The JAK/STAT pathway

Cancer Surveys

Volumn 27, Issue , 1996, Pages 139-163

  Wilks, Andrew F ^a   Oates, Andrew C ^a  

^a ROYAL MELBOURNE HOSPITAL   (Australia)

Author keywords

[No Author keywords available]

Indexed keywords

CYTOKINE RECEPTOR;

DNA TRANSCRIPTION; GENE MUTATION; GENETIC SCREENING; PRIORITY JOURNAL; PROTEIN PHOSPHORYLATION; REGULATORY MECHANISM; REVIEW; SIGNAL TRANSDUCTION; SOMATIC CELL GENETICS;

ANIMALS; INTERFERONS; PROTEIN-TYROSINE KINASES; SIGNAL TRANSDUCTION; TRANS-ACTIVATORS;

EID: 0029911962     PISSN: 02612429     EISSN: None     Source Type: Journal    
DOI: None     Document Type: Review

References (113)

1. Ackrill AM, Reid LE, Gilbert CS et al (1991) Differential response of the human 6-16 and 9-27 genes to alpha and gamma interferons. Nucleic Acids Research 19 591-598
2. Akira S, Nishio Y, Inoue M et al (1994) Molecular cloning of APRF, a novel IFN-stimulated gene factor 3 p91-related transcription factor involved in the gp130-mediated signaling pathway. Cell 77 63-71
3. Argetsinger LS, Campbell GS, Yang X et al (1993) Identification of JAK2 as a growth hormone receptor-associated tyrosine kinase. Cell 74 237-244
4. Argetsinger LS, Hsu GW, Myers MG Jr, Billestrup N, White MF and Carter Su C (1995) Growth hormone, interferon-gamma, and leukemia inhibitory factor promoted tyrosyl phosphorylation of insulin receptor substrate-1. Journal of Biological Chemistry 270 14685-14692
5. Bazan JF (1990) Structural design and molecular evolution of a cytokine receptor superfamily. Proceedings of the National Academy of Sciences of the USA 87 6934-6938
6. Bhat GJ, Thekkumkara TJ, Thomas WG, Conrad KM and Baker KM (1994) Angiotensin II stimulates sis-inducing factor-like DNA binding activity. Evidence that the AT1A receptor activates transcription factor-Stat91 and/or a related protein. Journal of Biological Chemistry 269 31443-31449
7. 1A cells. Cross-talk between angiotensin II and interleukin-6 nuclear signaling. Journal of Biological Chemistry 270 19059-19065
8. Biggs J, Tripoulos N, Hersperger H, Dearolf C and Shearn A (1988) Analysis of the lethal interaction between the prune and killer of prune mutations of Drosophila. Cell 2 1333-1343
9. Biggs J, Hersperger E, Steeg PS, Liotta LA and Shearn A (1990) A Drosophila gene that is homologous to a mammalian gene associated with tumor metastasis codes for a nucleoside diphosphate kinase. Cell 63 933-940
10. Binari R and Perrimon N (1994) Stripe specific regulation of pair-rule genes by hopscotch, a putative JAK family tyrosine kinase. Genes and Development 8 300-312
11. Boulton TG, Zhong Z, Wen Z, Darnell JE Jr, Stahl N and Yancopoulos GD (1995) STAT3 activation by cytokines utilizing gp130 and related transducers involves a secondary modification requiring an H7-sensitive kinase. Proceedings of the National Academy of Sciences of the USA 92 6915-6919
12. Chinkers M and Garbers DL (1989) The protein kinase domain of the ANP receptor is required for signaling. Science 245 1392-1394
13. Corwin HO and Hanratty WP (1976) Characterization of a unique lethal tumorous mutation in drosophila. Molecular and General Genet ics 144 345-347
14. Dale TC, Rosen JM, Guille MJ et al (1989) Overlapping sites for constitutive and induced DNA binding factors involved in interferon-stimulated transcription. EMBO Journal 8 831-839
15. DaSilva L, Howard OMZ, Rui H, Kirken RA and Farrar WL (1994) Growth signaling and JAK2 association mediated by membrane-proximal cytoplasmic regions of prolactin receptors. Journal of Biological Chemistry 269 18267-18270
16. David M, Petricoin E III, Benjamin C, Pine R, Weber MJ and Larner AC (1995) Requirement for MAP kinase (ERK2) activity in interferon alpha- and interferon beta-stimulated gene expression through stat proteins. Science 269 1721-1723
