The fat and warts signaling pathways: New insights into their regulation, mechanism and conservation

Development

Volumn 135, Issue 17, 2008, Pages 2827-2838

  Reddy, B V V G ^a   Irvine, Kenneth D ^a  

^a RUTGERS UNIVERSITY   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

CYTOPLASM PROTEIN; DNA; DROSOPHILA PROTEIN; EXPANDED; MERLIN; PHOSPHOTRANSFERASE; PROTEIN FAT; PROTEIN KINASE; PROTEIN KINASE HIPPO; PROTEIN KINASE WARTS; REGULATOR PROTEIN; TRANSCRIPTION FACTOR; TRANSCRIPTION FACTOR YORKIE; TUMOR SUPPRESSOR PROTEIN; UNCLASSIFIED DRUG;

BIOLOGICAL ACTIVITY; BREAST CANCER; BREAST CARCINOGENESIS; CELL POLARITY; CELLULAR DISTRIBUTION; CONTACT INHIBITION; DEVELOPMENTAL BIOLOGY; DNA DETERMINATION; DROSOPHILA; ENZYME PHOSPHORYLATION; ENZYME SUBSTRATE; GENE TARGETING; GENETIC CONSERVATION; GROWTH REGULATION; LIVER CANCER; MALIGNANT NEOPLASTIC DISEASE; MAMMALIAN GENETICS; MOLECULAR DYNAMICS; NONHUMAN; PRIORITY JOURNAL; PROSTATE CANCER; PROTEIN DNA BINDING; PROTEIN FUNCTION; PROTEIN LOCALIZATION; PROTEIN PROTEIN INTERACTION; REVIEW; SIGNAL TRANSDUCTION; TISSUE REGENERATION; TRANSCRIPTION REGULATION;

ANIMALS; CELL ADHESION MOLECULES; CONSERVED SEQUENCE; DROSOPHILA MELANOGASTER; DROSOPHILA PROTEINS; MAMMALS; NEOPLASMS; PROTEIN KINASES; SIGNAL TRANSDUCTION;

MAMMALIA;

EID: 48749118705     PISSN: 09501991     EISSN: None     Source Type: Journal    
DOI: 10.1242/dev.020974     Document Type: Review

References (126)

1. Abe, Y., Ohsugi, M., Haraguchi, K., Fujimoto, J. and Yamamoto, T. (2006). LATS2-Ajuba complex regulates gamma-tubulin recruitment to centrosomes and spindle organization during mitosis. FEBS Lett. 580, 782-788.
2. Adler, P. N., Chariton, J. and Liu, J. (1998). Mutations in the cadherin superfamily member gene dachsous cause a tissue polarity phenotype by altering frizzled signaling. Development 125, 959-968.
3. Baena-Lopez, L. A., Baonza, A. and Garcia-Bellido, A. (2005). The orientation of cell divisions determines the shape of Drosophila organs. Curr. Biol. 15, 1640-1644.
4. Barolo, S. and Posakony, J. W. (2002). Three habits of highly effective signaling pathways: principles of transcriptional control by developmental cell signaling. Genes Dev 16, 1167-1181.
5. Basu, S., Totty, N. F., Irwin, M. S., Sudol, M. and Downward, J. (2003). Akt phosphorylates the Yes-associated protein, YAP, to induce interaction with 14-3-3 and attenuation of p73-mediated apoptosis. Mol. Cell 11, 11-23.
6. Bennett, F. C. and Harvey, K. F. (2006). Fat cadherin modulates organ size in Drosophila via the Salvador/Warts/Hippo signaling pathway. Curr. Biol. 16, 2101-2110.
7. Boedigheimer, M. and Laughon, A. (1993). Expanded: a gene involved in the control of cell proliferation in imaginal discs. Development 118, 1291-1301.
8. Bothos, J., Tuttle, R. L., Ottey, M., Luca, F. C. and Halazonetis, T. D. (2005). Human LATS1 is a mitotic exit network kinase. Cancer Res. 65, 6568-6575.
9. Bridges, C. B. and Morgan, T. H. (1919). Contributions to the genetics of Drosophila melanogaster. II. The second-chromosome group of mutant characters. Carnegie Inst. Wash. Pub. 278, 123-304.
