Planar cell polarity in drosophila

Organogenesis

Volumn 7, Issue 3, 2011, Pages

  Maung, Saw Myat Thanda W ^a   Jenny, Andreas ^a  

^a ALBERT EINSTEIN COLLEGE OF MEDICINE   (United States)

Author keywords

Abdomen; Dachsous; Drosophila; Eye; Fat; Fizzled; Non canonical Wnt signaling; Planar cell polarity; Protein asymmetry; Wing

Indexed keywords

ABDOMEN; CELL CONTACT; CELL POLARITY; DROSOPHILA MELANOGASTER; NONHUMAN; PHENOTYPE; PLANAR CELL POLARITY; PROTEIN LOCALIZATION; PROTEIN PROTEIN INTERACTION; REVIEW; SENSE ORGAN; SIGNAL TRANSDUCTION; ANIMAL; BIOLOGICAL MODEL; CYTOLOGY; CYTOSKELETON; DROSOPHILA; EPITHELIUM; GASTRULATION; HUMAN; METABOLISM; PHYSIOLOGY; STEREOCILIUM;

DROSOPHILA MELANOGASTER; VERTEBRATA;

DROSOPHILA PROTEIN;

ANIMALS; CELL POLARITY; CYTOSKELETON; DROSOPHILA; DROSOPHILA PROTEINS; EPITHELIUM; GASTRULATION; HUMANS; MODELS, BIOLOGICAL; PHENOTYPE; STEREOCILIA;

EID: 80054680272     PISSN: 15476278     EISSN: 15558592     Source Type: Journal    
DOI: 10.4161/org.7.3.18143     Document Type: Review

References (250)

1. Eaton S, Simons K. Apical, basal and lateral cues for epithelial polarization. Cell 1995; 82:5-8; PMID:7606785; DOI:10.1016/0092-8674(95)90045-4.
2. Wodarz A, Nathke I. Cell polarity in development and cancer. Nat Cell Biol 2007; 9:1016-24; PMID:17762893; DOI:10.1038/ncb433.
3. Lecuit T, Lenne PF. Cell surface mechanics and the control of cell shape, tissue patterns and morphogenesis. Nat Rev Mol Cell Biol 2007; 8:633-44; PMID:17643125; DOI:10.1038/nrm2222.
4. Zallen JA, Blankenship JT. Multicellular dynamics during epithelial elongation. Semin Cell Dev Biol 2008; 19:263-70; PMID:18343171; DOI:10.1016/j. semcdb.2008.01.005.
5. Henderson DJ, Chaudhry B. Getting to the heart of planar cell polarity signaling. Birth Defects Res A Clin Mol Teratol 2011; 91:460-7; PMID:21538810; DOI:10.1002/bdra.20792.
6. Vandenberg AL, Sassoon DA. Non-canonical Wnt signaling regulates cell polarity in female reproductive tract development via van gogh-like 2. Development 2009; 136:1559-70; PMID:19363157; DOI:10.1242/ dev.034066.
7. Yates LL, Papakrivopoulou J, Long DA, Goggolidou P, Connolly JO, Woolf AS, et al. The planar cell polarity gene Vangl2 is required for mammalian kidneybranching morphogenesis and glomerular maturation. Hum Mol Genet 2010; 19:4663-76; PMID:20843830; DOI:10.1093/hmg/ddq397.
8. Yates LL, Schnatwinkel C, Murdoch JN, Bogani D, Formstone CJ, Townsend S, et al. The PCP genes Celsr1 and Vangl2 are required for normal lung branching morphogenesis. Hum Mol Genet 2010; 19:2251- 67; PMID:20223754; DOI:10.1093/hmg/ddq104.
9. Karner CM, Chirumamilla R, Aoki S, Igarashi P, Wallingford JB, Carroll TJ. Wnt9b signaling regulates planar cell polarity and kidney tubule morphogenesis. Nat Genet 2009; 41:793-9; PMID:19543268; DOI:10.1038/ng.400.
10. Gray RS, Roszko I, Solnica-Krezel L. Planar cell polarity: coordinating morphogenetic cell behaviors with embryonic polarity. Dev Cell 2011; 21:120-33; PMID:21763613; DOI:10.1016/j.devcel. 2011.06.011.
11. Copp AJ, Greene ND, Murdoch JN. The genetic basis of mammalian neurulation. Nat Rev Genet 2003; 4:784-93; PMID:13679871; DOI:10.1038/nrg1181.
12. Doudney K, Stanier P. Epithelial cell polarity genes are required for neural tube closure. Am J Med Genet C Semin Med Genet 2005; 135:42-7; PMID:15800847; DOI:10.1002/ajmg.c.30052.
13. Ross AJ, May-Simera H, Eichers ER, Kai M, Hill J, Jagger DJ, et al. Disruption of Bardet-Biedl syndrome ciliary proteins perturbs planar cell polarity in vertebrates. Nat Genet 2005; 37:1135-40; PMID:16170314; DOI:10.1038/ng1644.