17. Decker T, Lew DJ, Cheng Y-SE, Levy DE and Darnell JE (1989) Interactions of alpha- and gamma-interferon in the transcriptional regulation of the gene encoding a guanylate-binding protein. EMBO Journal 8 2009-2014
18. Eilers A, Georgellis D, Klose B et al (1995) Differentiation-regulated serine phosphorylation of STAT1 promotes GAF activation in macrophages. Molecular and Cellular Biology 15 3579-3586
19. Firmbach Kraft I, Byers M, Shows T, Dalla Favera R and Krolewski JJ (1990) tyk2, prototype of a novel class of non-receptor tyrosine kinase genes. Oncogene 5 1329-1336
20. Frank SJ, Gilliland G, Kraft AS and Arnold CS (1994) Interaction of the growth hormone receptor cytoplasmic domain with the JAK2 tyrosine kinase. Endocrinology 135 2228-2239
21. Friedman RL, Manly SP, McMahon M, Kerr IM and Stark GR (1984) Transcriptional and post-transcriptional regulation of interferon-induced gene expression in human cells. Cell 38 745-755
22. Fu X-Y, Schindler C, Improta T, Aebersold R and Darnell JE Jr (1992) The proteins of ISGF-3, the interferon α-induced transcriptional activator, define a gene family involved in signal transduction. Proceedings of the National Academy of Sciences of the USA 89 7840-7843
23. Fukunaka R, Ishizaka-Ikeda E, Seto Y and Nagata S (1990) Expression cloning of a receptor for murine granulocyte colony stimulating factor. Cell 61 341-350
24. Greenlund AC, Farrar MA, Viviano BL and Schreiber RD (1994) Ligand-induced IFNγ receptor tyrosine phosphorylation couples the receptor to its signal transduction system (p91). EMBO Journal 13 1591-1600
25. Greenlund AC, Morales MO, Viviano BL, Yan H, Krolewski J, Schreiber RD (1995) Stat recruitment by tyrosine-phosphorylated cytokine receptors: an ordered reversible affinitydriven process. Immunity 2 677-687
26. Guyer NB, Severns CW, Wong P, Feghali CA and Wright TM (1995) IFN-gamma induces a p91/Stat1 alpha-related transcription factor with distinct activation and binding properties. Journal of Immunology 155 3472-3480
27. Hackstein JHP (1992) The lethal prune/killer-of-prune interaction of Drosophila causes a syndrome resembling human neurofibromatosis (NFI). European Journal of Cell Biology 58 429-444
28. Hanks SK and Quinn AM (1991) Protein kinase catalytic domain sequence data-base: identification of conserved features of primary structure and classification of family members. Methods in Enzymology 200 38-62
29. Hanratty WP and Dearolf CR. (1993) The Drosophila Tumorous-lethal hematopoietic oncogene is a dominant mutation in the hopscotch locus. Molecular and General Genetics 238 33-37
30. Hanratty WP and Ryerse JS (1981) A genetic melanotic neoplasm of Drosophila melanogaster. Developmental Biology 83 238-249
31. Haque SJ and Williams BRG (1994) Identification and characterization of an interferon (IFN)-stimulated response element-IFN-stimulated gene factor 3-independent signaling pathway for IFN-alpha. Journal of Biological Chemistry 269 19523-19529
32. Harpur AG, Andres A-C, Zeimiecki A, Aston R and Wilks AF (1992) JAK2, a third member of the JAK family of protein tyrosine kinases. Oncogene 7 1347-1353
33. Harrison DA, Binari R, Nahreini TS, Gilman M and Perrimon N (1995) Activation of a Drosophila Janus kinase (JAK) causes hematopoietic neoplasia and developmental defects. EMBO Journal 14 2857-2865
34. He TC, Jiang N, Zhuang H and Wojchowski DM (1995) Erythropoietin-induced recruitment of She via a receptor phosphotyrosine-independent, Jak2-associated pathway. Journal of Biological Chemistry 270 11055-11061
35. Heim MH, Kerr IM, Stark GR and Darnell JE (1995) Contribution of STAT SH2 groups to specific interferon signaling by the Jak-STAT pathway. Science 267 1347-1349
36. Horvath CM, Wen Z and Darnell JE Jr (1995) A STAT protein domain that determines DNA sequence recognition suggests a novel DNA-binding domain. Genes and Development 9 984-994