10. Brodsky, M. H. and Steller, H. (1996). Positional information along the dorsal-ventral axis of the Drosophila eye: graded expression of the four-jointed gene. Dev. Biol. 173, 428-446.
11. Bryant, P. J., Huettner, B., Held, L. I., Jr, Ryerse, J. and Szidonya, J. (1988). Mutations at the fat locus interfere with cell proliferation control and epithelial morphogenesis in Drosophila. Dev. Biol. 129, 541-554.
12. Buckles, G. R., Rauskolb, C., Villano, J. L. and Katz, F. N. (2001). four-jointed interacts with dachs, abelson and enabled and feeds back onto the Notch pathway to affect growth and segmentation in the Drosophila leg. Development 128, 3533-3542.
13. Callus, B. A., Verhagen, A. M. and Vaux, D. L. (2006). Association of mammalian sterile twenty kinases, Mst1 and Mst2, with hSalvaclor via C-terminal coiled-coil domains, leads to its stabilization and phosphorylation. FEBS J. 273, 4264-4276.
14. Camargo, F. D., Golkhale, S., Johnnidis, J. B., Fu, D., Bell, G. W., Jaenisch, R. and Brummelkamp, T. R. (2007). YAP1 increases organ size and expands undifferentiated progenitor cells. Curr. Biol. 17, 2054-2060.
15. Campbell, S., Inamdar, M., Rodrigues, V., Raghavan, V., Palazzolo, M. and Chovnick, A. (1992). The scalloped gene encodes a novel, evolutionarily conserved transcription factor required for sensory organ differentiation in Drosophila. Genes Dev. 6, 367-379.
16. Casal, J., Struhl, G. and Lawrence, P. (2002). Developmental compartments and planar polarity in Drosophila. Curr. Biol. 12, 1189-1198.
17. Casal, J., Lawrence, P. A. and Struhl, G. (2006). Two separate molecular systems, Dachsous/Fat and Starry night/Frizzled, act independently to confer planar cell polarity. Development 133, 4561-4572.
18. Castillejo-Lopez, C., Arias, W. M. and Baumgartner, S. (2004). The fat-like gene of Drosophila is the true orthologue of vertebrate fat cadherins and is involved in the formation of tubular organs. J. Biol. Chem. 279, 24034-24043.
19. Chan, E. H., Nousiainen, M., Chalamalasetty, R. B., Schafer, A., Nigg, E. A. and Sillje, H. H. (2005). The Ste20-like kinase Mst2 activates the human large tumor suppressor kinase Lats1. Oncogene 24, 2076-2086.
20. Cho, E. and Irvine, K. D. (2004). Action of fat, four-jointed, dachsous and dachs in distal-to-proximal wing signaling. Development 131, 4489-4500.
21. Cho, E., Feng, Y., Rauskolb, C., Maitra, S., Fehon, R. and Irvine, K. D. (2006). Delineation of a Fat tumor suppressor pathway. Nat. Genet. 38, 1142-1150.
22. Clark, H. F., Brentrup, D., Schneitz, K., Bieber, A., Goodman, C. and Noll, M. (1995). Dachsous encodes a member of the cadherin superfamily that controls imaginal disc morphogenesis in Drosophila. Genes Dev. 9, 1530-1542.
23. Dong, J., Feldmann, G., Huang, J., Wu, S., Zhang, N., Comerford, S. A., Gayyed, M. F., Anders, R. A., Maitra, A. and Pan, D. (2007). Elucidation of a universal size-control mechanism in Drosophila and mammals. Cell 130, 1120-1133.
24. Emoto, K., Parrish, J. Z., Jan, L. Y. and Jan, Y. N. (2006). The tumour suppressor Hippo acts with the NDR kinases in dendritic tiling and maintenance. Nature 443, 210-213.
25. Espanel, X. and Sudol, M. (2001). Yes-associated protein and p53-binding protein-2 interact through their WW and SH3 domains. J. Biol. Chem. 276, 14514-14523.
26. Fanto, M., Clayton, L., Meredith, J., Hardiman, K., Charroux, B., Kerridge, S. and McNeill, H. (2003). The tumor-suppressor and cell adhesion molecule Fat controls planar polarity via physical interactions with Atrophin, a transcriptional co-repressor. Development 130, 763-774.
27. Feng, Y. and Irvine, K. D. (2007). Fat and expanded act in parallel to regulate growth through warts. Proc. Natl. Acad. Sci. USA 104, 20362-20367.