14. Simons M, Mlodzik M. Planar Cell Polarity Signaling: From Fly Development to Human Disease. Annu Rev Genet 2008; 42:517-40; PMID:18710302; DOI:10.1146/annurev.genet.42.110807.091432.
15. Qian D, Jones C, Rzadzinska A, Mark S, Zhang X, Steel KP, et al. Wnt5a functions in planar cell polarity regulation in mice. Dev Biol 2007; 306:121-33; PMID:17433286; DOI:10.1016/j.ydbio.2007.03.011.
16. Wang J, Mark S, Zhang X, Qian D, Yoo SJ, Radde- Gallwitz K, et al. Regulation of polarized extension and planar cell polarity in the cochlea by the vertebrate PCP pathway. Nat Genet 2005; 37:980-5; PMID:16116426; DOI:10.1038/ng1622.
17. Wang Y, Guo N, Nathans J. The role of Frizzled3 and Frizzled6 in neural tube closure and in the planar polarity of inner-ear sensory hair cells. J Neurosci 2006; 26:2147-56; PMID:16495441; DOI:10.1523/ JNEUROSCI.4698-05.2005.
18. Wu J, Mlodzik M. A quest for the mechanism regulating global planar cell polarity of tissues. Trends Cell Biol 2009; 19:295-305; PMID:19560358; DOI:10.1016/j. tcb.2009.04.003.
19. Caddy J, Wilanowski T, Darido C, Dworkin S, Ting SB, Zhao Q, et al. Epidermal wound repair is regulated by the planar cell polarity signaling pathway. Dev Cell 2010; 19:138-47; PMID:20643356; DOI:10.1016/j. devcel.2010.06.008.
20. Lawrence PA, Shelton PMJ. The determination of polarity in the developing insect retina. J Embryol Exp Morphol 1975; 33:471-86; PMID:1176856.
21. Vinson CR, Adler PN. Directional non-cell autonomy and the transmission of polarity information by the frizzled gene of Drosophila. Nature 1987; 329:549-51; PMID:3116434; DOI:10.1038/329549a0.
22. Adler P. Planar polarity in the Drosophila wing. In: Mlodzik M, Ed. Planar cell polarization during development. San Diego: Elsevier 2005; 1-14.
23. Aldaz S, Escudero LM, Freeman M. Live imaging of Drosophila imaginal disc development. Proc Natl Acad Sci USA 2010; 107:14217-22; PMID:20660765; DOI:10.1073/pnas.1008623107.
24. Garcia-Bellido A, Merriam JR. Parameters of the wing imaginal disc development of Drosophila melanogaster. Dev Biol 1971; 24:61-87; PMID:5001010; DOI:10.1016/0012-1606(71)90047-9.
25. Schubiger M, Palka J. Changing spatial patterns of DNA replication in the developing wing of Drosophila. Dev Biol 1987; 123:145-53; PMID:3622926; DOI:10.1016/0012-1606(87)90436-2.
26. Aigouy B, Farhadifar R, Staple DB, Sagner A, Roper JC, Julicher F, et al. Cell flow reorients the axis of planar polarity in the wing epithelium of Drosophila. Cell 2010; 142:773-86; PMID:20813263; DOI:10.1016/j. cell.2010.07.042.
27. Guild GM, Connelly PS, Ruggiero L, Vranich KA, Tilney LG. Actin filament bundles in Drosophila wing hairs: hairs and bristles use different strategies for assembly. Mol Biol Cell 2005; 16:3620-31; PMID:15917291; DOI:10.1091/mbc.E05-03-0185.
28. Mitchell HK, Roach J, Petersen NS. The morphogenesis of cell hairs on Drosophila wings. Dev Biol 1983; 95:387-98; PMID:6402398; DOI:10.1016/0012- 1606(83)90040-4.
29. Eaton S, Wepf R, Simons K. Roles for Rac1 and Cdc42 in planar polarization and hair outgrowth in the wing of Drosophila. J Cell Biol 1996; 135:1277-89; PMID:8947551; DOI:10.1083/jcb.135.5.1277.
30. Turner CM, Adler PN. Distinct roles for the actin and microtubule cytoskeletons in the morphogenesis of epidermal hairs during wing development in Drosophila. Mech Dev 1998; 70:181-92; PMID:9510034; DOI:10.1016/S0925-4773(97) 00194-9.
31. Wong LL, Adler PN. Tissue polarity genes of Drosophila regulate the subcellular location for prehair initiation in pupal wing cells. J Cell Biol 1993; 123:209-21; PMID:8408199; DOI:10.1083/jcb.123.1.209.
32. Bellaiche Y, Schweisguth F. Lineage diversity in the Drosophila nervous system. Curr Opin Genet Dev 2001; 11:418-23; PMID:11448628; DOI:10.1016/ S0959-437X(00)00212-4.