37. Horvitz HR and Sternberg PW (1991) Multiple intercellular signalling systems control the development of the Caenorhahditis elegans vulva. Nature 351 535-541
38. Hou J, Schindler U, Henzel WJ, Ho TC, Brasseur M and McKnight SL (1994) An interleukin-4-induced transcription factor: IL-4 Stat. Science 265 1701-1706
39. Igarashi K, David M, Lamer AC and Finbloom DS (1993) In vitro activation of a transcription factor by gamma interferon requires a membrane-associated tyrosine kinase and is mimicked by vanadate. Molecular and Cellular Biology 13 3984-3989
40. Igarashi K-I, Garotta G, Ozmen L et al (1994) Interferon-gamma induces tyrosine phosphorylation of interferon-gamma receptor and regulated association of protein tyrosine kinases, Jak1 and Jak2, with its receptor. Journal of Biological Chemistry 269 14333-14336
41. Imam AMA, Ackrill AM, Dale TC, Kerr IM and Stark GR (1990) Transcription factors induced by interferons alpha and gamma. Nucleic Acids Research 18 6573-6580
42. Improta T, Schindler C, Horvath CM, Kerr IM, Stark GR and Darnell JE Jr (1994) Transcription factor ISGF-3 formation requires phosphorylated Stat91 protein, but Stat113 protein is phosphorylated independently of Stat91 protein. Proceedings of the National Academy of Sciences of the USA 91 4776-4780
43. Kawamura M, McVicar DW, Johnston JA et al (1994) Molecular cloning of L-JAK, a Janus family protein-tyrosine kinase expressed in natural killer cells and activated leukocytes. Proceedings of the National Academy of Sciences of the USA 91 6374-6378
44. Kessler DS, Veals SA, Fu X-Y and Levy DE (1990) Interferon alpha regulates nuclear translocation and DNA-binding affinity of ISGF3, a multimeric transcriptional activator. Genes and Development 4 1753-1765
45. Klingmuller U, Lorenz U, Cantley LC, Neel BG and Lodish HF (1995) Specific recruitment of SH-PTP1 to the erythropoietin receptor causes inactivation of JAK2 and termination of proliferative signals. Cell 80 729-738
46. Lamb P, Seidel HM, Haslam J et al (1995) STAT protein complexes activated by interferon-gamma and gp130 signaling molecules differ in their sequence preferences and transcriptional induction properties. Nucleic Acids Research 23 3283-3289
47. Lascu I, Chaffotte A, Limbourg-Bouchon B and Veron M (1992) A Pro/Ser substitution in nucleoside diphosphate kinase of Drosophila melanogaster (mutation killer of prune) affects stability but not catalytic efficiency of the enzyme. Journal of Biological Chemistry 267 12775-12781
48. Leaman DW, Pisharody S, Flickinger TW et al (1996) Roles of JAKs in activation of STATs and stimulation of c-fos gene expression by epidermal growth factor. Molecular and Cellular Biology 16 369-375
49. Lebrun JJ, Ali S, Coffin V, Ullrich A and Kelly PA (1995a) A single phosphotyrosine residue of the prolactin receptor is responsible for activation of gene transcription. Proceedings of the National Academy of Sciences of the USA 92 4031-4035
50. Lebrun JJ, Ali S, Ullrich A and Kelly PA (1995b) Proline-rich sequence-mediated Jak2 association to the prolactin receptor is required but not sufficient for signal transduction. Journal of Biological Chemistry 270 10664-10670
51. Leung S, Qureshi SA, Kerr IM, Darnell JE Jr and Stark GR (1995) Role of STAT2 in the alpha interferon signaling pathway. Molecular and Cellular Biology 15 1312-1317
52. Levy DE, Kessler DS, Pine R and Darnell JE Jr (1989) Cytoplasmic activation of ISGF3, the positive regulator of interferon alpha-stimulated transcription, reconstituted in vitro. Genes and Development 3 1362-1371
53. Lin JX, Migone TS, Tsang M et al (1995) The role of shared receptor motifs and common Stat proteins in the generation of cytokine pleiotropy and redundancy by IL-2, IL-4, IL-7, IL-13, and IL-15. Immunity 2 331-339