28. Ferrigno, O., Lallemand, F., Verrecchia, F., L'Hoste, S., Camonis, J., Atfi, A. and Mauviel, A. (2002). Yes-associated protein (YAP65) interacts with Smad7 and potentiates its inhibitory activity against TGF-beta/Smad signaling. Oncogene 21, 4879-4884.
29. Giot, L., Bader, J. S., Brouwer, C., Chaudhurl, A., Kuang, B., Li, Y., Hao, Y. L., Ooi, C. E., Godwin, B., Vitols, E. et al. (2003). A protein interaction map of Drosophila melanogaster. Science 302, 1727-1736.
30. Glantschnig, H., Rodan, G. A. and Reszka, A. A. (2002). Mapping of MST1 kinase sites of phosphorylation. Activation and autophosphorylation. J. Biol. Chem. 277, 42987-42996.
31. Goulev, Y., Fauny, J. D., Gonzalez-Marti, B., Flagiello, D., Silber, J. and Zider, A. (2008). SCALLOPED interacts with YORKIE, the nuclear effector of the hippo tumor-suppressor pathway in Drosophila. Curr. Biol. 18, 435-441.
32. Graves, J. D., Gotoh, Y., Draves, K. E., Ambrose, D., Han, D. K., Wright, M., Chernoff, J., Clark, E. A. and Krebs, E. G. (1998). Caspase-mediated activation and induction of apoptosis by the mammalian Ste20-like kinase Mst1. EMBO J. 17, 2224-2234.
33. Graves, J. D., Draves, K. E., Gotoh, Y., Krebs, E. G. and Clark, E. A. (2001). Both phosphorylation and caspase-mediated cleavage contribute to regulation of the Ste20-like protein kinase Mst1 during CD95/Fas-induced apoptosis. J. Biol. Chem. 276, 14909-14915.
34. Haider, G. and Carroll, S. B. (2001). Binding of the Vestigial co-factor switches the DNA-target selectivity of the Scalloped selector protein. Development 128, 3295-3305.
35. Halder, G., Polaczyk, P., Kraus, M. E., Hudson, A., Kim, J., Laughon, A. and Carroll, S. (1998). The Vestigial and Scalloped proteins act together to directly regulate wing-specific gene expression in Drosophila. Genes Dev. 12, 3900-3909.
36. Hamaratoglu, F., Willecke, M., Kango-Singh, M., Nolo, R., Hyun, E., Tao, C., Jafar-Nejad, H. and Haider, G. (2006). The tumour-suppressor genes NF2/Merlin and Expanded act through Hippo signalling to regulate cell proliferation and apoptosis. Nat. Cell. Biol. 8, 27-36.
37. Hao, Y., Chun, A., Cheung, K., Rashidi, B. and Yang, X. (2008). Tumor suppressor LATS1 is a negative regulator of oncogene YAP. J. Biol. Chem. 283, 5496-5509.
38. Harvey, K. F., Pfleger, C. M. and Hariharan, I. K. (2003). The Drosophila Mst ortholog, hippo, restricts growth and cell proliferation and promotes apoptosis. Cell 114, 457-467.
39. Held, L. I., Duarte, C. M. and Derakshanian, K. (1986). Extra tarsal joints and abnormal cuticular polarities in various mutants of Drosophila melanogaster. Roux's Arch. Dev. Biol. 195, 145-157.
40. Hirabayashi, S., Nakagawa, K., Sumita, K., Hidaka, S., Kawai, T., Ikeda, M., Kawata, A., Ohno, K. and Hata, Y. (2008). Threonine 74 of MOB1 is a putative key phosphorylation site by MST2 to form the scaffold to activate nuclear Dbf2-related kinase 1. Oncogene (in press).
41. Hirota, T., Morisaki, T., Nishiyama, Y., Marumoto, T., Tada, K., Hara, T., Masuko, N., Inagaki, M., Hatakeyama, K. and Saya, H. (2000). Zyxin, a regulator of actin filament assembly, targets the mitotic apparatus by interacting with h-warts/LATS1 tumor suppressor. J. Cell Biol. 149, 1073-1086.
42. Huang, J., Wu, S., Barrera, J., Matthews, K. and Pan, D. (2005). The Hippo signaling pathway coordinately regulates cell proliferation and apoptosis by inactivating Yorkie, the Drosophila Homolog of YAP. Cell 122, 421-434.