33. Gonczy P. Mechanisms of asymmetric cell division: flies and worms pave the way. Nat Rev Mol Cell Biol 2008; 9:355-66; PMID:18431399; DOI:10.1038/nrm2388.
34. Bellaiche Y, Beaudoin-Massiani O, Stuttem I, Schweisguth F. The planar cell polarity protein Strabismus promotes Pins anterior localization during asymmetric division of sensory organ precursor cells in Drosophila. Development 2004; 131:469-78; PMID:14701683; DOI:10.1242/dev.00928.
35. Bellaiche Y, Gho M, Kaltschmidt JA, Brand AH, Schweisguth F. Frizzled regulates localization of cellfate determinants and mitotic spindle rotation during asymmetric cell division. Nat Cell Biol 2001; 3:50-7; PMID:11146626; DOI:10.1038/35050558.
36. Segalen M, Johnston CA, Martin CA, Dumortier JG, Prehoda KE, David NB, et al. The Fz-Dsh planar cell polarity pathway induces oriented cell division via Mud/NuMA in Drosophila and zebrafish. Dev Cell 2010; 19:740-52; PMID:21074723; DOI:10.1016/j. devcel.2010.10.004.
37. Olguín PGA, Mlodzik M. Intertissue mechanical stress affects Frizzled-mediated planar cell polarity in the Drosophila notum epidermis. Curr Biol 2011; 21:236-42; PMID:21276726; DOI:10.1016/j.cub.2011.01.001.
38. Lawrence PA, Johnston P. Cell lineage of the Drosophila abdomen: the epidermis, oenocytes and ventral muscles. J Embryol Exp Morphol 1982; 72:197-208; PMID:7183739.
39. Jenny A. Planar cell polarity signaling in the Drosophila eye. Curr Top Dev Biol 2010; 93:189-227; PMID:20959167; DOI:10.1016/B978-0-12-385044-7.00007- 2.
40. Seifert JR, Mlodzik M. Frizzled/PCP signalling: a conserved mechanism regulating cell polarity and directed motility. Nat Rev Genet 2007; 8:126-38; PMID:17230199; DOI:10.1038/nrg2042.
41. Cagan R. Principles of Drosophila eye differentiation. Curr Top Dev Biol 2009; 89:115-35; PMID:19737644; DOI:10.1016/S0070-2153(09)89005-4.
42. Singh A, Lim J, Choi K. Dorsoventral boundary for organizing growth and planar polarity in the Drosophila eye. In: Mlodzik M, Ed. Planar cell polarization during development. San Diego: Elsevier 2005; 59-90. (Pubitemid 44725679)
43. Wehrli M, Tomlinson A. Independent regulation of anterior/posterior and equatorial/polar polarity in the Drosophila eye; evidence for the involvement of Wnt signaling in the equatorial/polar axis. Development 1998; 125:1421-32; PMID:9502723.
44. Tomlinson A, Ready DF. Neuronal differentiation in the Drosophila ommatidium. Dev Biol 1987; 120:366-76; PMID:17985475; DOI:10.1016/0012-1606(87) 90239-9.
45. Baker NE, Mlodzik M, Rubin GM. Spacing differentiation in the developing Drosophila eye: a fibrinogenrelated lateral inhibitor encoded by scabrous. Science 1990; 250:1370-7; PMID:2175046; DOI:10.1126/science.2175046.
46. Baker NE, Zitron AE. Drosophila eye development: Notch and Delta amplify a neurogenic pattern conferred on the morphogenetic furrow by scabrous. Mech Dev 1995; 49:173-89; PMID:7734391; DOI:10.1016/0925-4773(94)00314-D.
47. Baker NE, Sung Y, Han D. Evolution of proneural atonal expression during distinct regulatory phases in the developing Drosophila eye. Curr Biol 1996; 6:1290-301; PMID:8939576; DOI:10.1016/S0960- 9822(02)70715-X.
48. Roignant JY, Treisman JE. Pattern formation in the Drosophila eye disc. Int J Dev Biol 2009; 53:795-804; PMID:19557685; DOI:10.1387/ijdb.072483jr.
49. Casal J, Lawrence PA, Struhl G. Two separate molecular systems, Dachsous/Fat and Starry night/Frizzled, act independently to confer planar cell polarity. Development 2006; 133:4561-72; PMID:17075008; DOI:10.1242/dev.02641.
50. Lawrence PA, Struhl G, Casal J. Planar cell polarity: one or two pathways? Nat Rev Genet 2007; 8:555-63; PMID:17563758; DOI:10.1038/nrg2125.
51. Rawls AS, Guinto JB, Wolff T. The cadherins fat and dachsous regulate dorsal/ventral signaling in the Drosophila eye. Curr Biol 2002; 12:1021- 6; PMID:12123577; DOI:10.1016/S0960- 9822(02)00893-X.