54. Loh JE, Chang C-H, Fodor WL and Flavell RA (1992) Dissection of the interferon gamma-MHC class II signal transduction pathway reveals that type I and type II interferon systems share common signalling component(s). EMBO Journal 11 1351-1363
55. Tum-1 jak kinase causes leukemia-like hematopoietic defects. EMBO Journal 14 1412-1420
56. McKendry R, John J, Flavell D, Muller M, Kerr IM and Stark GR (1991) High-frequency mutagenesis of human cells and characterization of a mutant unresponsive to both alpha and gamma interferons. Proceedings of the National Academy of Sciences of the USA 88 11455-11459
57. Malmqvist M (1993) Biospecific interaction analysis using biosensor technology. Nature 361 186-187
58. Marrero MB, Schieffer B, Paxton WG et al (1995) Direct stimulation of Jak/STAT pathway by the angiotensin II AT1 receptor. Nature 375 247-250
59. Matsuguchi T, Inhorn RC, Carlesso N, Xu G, Druker B and Griffin JD (1995) Tyrosine phosphorylation of p95Vav in myeloid cells is regulated by GM-CSF, IL-3 and steel factor and is constitutively increased by p210BCR/ABL. EMBO Journal 14 257-265
60. Muller M, Brisce J, Laxton C et al (1993a) The protein tyrosine kinase JAK1 complements defects in interferon-alpha/beta and -gamma signal. Nature 366 129-135
61. Muller M, Laxton C, Briscoe J et al (1993b) Complementation of a mutant cell line: central role of the 91 kDa polypeptide of ISGF3 in the interferon-α and γ signal transduction pathways. EMBO Journal 12 4221-4228
62. Nappi AJ and Silvers M (1984) Cell surface changes associated with cellular immune reactions in Drosophila. Science 225 1166-1168
63. Nicholson SE, Oates AC, Harpur AG, Ziemiecki A, Wilks AF and Layton JE (1994) Tyrosine kinase JAK1 is associated with the granulocyte-colony-stimulating factor receptor and both become tyrosine-phosphorylated after receptor activation. Proceedings of the National Academy of Sciences of the USA 91 2985-2988
64. Novick D, Cohen B and Rubinstein M (1994) The human interferon alpha/beta receptor: characterization and molecular cloning. Cell 77 391-400
65. Pankratz MJ and Jackle H (1993) Blastoderm segmentation, In: Bate M and Martinez Arias A (eds). The Development of Drosophila melanogaster vol. 1 pp 467-516, Cold Spring Harbor Labortory Press, Cold Spring Harbor, New York
66. Pawson T (1992) SH2 and SH3 domains. Current Opinion in Structural Biology 2 432-437
67. Pawson T (1994) SH2 and SH3 domains in signal transduction. Advances in Cancer Research 64 87-110
68. Pearse RN, Feinman R and Ravetch JV (1991) Characterization of the promoter of the human gene encoding the high affinity IgG receptor: transcriptional induction by gamma interferon is mediated through common DNA responsible elements. Proceedings of the National Academy of Sciences of the USA 88 11305-11309
69. Pellegrini S, John J, Shearer M, Kerr IM and Stark GR (1989) Use of a selectable marker regulated by alpha interferon to obtain mutations in the signaling pathway. Molecular and Cellular Biology 9 4605-4612
70. Perez C, Wietzerbin J and Benech PD (1993) Two cis-DNA elements involved in myeloid-cell-specific expression and gamma interferon (IFNγ) activation of the human high-affinity Fc gamma receptor gene: a novel IFN regulatory mechanism. Molecular and Cellular Biology 13 2182-2192
71. Perrimon N and Mahowald AP (1986) 1 (1) hopscotch, a larval-pupal zygotic lethal with a specific maternal effect on segmentation in Drosophila. Developmental Biology 118 28-41
72. Perrimon N, Engstrom L and Mahowald AP (1989) Zygotic lethals with specific maternal effect phenotypes in drosophila melanogaster. I. Loci on the X chromosome. Genetics 121 333-352
73. Qureshi SA, Salditt Georgieff M and Darnell JE (1995) Tyrosine-phosphorylated Stat1 and Stat2 plus a 48-kDa protein all contact DNA in forming interferon-stimulated-gene factor 3. Proceedings of the National Academy of Sciences of the USA 92 3829-3833