43. Ishikawa, H. O., Takeuchi, H., Haltiwanger, R. S. and Irvine, K. D. (2008). Four-jointed is a Golgi kinase that phosphorylates a subset of cadherin domains. Science (in press).
44. Jia, J., Zhang, W., Wang, B., Trinko, R. and Jiang, J. (2003). The Drosophila Ste20 family kinase dMST functions as a tumor suppressor by restricting cell proliferation and promoting apoptosis. Genes Dev. 17, 2514-2519.
45. Justice, R. W., Zilian, O., Woods, D. F., Noll, M. and Bryant, P. J. (1995). The Drosophila tumor suppressor gene warts encodes a homolog of human myotonic dystrophy kinase and is required for the control of cell shape and proliferation. Genes Dev. 9, 534-546.
46. Kango-Singh, M., Nolo, R., Tao, C., Verstreken, P., Hiesinger, P. R., Bellen, H. J. and Haider, G. (2002). Shar-pei mediates cell proliferation arrest during imaginal disc growth in Drosophila. Development 129, 5719-5730.
47. Katoh, Y. and Katoh, M. (2006). Comparative integromics on FAT1, FAT2, FAT3 and FAT4. Int. J. Mol. Med. 18, 523-528.
48. Klein, T. J. and Mlodzik, M. (2005). Planar cell polarization: an emerging model points in the right direction. Annu. Rev. Cell Dev. Biol. 21, 155-176.
49. Komuro, A., Nagai, M., Navin, N. E. and Sudol, M. (2003). WW domain-containing protein YAP associates with ErbB-4 and acts as a co-transcriptional activator for the carboxyl-terminal fragment of ErbB-4 that translocates to the nucleus. J. Biol. Chem. 278, 33334-33341.
50. Lai, Z. C., Wei, X., Shimizu, T., Ramos, E., Rohrbaugh, M., Nikolaidis, N., Ho, L. L. and Li, Y. (2005), Control of cell proliferation and apoptosis by mob as tumor suppressor, mats. Cell 120, 675-685.
51. LaJeunesse, D. R., McCartney, B. M. and Fehon, R. G. (1998). Structural analysis of Drosophila merlin reveals functional domains important for growth control and subcellular localization. J. Cell Biol. 141, 1589-1599.
52. Lee, J. H., Kim, T. S., Yang, T. H., Koo, B. K., Oh, S. P., Lee, K. P., Oh, H. J., Lee, S. H., Kong, Y. Y., Kim, J. M. et al. (2008). A crucial role of WW45 in developing epithelial tissues in the mouse. EMBO J. 27, 1231-1242.
53. Lee, K. K. and Yonehara, S. (2002). Phosphorylation and dimerization regulate nucleocytoplasmic shuttling of mammalian STE20-like kinase (MST). J. Biol. Chem. 277, 12351-12358.
54. Lei, Q. Y., Zhang, H., Zhao, B., Zha, Z. Y., Bai, F., Pei, X. H., Zhao, S., Xiong, Y. and Guan, K. L. (2008). TAZ promotes cell proliferation and epithelial-mesenchymal transition and is inhibited by the hippo pathway. Mol. Cell. Biol. 28, 2426-2436.
55. Liu, X., Grammont M. and Irvine, K. D. (2000). Roles for scalloped and vestigial in regulating cell affinity and interactions between the wing blade and the wing hinge. Dev. Biol. 228, 287-303.
56. Ma, D., Yang, C. H., McNeill, H., Simon, M. A. and Axelrod, J. D. (2003). Fidelity in planar cell polarity signalling. Nature 421, 543-547.
57. MacDougall, N., Lad, Y., Wilkie, G. S., Francis-Lang, H., Sullivan, W. and Davis, I. (2001). Merlin, the Drosophila homologue of neurofibromatosis-2, is specifically required in posterior follicle cells for axis formation in the oocyte. Development 128, 665-673.
58. Mackintosh, C. (2004). Dynamic interactions between 14-3-3 proteins and phosphoproteins regulate diverse cellular processes. Biochem. J. 381, 329-342.
59. Mahoney, P. A., Weber, U., Onofrechuk, P., Biessmann, H., Bryant, P. J. and Goodman, C. S. (1991). The fat tumor suppressor gene in Drosophila encodes a novel member of the cadherin gene superfamily. Cell 67, 853-868.
