 189, pp. 111-121.
50. Ghose, R., Shekhtman, A., Goger, MJ and Cowburn, D. (2001) A novel, specific interaction involving the Csk SH3 domain and its natural ligand. Nat Struct Biol, 8, pp. 998-1004.
51. Hill, RJ, Zozulya, S., Lu, YL, Ward, K., Gishizky, M. and Jallal, B. (2002) The lymphoid protein tyrosine phosphatase Lyp interacts with the adaptor molecule Grb2 and functions as a negative regulator of T-cell activation. Exp Hematol, 30, pp. 237-244.
52. Spencer, S., Dowbenko, D., Cheng, J., Brush, J., Utzig, S., Simanis, V. and Lasky, LA (1997) PSTPIP: A tyrosine phosphorylated cleavage furrow-associated protein that is a substrate for a PEST tyrosine phosphatase. J Cell Biol, 138, pp. 845-860.
53. Dowbenko, D. and Lasky, LA (1998) PSTPIP 2, a second tyrosine phosphorylated, cytoskeletal-associated protein that binds a PEST-type protein-tyrosine phosphatase. J Biol Chem, 273, pp. 30487-30496.
54. Cote, JF, Chung, PL, Theberge, JF, Halle, M., Spencer, S., Lasky, LA and Tremblay, ML (2002) PSTPIP is a substrate of PTP-PEST and serves as a scaffold guiding PTP-PEST toward a specific dephosphorylation of WASP. J Biol Chem, 277, pp. 2973-2986.
55. Katrekar, K., Honigberg, LA, Smith, AM, Conn, MT, Tang, J., Jeffrey, D., Mortara, K., Sampang, J., Williams, SR, Buggy, J. and Clark, JM (2006) Identification of substrates of human protein tyrosine phosphatase PTPN22. J Biol Chem, 281, pp. 11002-11010.
56. Davidson, D., Bakinowski, M., Thomas, ML, Horejsi, V. and Veillette, A. (2003) Phosphorylationdependent regulation of T-cell activation by PAG/ Cbp, a lipid raft-associated transmembrane adaptor. Mol Cell Biol, 23, pp. 2017-2028.
57. Huo, J., Tan, JE and Lam, KP (2005) Cbp deficiency alters Csk localization in lipid rafts but does not affect T-cell development. Mol Cell Biol, 25, pp. 8486-8495.
58. Dobenecker, MW, Schmedt, C., Okada, M. and Tarakhovsky, A. (2005) The ubiquitously expressed Csk adaptor protein Cbp is dispensable for embryogenesis and T-cell development and function. Mol Cell Biol, 25, pp. 10533-10542.
59. Hasegawa, K., Martin, F., Huang, G., Tumas, D., Diehl, L. and Chan, AC (2004) PEST domainenriched tyrosine phosphatase (PEP) regulation of effector/memory T cells. Science, 303, pp. 685-689.
60. Vang, T., Congia, M., Macis, MD, Musumeci, L., Orru, V., Zavattari, P., Nika, K., Tautz, L., Taskén, K., Cucca, F., Mustelin, T. and Bottini, N. (2005) Autoimmune-associated lymphoid tyrosine phosphatase is a gain-of-function variant. Nat Genet, 37, pp. 1317-1319.
61. Burkhardt, H., Huffmeier, U., Spriewald, B., Bohm, B., Rau, R., Kallert, S., Engstrom, A., Holmdahl, R. and Reis, A. (2006) Association between protein tyrosine phosphatase 22 variant R620W in conjunction with the HLA-DRB1 shared epitope and humoral autoimmunity to an immunodominant epitope of cartilage-specific type II collagen in early rheumatoid arthritis. Arthritis Rheum, 54, pp. 82-89.
62. Sakaguchi, N., Takahashi, T., Hata, H., Nomura, T., Tagami, T., Yamazaki, S., Sakihama, T., Matsutani, T., Negishi, I., Nakatsuru, S. and Sakaguchi, S. (2003) Altered thymic T-cell selection due to a mutation of the ZAP-70 gene causes autoimmune arthritis in mice. Nature, 426, pp. 454-460.
63. Gomes, LM, Anaya, JM and Martin, J. (2005) Genetic influence of PTPN22 R620W polymorphism in tuberculosis. Hum Immunol, 66, pp. 1242-1247.
64. Chapman, SJ, Khor, CC, Vannberg, FO, Maskell, NA, Davies, CW, Hedley, EL, Segal, S., Moore, CE, Knox, K., Day, NP, Gillespie, SH, Crook, DW, Davies, RJ and Hill, AV (2006) PTPN22 and invasive bacterial disease. Nat Genet, 38, pp. 499-500.
65. Mustelin, T. (2006) Are other protein tyrosine phosphatases than PTPN22 associated with autoimmunity?. Sem Immunol, 18, pp. 254-260.
66. Mustelin, T., Vang, T. and Bottini, N. (2005) Protein tyrosine phosphatases and the immune response. Nat Rev Immunol, 5, pp. 43-57.