52. Strutt H, Strutt D. Nonautonomous planar polarity patterning in Drosophila: dishevelled-independent functions of frizzled. Dev Cell 2002; 3:851- 63; PMID:12479810; DOI:10.1016/S1534- 5807(02)00363-5.
53. Yang C, Axelrod JD, Simon MA. Regulation of Frizzled by Fat-like Cadherins during Planar Polarity Signaling in the Drosophila Compound Eye. Cell 2002; 108:675-88; PMID:11893338; DOI:10.1016/ S0092-8674(02)00658-X.
54. Zeidler MP, Perrimon N, Strutt DI. The fourjointed gene is required in the Drosophila eye for ommatidial polarity specification. Curr Biol 1999; 9:1363-72; PMID:10607560; DOI:10.1016/S0960- 9822(00)80081-0.
55. Clevers H. Wnt/beta-catenin signaling in development and disease. Cell 2006; 127:469-80; PMID:17081971; DOI:10.1016/j.cell.2006.10.018.
56. Heisenberg CP, Tada M, Rauch GJ, Saude L, Concha ML, Geisler R, et al. Silberblick/Wnt11 mediates convergent extension movements during zebrafish gastrulation. Nature 2000; 405:76-81; PMID:10811221; DOI:10.1038/35011068.
57. Gros J, Serralbo O, Marcelle C. WNT11 acts as a directional cue to organize the elongation of early muscle fibres. Nature 2009; 457:589-93; PMID:18987628; DOI:10.1038/nature07564.
58. Chen WS, Antic D, Matis M, Logan CY, Povelones M, Anderson GA, et al. Asymmetric homotypic interactions of the atypical cadherin flamingo mediate intercellular polarity signaling. Cell 2008; 133:1093-105; PMID:18555784; DOI:10.1016/j.cell.2008.04.048.
59. Lawrence PA, Casal J, Struhl G. Towards a model of the organisation of planar polarity and pattern in the Drosophila abdomen. Development 2002; 129:2749-60; PMID:12015301.
60. Lim J, Norga KK, Chen Z, Choi KW. Control of planar cell polarity by interaction of DWnt4 and fourjointed. Genesis 2005; 42:150-61; PMID:15986451; DOI:10.1002/gene.20142.
61. Chae J, Kim MJ, Goo JH, Collier S, Gubb D, Charlton J, et al. The Drosophila tissue polarity gene starry night encodes a member of the protocadherin family. Development 1999; 126:5421-9; PMID:10556066.
62. Feiguin F, Hannus M, Mlodzik M, Eaton S. The Ankyrin repeat protein Diego mediates Frizzleddependent planar polarization. Dev Cell 2001; 1:93-101; PMID:11703927; DOI:10.1016/S1534- 5807(01)00010-7.
63. Gubb D, García-Bellido A. A genetic analysis of the determination of cuticular polarity during development in Drosophila melanogaster. J Embryol Exp Morphol 1982; 68:37-57; PMID:6809878.
64. Gubb D, Green C, Huen D, Coulson D, Johnson G, Tree D, et al. The balance between isoforms of the prickle LIM domain protein is critical for planar polarity in Drosophila imaginal discs. Genes Dev 1999; 13:2315-27; PMID:10485852; DOI:10.1101/ gad.13.17.2315.
65. Krasnow RE, Adler PN. A single frizzled protein has a dual role in tissue polarity. Development 1994; 120:1883-93; PMID:7924994.
66. Krasnow RE, Wong LL, Adler PN. dishevelled is a component of the frizzled signalling pathway in Drosophila. Development 1995; 121:4095-102; PMID:8575310.
67. Taylor J, Abramova N, Charlton J, Adler PN. Van Gogh: a new Drosophila tissue polarity gene. Genetics 1998; 150:199-210; PMID:9725839.
68. Usui T, Shima Y, Shimada Y, Hirano S, Burgess RW, Schwarz TL, et al. Flamingo, a seven-pass transmembrane cadherin, regulates planar cell polarity under the control of Frizzled. Cell 1999; 98:585-95; PMID:10490098; DOI:10.1016/S0092- 8674(00)80046-X.
69. Vinson CR, Conover S, Adler PN. A Drosophila tissue polarity locus encodes a protein containing seven potential transmembrane domains. Nature 1989; 338:263-4; PMID:2493583; DOI:10.1038/338263a0.
70. Wolff T, Rubin GM. strabismus, a novel gene that regulates tissue polarity and cell fate decisions in Drosophila. Development 1998; 125:1149-59; PMID:9463361.
71. Zheng L, Zhang J, Carthew RW. frizzled regulates mirror- symmetric pattern formation in the Drosophila eye. Development 1995; 121:3045-55; PMID:7555730.
72. Lawrence PA, Casal J, Struhl G. Cell interactions and planar polarity in the abdominal epidermis of Drosophila. Development 2004; 131:4651-64; PMID:15329345; DOI:10.1242/dev.01351.