74. Reid LE, Brasnett AH, Gilbert CS et al (1989) A single DNA response element can confer inducibility by both a- and γ-interferons. Proceedings of the National Academy of Sciences of the USA 86 840-844
75. Rizki TM and Rizki RM (1978) Larval adipose tissue of homoeotic bithorax mutants of Drosophila. Developmental Biology 65 476-482
76. Rui H, Kirken RA and Farrar WL (1994) Activation of receptor-associated tyrosine kinase JAK2 by prolactin journal of Biological Chemistry 269 5364-5368
77. Russell SM, Johnston JA, Noguchi M et al (1994) Interaction of IL-2Rbβ and gamma chains with Jak1 and Jak3: implications for XSCID and XCID. Science 266 1042-1045
78. Schaefer TS, Sanders LK and Nathans D (1995) Cooperative transcriptional activity of Jun and STAT3beta, a short form of Stat 3. Proceedings of the National Academy of Science USA 92 9097-9101
79. Schindler C, Fu X-F, Improta T, Aebersold R and Darnell JE Jr (1992) Proteins of transcription factor ISGF-3: one gene encodes the 91- and 84-kDa ISGF-3 proteins that are activated by interferon α. Proceedings of the National Academy of Sciences of the USA 89 7836-7839
80. Schindler U, Wu P, Rothe M, Brasseur M and McKnight SL (1995) Components of a Stat recognition code: evidence for two layers of molecular selectivity. Immunity 2 689-697
81. Shuai K, Stark GR, Kerr IM and Darnell Jr JE (1993) A single phosphotyrosine residue of Stat91 required for gene activation by interferon-gamma. Science 261 1744-1746
82. Shuai K, Horvath CM, Huang LH, Qureshi SA, Cowburn D and Darnell JE Jr (1994) Interferon activation of the transcription factor Stat91 involves dimerization through SH2 - phosphotyrosyl peptide interactions. Cell 76 821-828
83. Simon MA, Bowtell DD, Dodson GS, Laverty TR and Rubin GM (1991) Rasl and a putative guanine nucleotide exchange factor perform crucial steps in signaling by the sevenless protein tyrosine kinase. Cell 67 701-716
84. Sims SH, Cha Y, Romine MF, Gao P-Q, Gottlieb K and Deisseroth AB (1993) A novel interferon-inducible domain: structural and functional analysis of the human interferon regulatory factor 1 gene promoter. Molecular and Cellular Biology 13 690-702
85. Stahl N, Boulton TG, Farruggella T et al (1994) Association and activation of Jak-Tyk kinases by CNTF-LIF-IL-6 β receptor components. Science 263 92-95
86. Stahl N, Farruggella TJ, Boulton TG, Zhong Z, Darnell JE Jr and Yancopoulos GD (1995) Choice of STATs and other substrates specified by modular tyrosine-based motifs in cytokine receptors. Science 267 1349-1353
87. Sun XJ, Rothenberg P, Kahn CR et al (1991) Structure of the insulin receptor substrate IRS-1 defines a unique signal transduction protein. Nature 352 73-77
88. Sun XJ, Wang LM, Zhang Y et al (1995) Role of IRS-2 in insulin and cytokine signalling. Nature 377 173-177
89. Sweitzer SM, Calvo S, Kraus MH, Finbloom DS and Larner AC (1995) Characterization of a Stat-like DNA binding activity in Drosophila melanogaster. Journal of Biological Chemistry 270 16510-16513
90. Takahashi T and Shirasawa T (1994) Molecular cloning of rat JAK3, a novel member of the JAK family of protein tyrosine kinases. FEBS Let ters 342 124-128
91. Tanaka N, Aso H, Ohbo K et al (1994) Physical association of JAK1 and JAK2 kinases with the interleukin 2 receptor beta and gamma chains. Proceedings of the National Academy of Sciences of the USA 91 7271-7275
92. Tanner JW, Chen W, Young RL, Longmore GD and Shaw AS (1995) The conserved box 1 motif of cytokine receptors is required for association with JAK kinases. Journal of Biological Chemistry 270 6523-6530
93. Teng DHF, Engele CM and Venkatesh TR (1991) A product of the prune locus of Drosophila is similar to mammalian GTPase-activating protein. Nature 353 437-440
94. Tsui HW, Siminovitch KA, de Souza L and Tsui FW (1993) Motheaten and viable motheaten mice have mutations in the haematopoietic cell phosphatase gene. Nature Genetics 4 124-129