60. Mahoney, W. M., Jr, Hong, J. H., Yaffe, M. B. and Farrance, I. K. (2005). The transcriptional co-activator TAZ interacts differentially with transcriptional enhancer factor-1 (TEF-1) family members. Biochem. J. 388, 217-225.
61. Maitra, S., Kulikauskas, R. M., Gavilan, H. and Fehon, R. G. (2006). The tumor suppressors Merlin and expanded function cooperatively to modulate receptor endocytosis and signaling. Curr. Biol. 16, 702-709.
62. Mangeat, P., Roy, C. and Martin, M. (1999). ERM proteins in cell adhesion and membrane dynamics. Trends Cell. Biol. 9, 187-192.
63. Mao, Y., Rauskolb, C., Cho, E., Hu, W. L., Hayter, H., Minihan, G., Katz, F. N. and Irvine, K. D. (2006). Dachs: an unconventional myosin that functions downstream of Fat to regulate growth, affinity and gene expression in Drosophila. Development 133, 2539-2551.
64. Matakatsu, H. and Blair, S. S. (2004). Interactions between Fat and Dachsous and the regulation of planar cell polarity in the Drosophila wing. Development 131, 3785-3794.
65. Matakatsu, H. and Blair, S. S. (2006). Separating the adhesive and signaling functions of the Fat and Dachsous protocadherins. Development 133, 2315-2324.
66. McCartney, B. M., Kulikauskas, R. M., LaJeunesse, D. R. and Fehon, R. G. (2000). The neurofibromatosis-2 homologue, Merlin, and the tumor suppressor expanded function together in Drosophila to regulate cell proliferation and differentiation. Development 127, 1315-1324.
67. McClatchey, A. I. and Giovannini, M. (2005). Membrane organization and tumorigenesis-the NF2 tumor suppressor, Merlin. Genes Dev 19, 2265-2277.
68. McPherson, J. P, Tamblyn, L., Elia, A., Migon, E., Shehabeldin, A., Matysiak-Zablocki, E., Lemmers, B., Salmena, L., Hakem, A., Fish, J. et al. (2004). Lats2/Kpm is required for embryonic development, proliferation control and genomic integrity. EMBO J. 23, 3677-3688.
69. Meignin, C., Alvarez-Garcia, I., Davis, I. and Palacios, I. M. (2007). The salvador-warts-hippo pathway is required for epithelial proliferation and axis specification in Drosophila. Curr. Biol. 17, 1871-1878.
70. Mikeladze-Dvali, T., Wernet, M. F., Pistillo, D., Mazzoni, E. O., Teleman, A. A., Chen, Y. W., Cohen, S. and Desplan, C. (2005). The growth regulators warts/lats and melted interact in a bistable loop to specify opposite fates in Drosophila R8 photoreceptors. Cell 122, 775-787.
71. Mohr, O. L. (1923), Modifications of the sex-ratio through a sex-linked semi-lethal in Drosophila melanogaster. (Besides notes on an autosomal section deficiency.) In Studia Mendeliana, pp. 266-287. Brunn, Czechoslovakia: Apud Typos.
72. Morisaki, T., Hirota, T., Iida, S., Marumoto, T., Hara, T., Nishiyama, Y., Kawasuzi, M., Hiraoka, T., Mimori, T., Araki, N. et al. (2002). WARTS tumor suppressor is phosphorylated by Cdc2/cyclin B at spindle poles during mitosis. FEBS Lett. 529, 319-324.
73. Morrison, H., Sherman, L. S., Legg, J., Banine, F., Isacke, C., Haipek, C. A., Gutmann, D. H., Ponta, H. and Herrlich, P. (2001). The NF2 tumor suppressor gene product, merlin, mediates contact inhibition of growth through interactions with CD44. Genes Dev. 15, 968-980.
74. Nishiyama, Y., Hirota, T., Morisaki, T., Hara, T., Marumoto, T., Iida, S., Makino, K., Yamamoto, H., Hiraoka, T,, Kitamura, N. et al. (1999). A human homolog of Drosophila warts tumor suppressor, h-warts, localized to mitotic apparatus and specifically phosphorylated during mitosis. FEBS Lett. 459, 159-165.