73. Adler PN, Taylor J, Charlton J. The domineering non-autonomy of frizzled and van Gogh clones in the Drosophila wing is a consequence of a disruption in local signaling. Mech Dev 2000; 96:197- 207; PMID:10960784; DOI:10.1016/S0925- 4773(00)00392-0.
74. Amonlirdviman K, Khare NA, Tree DR, Chen WS, Axelrod JD, Tomlin CJ. Mathematical modeling of planar cell polarity to understand domineering nonautonomy. Science 2005; 307:423-6; PMID:15662015; DOI:10.1126/science. 1105471.
75. Le Garrec JF, Lopez P, Kerszberg M. Establishment and maintenance of planar epithelial cell polarity by asymmetric cadherin bridges: a computer model. Dev Dyn 2006; 235:235-46; PMID:16258926; DOI:10.1002/ dvdy.20617.
76. Ciruna B, Jenny A, Lee D, Mlodzik M, Schier AF. Planar cell polarity signalling couples cell division and morphogenesis during neurulation. Nature 2006; 439:220-4; PMID:16407953; DOI:10.1038/ nature04375.
77. Devenport D, Fuchs E. Planar polarization in embryonic epidermis orchestrates global asymmetric morphogenesis of hair follicles. Nat Cell Biol 2008; 10:1257- 68; PMID:18849982; DOI:10.1038/ncb1784.
78. Gho M, Schweisguth F. Frizzled signalling controls orientation of asymmetric sense organ precursor cell divisions in Drosophila. Nature 1998; 393:178-81; PMID:9603522; DOI:10.1038/30265.
79. Mitchell B, Stubbs JL, Huisman F, Taborek P, Yu C, Kintner C. The PCP pathway instructs the planar orientation of ciliated cells in the Xenopus larval skin. Curr Biol 2009; 19:924-9; PMID:19427216; DOI:10.1016/j.cub.2009.04.018.
80. Guo N, Hawkins C, Nathans J. From The Cover: Frizzled6 controls hair patterning in mice. Proc Natl Acad Sci USA 2004; 101:9277-81; PMID:15169958; DOI:10.1073/pnas.0402802101.
81. Wallingford JB, Habas R. The developmental biology of Dishevelled: an enigmatic protein governing cell fate and cell polarity. Development 2005; 132:4421-36; PMID:16192308; DOI:10.1242/dev.02068.
82. Singh J, Yanfeng WA, Grumolato L, Aaronson SA, Mlodzik M. Abelson family kinases regulate Frizzled planar cell polarity signaling via Dsh phosphorylation. Genes Dev 2010; 24:2157-68; PMID:20837657; DOI:10.1101/gad. 1961010.
83. Axelrod JD. Unipolar membrane association of Dishevelled mediates Frizzled planar cell polarity signaling. Genes Dev 2001; 15:1182-7; PMID:11358862.
84. Wong HC, Bourdelas A, Krauss A, Lee HJ, Shao Y, Wu D, et al. Direct binding of the PDZ domain of Dishevelled to a conserved internal sequence in the C-terminal region of Frizzled. Mol Cell 2003; 12:1251-60; PMID:14636582; DOI:10.1016/S1097-2765(03)00427-1.
85. Wu J, Jenny A, Mirkovic I, Mlodzik M. Frizzled- Dishevelled signaling specificity outcome can be modulated by Diego in Drosophila. Mech Dev 2008; 125:30-42; PMID:18065209; DOI:10.1016/j. mod.2007.10.006.
86. Jenny A, Reynolds-Kenneally J, Das G, Burnett M, Mlodzik M. Diego and Prickle regulate Frizzled planar cell polarity signalling by competing for Dishevelled binding. Nat Cell Biol 2005; 7:691-7; PMID:15937478; DOI:10.1038/ncb1271.
87. Tree DR, Shulman JM, Rousset R, Scott MP, Gubb D, Axelrod JD. Prickle mediates feedback amplification to generate asymmetric planar cell polarity signaling. Cell 2002; 109:371-81; PMID:12015986; DOI:10.1016/ S0092-8674(02)00715-8.
88. Gubb D. Cellular polarity, mitotic synchrony and axes of symmetry during growth. Where does the information come from? Int J Dev Biol 1998; 42:369-77; PMID:9654021.
89. Jenny A, Darken RS, Wilson PA, Mlodzik M. Prickle and Strabismus form a functional complex to generate a correct axis during planar cell polarity signaling. EMBO J 2003; 22:4409-20; PMID:12941693; DOI:10.1093/emboj/cdg424.
90. Jenny A, Mlodzik M. Planar cell polarity signaling: a common mechanism for cellular polarization. Mt Sinai J Med 2006; 73:738-50; PMID:17008934.