95. Ullrich A and Schlessinger J (1990) Signal transduction by receptors with tyrosine kinase activity. Cell 61 203-212
96. VanderKuur JA, Wang X, Zhang L et al (1994) Domains of the growth hormone receptor required for association and activation of JAK2 tyrosine kinase. Journal of Biological Chemistry 269 21709-21717
97. Veals SA, Schindler C, Leonard D et al (1992) Subunit of an alpha-interferon-responsive transcription factor is related to interferon regulatory factor and Myb families of DNA-binding proteins. Molecular and Cellular Biology 12 3315-3324
98. Velazquez L, Fellous M, Stark GR and Pellegrini S (1992) A protein tyrosine kinase in the interferon alpha/beta signaling pathway. Cell 70 313-322
99. Velazquez L, Mogensen KE, Barbieri G, Fellous M, Uze G and Pellegrini S (1995) Distinct domains of the protein tyrosine kinase tyk2 required for binding of interferon-alpha/beta and for signal transduction. Journal of Biological Chemistry 270 3327-3334
100. Wakao H, Gouilleux F and Groner B (1994) Mammary gland factor (MGF) is a novel member of the cytokine regulated transcription factor gene family and confers the prolactin response. EMBO Journal 13 2182-2191
101. Wakao H, Gouilleux F and Groner B (1995) Mammary gland factor (MGF) is a novel member of the cytokine regulated transcription factor gene family and confers the prolactin response. EMBO Journal 14 854-855
102. Wang LM, Myers MG Jr, Sun XJ, Aaronson SA, White M and Pierce JH (1993) IRS-1: essential for insulin- and IL-4-stimulated mitogenesis in hematopoietic cells. Science 261 1591-1594
103. Watling D, Guschin D, Muller M et al (1993) Complementation by the protein tyrosine kinase JAK2 of a mutant cell line defective in the interferon-γ signal transduction pathway. Nature 366 166-170
104. Wen Z, Zhong Z and Darnell JE Jr (1995) Maximal activation of transcription by Stat1 and Stat3 requires both tyrosine and serine phosphorylation. Cell 82 241-250
105. Wilks AF (1989) Two putative protein-tyrosine kinases identified by application of the polymerase chain reaction. Proceedings of the National Academy of Sciences of the USA 86 1603-1607
106. Wilks AF, Harpur AG, Kurban RR, Ralph SJ, Zurcher G and Ziemiecki A (1991) Two novel protein-tyrosine kinases, each with a second phosphotransferase-related catalytic domain, define a new class of protein kinase. Molecular and Cellular Biology 11 2057-2065
107. Witthuhn BA, Quelle FW, Silvennoinen O et al (1993) JAK2 associates with the erythropoietin receptor and is tyrosine phosphorylated and activated following stimulation with erythropoietin. Cell 74 227-236
108. Yan R, Qureshi S, Zhong Z, Wen Z and Daenell JE (1995) The genomic structure of the STAT genes: multiple exons in coincident sites in Stat1 and Stat2. Nucleic Acids Research 23 459-463
109. Zhang X, Blenis J, Li HC, Schindler C and Chen Kiang S (1995) Requirement of serine phosphorylation for formation of STAT-promoter complexes. Science 267 1990-1994
110. Zhao Y, Wagner F, Frank SA and Kraft AS (1995) The amino-terminal portion of the JAK2 protein kinase is necessary for binding and phosphorylation of the granulocyte-macrophage colony stimulating factor β chain. Journal of Biological Chemistry 270 13814-13818
111. Zhong Z, Wen Z and Darnell Jr JE (1994a). Stat3 and Stat4: members of the family of signal transducers and activators of transcription. Proceedings of the National Academy of Sciences of the USA 91 4806-4810
112. Zhong Z, Wen Z and Darnell JE (1994b) Stat3: a STAT family member activated by tyrosine phosphorylation in response to epidermal growth factor and interleukin-6. Science 264 95-98
113. K-pn a homologue of the metastasis suppressor gene nm23, suppresses the Tum-1 haematopoietic oncogene. Nature Genetics 4 195-201