75. Nolo, R., Morrison, C. M., Tao, C., Zhang, X. and Halder, G. (2006). The bantam microRNA is a target of the hippo tumor-suppressor pathway. Curr. Biol. 16, 1895-1904.
76. Oh, H. and Irvine, K. D. (2008). In vivo regulation of Yorkie phosphorylation and localization. Development 135, 1081-1088.
77. Overholtzer, M., Zhang, J., Smolen, G. A., Muir, B., Li, W., Sgroi, D. C., Deng, C. X., Brugge, J. S. and Haber, D. A. (2006). Transforming properties of YAP, a candidate oncogene on the chromosome 11q22 amplicon. Proc. Natl. Acad. Sci. USA 103, 12405-12410.
78. Pantalacci, S., Tapon, N. and Leopold, P. (2003). The Salvador partner Hippo promotes apoptosis and cell-cycle exit in Drosophila. Nat. Cell Biol. 5, 921-927.
79. Paumard-Rigal, S., Zider, A., Vaudin, P. and Silber, J. (1998). Specific interactions between vestigial and scalloped are required to promote wing tissue proliferation in Drosophila melanogaster. Dev Genes Evol. 208, 440-446.
80. Pellock, B. J., Buff, E., White, K. and Hariharan, I. K. (2007). The Drosophila tumor suppressors Expanded and Merlin differentially regulate cell cycle exit, apoptosis, and Wingless signaling. Dev Biol. 304, 102-115.
81. Polesello, C. and Tapon, N. (2007). Salvador-warts-hippo signaling promotes Drosophila posterior follicle cell maturation downstream of notch. Curr. Biol. 17, 1864-1870.
82. Polesello, C., Huelsmann, S., Brown, N. H. and Tapon, N. (2006). The Drosophila RASSF homolog antagonizes the hippo pathway. Curr. Biol. 16, 2459-2465.
83. Praskova, M., Khoklatchev, A., Ortiz-Vega, S. and Avruch, J. (2004). Regulation of the MST1 kinase by autophosphorylation, by the growth inhibitory proteins, RASSF1 and NORE1, and by Ras. Biochem. J. 381, 453-462.
84. Praskova, M., Xia, F. and Avruch, J. (2008). MOBKL1A/MOBKL1B phosphorylation by MST1 and MST2 inhibits cell proliferation. Curr. Biol. 18, 311-321.
85. Rawls, A. S., Guinto, J. B. and Wolff, T. (2002). The cadherins fat and dachsous regulate dorsal/ventral signaling in the Drosophila eye. Curr. Biol. 12, 1021-1026.
86. Rock, R., Schrauth, S. and Gessler, M. (2005). Expression of mouse dchs1, fjx1, and fat-j suggests conservation of the planar cell polarity pathway identified in Drosophila. Dev. Dyn. 234, 747-755.
87. Rogulja, D., Rauskolb, C. and Irvine, K. D. (2008). Morphogen control of wing growth through the Fat signaling pathway. Dev Cell (in press).
88. Saburi, S., Hester, I., Eremina, V., Fischer, E., Pontoglio, M., Gessler, M., Quaggin, S., Harrison, R., Mount, R. and McNeill, H. (2008). Loss of Fat4 disrupts PCP signalling and oriented cell division, leading to cystic kidney disease. Nat. Genet (in press).
89. Seidel, C., Schagdarsurengin, U., Blumke, K., Wurl, R, Pfeifer, G. P., Hauptmann, S., Taubert, H. and Dammann, R. (2007). Frequent hypermethylation of MST1 and MST2 in soft tissue sarcoma. Mol. Carcinog. 46, 865-871.
90. Shimizu, T., Ho, L. L. and Lai, Z. C. (2008). The mob as tumor suppressor gene is essential for early development and regulates tissue growth in Drosophila. Genetics 178, 957-965.
91. Silva, E., Tsatskis, Y., Gardano, L., Tapon, N. and McNeill, H. (2006). The tumor-suppressor gene fat controls tissue growth upstream of expanded in the hippo signaling pathway. Curr. Biol. 16, 2081-2089.
92. Simmonds, A. J., Liu, X., Soanes, K. H., Krause, H. M., Irvine, K. D. and Bell, J. B. (1998). Molecular interactions between Vestigial and Scalloped promote wing formation in Drosophila. Genes Dev. 12, 3815-3820.