91. Das G, Reynolds-Kenneally J, Mlodzik M. The Atypical Cadherin Flamingo Links Frizzled and Notch Signaling in Planar Polarity Establishment in the Drosophila Eye. Dev Cell 2002; 2:655-66; PMID:12015972; DOI:10.1016/S1534- 5807(02)00147-8.
92. Bastock R, Strutt H, Strutt D. Strabismus is asymmetrically localised and binds to Prickle and Dishevelled during Drosophila planar polarity patterning. Development 2003; 130:3007-14; PMID:12756182; DOI:10.1242/dev.00526.
93. Das G, Jenny A, Klein TJ, Eaton S, Mlodzik M. Diego interacts with Prickle and Strabismus/Van Gogh to localize planar cell polarity complexes. Development 2004; 131:4467-76; PMID:15306567; DOI:10.1242/ dev.01317.
94. Strutt DI. Asymmetric localization of frizzled and the establishment of cell polarity in the Drosophila wing. Mol Cell 2001; 7:367-75; PMID:11239465; DOI:10.1016/S1097-2765(01)00184-8.
95. Strutt H, Strutt D. Differential stability of flamingo protein complexes underlies the establishment of planar polarity. Curr Biol 2008; 18:1555-64; PMID:18804371; DOI:10.1016/j.cub.2008.08.063.
96. Wu J, Mlodzik M. The frizzled extracellular domain is a ligand for Van Gogh/Stbm during nonautonomous planar cell polarity signaling. Dev Cell 2008; 15:462-9; PMID:18804440; DOI:10.1016/j.devcel. 2008.08.004.
97. Strutt D, Johnson R, Cooper K, Bray S. Asymmetric localization of frizzled and the determination of notchdependent cell fate in the Drosophila eye. Curr Biol 2002; 12:813-24; PMID:12015117; DOI:10.1016/ S0960-9822(02)00841- 2.
98. Lu B, Usui T, Uemura T, Jan L, Jan YN. Flamingo controls the planar polarity of sensory bristles and asymmetric division of sensory organ precursors in Drosophila. Curr Biol 1999; 9:1247-50; PMID:10556092; DOI:10.1016/S0960- 9822(99)80505-3.
99. Deans MR, Antic D, Suyama K, Scott MP, Axelrod JD, Goodrich LV. Asymmetric distribution of pricklelike 2 reveals an early underlying polarization of vestibular sensory epithelia in the inner ear. J Neurosci 2007; 27:3139-47; PMID:17376975; DOI:10.1523/ JNEUROSCI.5151-06.2007.
100. Montcouquiol M, Sans N, Huss D, Kach J, Dickman JD, Forge A, et al. Asymmetric localization of Vangl2 and Fz3 indicate novel mechanisms for planar cell polarity in mammals. J Neurosci 2006; 26:5265-75; PMID:16687519; DOI:10.1523/ JNEUROSCI.4680-05.2006.
101. Yin C, Kiskowski M, Pouille PA, Farge E, Solnica- Krezel L. Cooperation of polarized cell intercalations drives convergence and extension of presomitic mesoderm during zebrafish gastrulation. J Cell Biol 2008; 180:221-32; PMID:18195109; DOI:10.1083/ jcb.200704150.
102. Goodrich LV, Strutt D. Principles of planar polarity in animal development. Development 2011; 138:1877- 92; PMID:21521735; DOI:10.1242/dev. 054080.
103. Vladar EK, Antic D, Axelrod JD. Planar Cell Polarity Signaling: The Developing Cell's Compass. Cold Spring Harb Perspect Biol 2009; 1:2964; PMID:20066108; DOI:10.1101/cshperspect.a002964.
104. Classen AK, Anderson KI, Marois E, Eaton S. Hexagonal packing of Drosophila wing epithelial cells by the planar cell polarity pathway. Dev Cell 2005; 9:805-17; PMID:16326392; DOI:10.1016/j.devcel. 2005.10.016.
105. Strutt H, Warrington SJ, Strutt D. Dynamics of core planar polarity protein turnover and stable assembly into discrete membrane subdomains. Dev Cell 2011; 20:511-25; PMID:21497763; DOI:10.1016/j.devcel. 2011.03.018.
106. Shimada Y, Yonemura S, Ohkura H, Strutt D, Uemura T. Polarized transport of Frizzled along the planar microtubule arrays in Drosophila wing epithelium. Dev Cell 2006; 10:209-22; PMID:16459300; DOI:10.1016/j.devcel.2005.11.016.
107. Harumoto T, Ito M, Shimada Y, Kobayashi TJ, Ueda HR, Lu B, et al. Atypical cadherins Dachsous and Fat control dynamics of noncentrosomal microtubules in planar cell polarity. Dev Cell 2010; 19:389-401; PMID:20817616; DOI:10.1016/j.devcel. 2010.08.004.