93. Simon, M. A. (2004). Planar cell polarity in the Drosophila eye is directed by graded Four-jointed and Dachsous expression. Development 131, 6175-6184.
94. St John, M. A., Tao, W., Fei, X., Fukumoto, R., Carcangiu, M. L., Brownstein, D. G., Parlow, A. F., McGrath, J. and Xu, T. (1999). Mice deficient of Lats1 develop soft-tissue sarcomas, ovarian tumours and pituitary dysfunction. Nat. Genet. 21, 182-186.
95. Strano, S., Munarriz, E., Rossi, M., Castagnoli, L., Shaul, Y., Sacchi, A., Oren, M., Sudol, M., Cesareni, G. and Blandino, G. (2001). Physical interaction with Yes-associated protein enhances p73 transcriptional activity. J. Biol. Chem. 276, 15164-15173.
96. Strutt, H. and Strutt, D. (2002). Nonautonomous planar polarity patterning in Drosophila: dishevelled-independent functions of frizzled. Dev. Cell 3, 851-863.
97. Strutt, H., Mundy, J., Hofstra, K. and Strutt, D. (2004). Cleavage and secretion is not required for Four-jointed function in Drosophila patterning. Development 131, 881-890.
98. Takahashi, Y., Miyoshi, Y., Takahata, C., Irahara, N., Taguchi, T., Tamaki, Y. and Noguchi, S. (2005). Down-regulation of LATS1 and LATS2 mRNA expression by promoter hypermethylation and its association with biologically aggressive phenotype in human breast cancers. Clin. Cancer Res. 11, 1380-1385.
99. Tao, W., Zhang, S., Turenchalk, G. S., Stewart, R. A., St John, M. A., Chen, W. and Xu, T. (1999). Human homologue of the Drosophila melanogaster lats tumour suppressor modulates CDC2 activity. Nat. Genet. 21, 177-181.
100. Tapon, N., Harvey, K. F., Bell, D. W., Wahrer, D. C., Schiripo, T. A., Haber, D. A. and Hariharan, I. K. (2002). salvador promotes both cell cycle exit and apoptosis in Drosophila and is mutated in human cancer cell lines. Cell 110, 467-478.
101. Thompson, B. J. and Cohen, S. M. (2006). The Hippo pathway regulates the bantam microRNA to control cell proliferation and apoptosis in Drosophila. Cell 126, 767-774.
102. Toji, S., Yabuta, N., Hosomi, T., Nishihara, S., Kobayashi, T., Suzuki, S., Tamai, K. and Nojima, H. (2004). The centrosomal protein Lats2 is a phosphorylation target of Aurora-A kinase. Genes Cells 9, 383-397.
103. Tyler, D. M. and Baker, N. E. (2007). Expanded and fat regulate growth and differentiation in the Drosophila eye through multiple signaling pathways. Dev. Biol. 305, 187-201.
104. Udan, R. S., Kango-Singh, M., Nolo, R., Tao, C. and Halder, G. (2003). Hippo promotes proliferation arrest and apoptosis in the Salvador/Warts pathway. Nat. Cell. Biol. 5, 914-920.
105. Vassilev, A., Kaneko, K. J., Shu, H., Zhao, Y. and DePamphilis, M. L. (2001). TEAD/TEF transcription factors utilize the activation domain of YAP65, a Src/Yes-associated protein localized in the cytoplasm. Genes Dev. 15, 1229-1241.
106. Vaudin, P., Delanoue, R., Davidson, I., Silber, J. and Zider, A. (1999). TONDU (TDU), a novel human protein related to the product of vestigial (vg) gene of Drosophila melanogaster interacts with vertebrate TEF factors and substitutes for Vg function in wing formation. Development 126, 4807-4816.
107. Villano, J. L. and Katz, F. N. (1995). four-jointed is required for intermediate growth in the proximal-distal axis in Drosophila. Development 121, 2767-2777.
108. Waddington, C. H. (1940). The genetic control of wing development in Drosophila. J. Genet. 41, 75-139.
109. Wei, X., Shimizu, T. and Lai, Z. C. (2007). Mob as tumor suppressor is activated by Hippo kinase for growth inhibition in Drosophila. EMBO J. 26, 1772-1781.