108. Narimatsu M, Bose R, Pye M, Zhang L, Miller B, Ching P, et al. Regulation of planar cell polarity by Smurf ubiquitin ligases. Cell 2009; 137:295-307; PMID:19379695; DOI:10.1016/j.cell.2009.02.025.
109. Adler PN, Krasnow RE, Liu J. Tissue polarity points from cells that have higher Frizzled levels towards cells that have lower Frizzled levels. Curr Biol 1997; 7:940-9; PMID:9382848; DOI:10.1016/S0960- 9822(06)00413-1.
110. Strutt D. Gradients and the specification of planar polarity in the insect cuticle. Cold Spring Harb Perspect Biol 2009; 1:489; PMID:20066114; DOI:10.1101/ cshperspect.a000489.
111. Strutt H, Strutt D. Long-range coordination of planar polarity in Drosophila. Bioessays 2005; 27:1218-27; PMID:16299762; DOI:10.1002/bies.20318.
112. Strutt D, Strutt H. Differential activities of the core planar polarity proteins during Drosophila wing patterning. Dev Biol 2007; 302:181-94; PMID:17045581; DOI:10.1016/j.ydbio.2006.09.026.
113. Boutros M, Mlodzik M. Dishevelled: at the crossroads of divergent intracellular signaling pathways. Mech Dev 1999; 83:27-37; PMID:10507837; DOI:10.1016/ S0925-4773(99)00046-5.
114. Fanto M, Weber U, Strutt DI, Mlodzik M. Nuclear signaling by Rac abd Rho GTPases is required in the establsihememt of epithelial planar polarity in the Drosophila eye. Curr Biol 2000; 10:979- 88; PMID:10985385; DOI:10.1016/S0960- 9822(00)00645-X.
115. Strutt DI, Weber U, Mlodzik M. The role of RhoA in tissue polarity and Frizzled signalling. Nature 1997; 387:292-5; PMID:9153394; DOI:10.1038/387292a0.
116. Winter CG, Wang B, Ballew A, Royou A, Karess R, Axelrod JD, et al. Drosophila Rho-associated kinase (Drok) links Frizzled-mediated planar cell polarity signaling to the actin cytoskeleton. Cell 2001; 105:81-91; PMID:11301004; DOI:10.1016/S0092- 8674(01)00298-7.
117. Weber U, Paricio N, Mlodzik M. Jun mediates Frizzled induced R3/R4 cell fate distinction and planar polarity determination in the Drosophila eye. Development 2000; 127:3619-29; PMID:10903185.
118. Weber U, Pataki C, Mihaly J, Mlodzik M. Combinatorial signaling by the Frizzled/PCP and Egfr pathways during planar cell polarity establishment in the Drosophila eye. Dev Biol 2008; 316:110-23; PMID:18291359; DOI:10.1016/j.ydbio. 2008.01.016.
119. Paricio N, Feiguin F, Boutros M, Eaton S, Mlodzik M. The Drosophila STE20-like kinase Misshapen is required downstream of the Frizzled receptor in planar polarity signaling. EMBO J 1999; 18:4669-78; PMID:10469646; DOI:10.1093/emboj/18.17.4669.
120. Cooper MTD, Bray SJ. Frizzled regulation of Notch signalling polarizes cell fate in the Drosophila eye. Nature 1999; 397:526-9; PMID:10028969; DOI:10.1038/17395.
121. Fanto M, Mlodzik M. Asymmetric Notch activation specifies photoreceptors R3 and R4 and planar polarity in the Drosophila eye. Nature 1999; 397:523-6; PMID:10028968; DOI:10.1038/17389.
122. Tomlinson A, Struhl G. Decoding vectorial information from a gradient: sequential roles of the receptors Frizzled and Notch in establishing planar polarity in the Drosophila eye. Development 1999; 126:5725-38; PMID:10572048.
123. Choi KW, Benzer S. Rotation of photoreceptor clusters in the developing Drosophila eye requires the nemo gene. Cell 1994; 78:125-36; PMID:8033204; DOI:10.1016/0092-8674(94)90579-7.
124. Mirkovic I, Gault WJ, Rahnama M, Jenny A, Gaengel K, Bessette D, et al. Nemo kinase phosphorylates betacatenin to promote ommatidial rotation and connects core PCP factors to E-cadherin-beta-catenin. Nat Struct Mol Biol 2011; 18:665-72; PMID:21552260; DOI:10.1038/nsmb.2049.
125. Brown KE, Freeman M. Egfr signalling defines a protective function for ommatidial orientation in the Drosophila eye. Development 2003; 130:5401-12; PMID:14507785; DOI:10.1242/dev.00773.
126. Gaengel K, Mlodzik M. Egfr signaling regulates ommatidial rotation and cell motility in the Drosophila eye via MAPK/Pnt signaling and the Ras effector Canoe/AF6. Development 2003; 130:5413-23; PMID:14507782; DOI:10.1242/dev.00759.