110. Willecke, M., Hamaratoglu, F., Kango-Singh, M., Udan, R., Chen, C. L., Tao, C., Zhang, X. and Halder, G. (2006). The fat cadherin acts through the hippo tumor-suppressor pathway to regulate tissue size. Curr. Biol. 16, 2090-2100.
111. Wu, S., Huang, J., Dong, J. and Pan, D. (2003). hippo encodes a Ste-20 family protein kinase that restricts cell proliferation and promotes apoptosis in conjunction with salvador and warts. Cell 114, 445-456.
112. Wu, S., Liu, Y., Zheng, Y., Dong, J. and Pan, D. (2008). The TEAD/ TEF family protein Scalloped mediates transcriptional output of the Hippo growth-regulatory pathway. Dev. Cell 14, 388-398.
113. Xu, T., Wang, W., Zhang, S., Stewart, R. A. and Yu, W. (1995). Identifying tumor suppressors in genetic mosaics: the Drosophila lats gene encodes a putative protein kinase. Development 121, 1053-1063.
114. Yagi, R., Chen, L. F., Shigesada, K., Murakami, Y. and Ito, Y. (1999). A WW domain-containing yes-associated protein (YAP) is a novel transcriptional coactivator. EMBO J. 18, 2551-2562.
115. Yang, C., Axelrod, J. D. and Simon, M. A. (2002). Regulation of Frizzled by Fat-like cadherins during planar polarity signaling in the Drosophila compound eye. Cell 108, 675-688.
116. Yang, X., Yu, K., Hao, Y., Li, D. M., Stewart, R., Insogna, K. L. and Xu, T. (2004). LATS1 tumour suppressor affects cytokinesis by inhibiting LIMK1. Nat. Cell Biol. 6, 609-617.
117. Yu, J., Poulton, J., Huang, Y. C. and Deng, W. M. (2008). The hippo pathway promotes Notch signaling in regulation of cell differentiation, proliferation, and oocyte polarity. PLoS ONE 3, e1761.
118. Zaidi, S. K., Sullivan, A. J., Medina, R., Ito, Y., van Wijnen, A. J., Stein, J. L., Lian, J. B. and Stein, G. S. (2004). Tyrosine phosphorylation controls Runx2-mediated subnuclear targeting of YAP to repress transcription. EMBO J. 23, 790-799.
119. Zeidler, M. P., Perrimon, N. and Strutt, D. I. (1999). The four-jointed gene is required in the Drosophila eye for ommatidial polarity specification. Curr. Biol. 9, 1363-1372.
120. Zeidler, M. P., Perrimon, N. and Strutt, D. I. (2000). Multiple roles for four-jointed in planar polarity and limb patterning. Dev Biol. 228, 181-196.
121. Zender, L., Spector, M. S., Xue, W., Flemming, P., Cordon-Cardo, C., Silke, J., Fan, S. T., Luk, J. M., Wigler, M., Hannon, G. J. et al. (2006). Identification and validation of oncogenes in liver cancer using an integrative oncogenomic 000000000approach. Cell 125, 1253-1267.
122. Zhang, J., Smolen, G. A. and Haber, D. A. (2008a). Negative regulation of YAP by LATS1 underscores evolutionary conservation of the Drosophila Hippo pathway. Cancer Res. 68, 2789-2794.
123. Zhang, L., Ren, F., Zhang, Q., Chen, Y., Wang, B. and Jiang, J. (2008b). The TEAD/TEF family of transcription factor Scalloped mediates Hippo signaling in organ size control. Dev Cell 14, 377-387.
124. Zhao, B., Wei, X., Li, W., Udan, R. S., Yang, Q., Kim, J., Xie, J., Ikenoue, T., Yu, J., Li, L. et al. (2007). Inactivation of YAP oncoprotein by the Hippo pathway is involved in cell contact inhibition and tissue growth control. Genes Dev. 21, 2747-2761.
125. Zhao, B., Ye, X., Yu, J., Li, L., Li, W., Li, S., Yu, J., Lin, J., Wang, C.-Y., Chinnaiyan, A. M. et al. (2008). TEAD mediates YAP dependent gene induction and growth control. Genes Dev. (in press).
126. Zilian, O., Frei, E., Burke, R., Brentrup, D., Gutjahr, T., Bryant, P. J. and Noll, M. (1999). double-time is identical to discs overgrown, which is required for cell survival, proliferation and growth arrest in Drosophila imaginal discs. Development 126, 5409-5420.