127. Strutt H, Strutt D. EGF signaling and ommatidial rotation in the Drosophila eye. Curr Biol 2003; 13:1451-7; PMID:12932331; DOI:10.1016/S0960- 9822(03)00545-1.
128. Adler PN. Planar signaling and morphogenesis in Drosophila. Dev Cell 2002; 2:525- 35; PMID:12015961; DOI:10.1016/S1534- 5807(02)00176-4.
129. Strutt D, Warrington SJ. Planar polarity genes in the Drosophila wing regulate the localisation of the FH3- domain protein Multiple Wing Hairs to control the site of hair production. Development 2008; 135:3103-11; PMID:18701542; DOI:10.1242/dev.025205.
130. Yan J, Huen D, Morely T, Johnson G, Gubb D, Roote J, et al. The multiple-wing-hairs gene encodes a novel GBD-FH3 domain-containing protein that functions both prior to and after wing hair initiation. Genetics 2008; 180:219-28; PMID:18723886; DOI:10.1534/ genetics.108.091314.
131. Higgs HN. Formin proteins: a domain-based approach. Trends Biochem Sci 2005; 30:342-53; PMID:15950879; DOI:10.1016/j.tibs.2005.04.014.
132. Adler PN, Zhu C, Stone D. Inturned localizes to the proximal side of wing cells under the instruction of upstream planar polarity proteins. Curr Biol 2004; 14:2046-51; PMID:15556868; DOI:10.1016/j. cub.2004.11.007.
133. Lu Q, Yan J, Adler PN. The Drosophila planar polarity proteins inturned and multiple wing hairs interact physically and function together. Genetics 2010; 185:549-58; PMID:20351219; DOI:10.1534/ genetics.110.114066.
134. Matusek T, Djiane A, Jankovics F, Brunner D, Mlodzik M, Mihaly J. The Drosophila formin DAAM regulates the tracheal cuticle pattern through organizing the actin cytoskeleton. Development 2006; 133:957-66; PMID:16469972; DOI:10.1242/dev.02266.
135. Yan J, Lu Q, Fang X, Adler PN. Rho1 has multiple functions in Drosophila wing planar polarity. Dev Biol 2009; 333:186-99; PMID:19576201; DOI:10.1016/j. ydbio.2009.06.027.
136. Simons M, Gault WJ, Gotthardt D, Rohatgi R, Klein TJ, Shao Y, et al. Electrochemical cues regulate assembly of the Frizzled/Dishevelled complex at the plasma membrane during planar epithelial polarization. Nat Cell Biol 2009; 11:286-94; PMID:19234454; DOI:10.1038/ncb1836.
137. Buechling T, Bartscherer K, Ohkawara B, Chaudhary V, Spirohn K, Niehrs C, et al. Wnt/Frizzled signaling requires dPRR, the Drosophila homolog of the prorenin receptor. Curr Biol 2010; 20:1263-8; PMID:20579883; DOI:10.1016/j.cub.2010.05.028.
138. +-ATPase- mediated acidification for Wnt signaling. Science 2010; 327:459- 63; PMID:20093472; DOI:10.1126/science.1179802.
139. Hermle T, Saltukoglu D, Grunewald J, Walz G, Simons M. Regulation of Frizzled-dependent planar polarity signaling by a V-ATPase subunit. Curr Biol 2010; 20:1269-76; PMID:20579879; DOI:10.1016/j. cub.2010.05.057.
140. Niehrs C, Boutros M. Trafficking, acidification and growth factor signaling. Sci Signal 2010; 3:26; PMID:20699474; DOI:10.1126/scisignal.3134pe26.
141. Adler PN, Charlton J, Liu J. Mutations in the cadherin superfamily member gene dachsous cause a tissue polarity phenotype by altering frizzled signaling. Development 1998; 125:959-68; PMID:9449678.
142. Ma D, Yang CH, McNeill H, Simon MA, Axelrod JD. Fidelity in planar cell polarity signalling. Nature 2003; 421:543-7; PMID:12540853; DOI:10.1038/ nature01366.
143. Ishikawa HO, Takeuchi H, Haltiwanger RS, Irvine KD. Four-jointed is a Golgi kinase that phosphorylates a subset of cadherin domains. Science 2008; 321:401-4; PMID:18635802; DOI:10.1126/science.1158159.
144. Strutt H, Mundy J, Hofstra K, Strutt D. Cleavage and secretion is not required for Four-jointed function in Drosophila patterning. Development 2004; 131:881-90; PMID:14757640; DOI:10.1242/dev.00996.
145. Zeidler MP, Perrimon N, Strutt DI. Multiple roles for four-jointed in planar polarity and limb patterning. Dev Biol 2000; 228:181-96; PMID:11112323; DOI:10.1006/dbio.2000.9940.
146. Brittle AL, Repiso A, Casal J, Lawrence PA, Strutt D. Four-jointed modulates growth and planar polarity by reducing the affinity of dachsous for fat. Curr Biol 2010; 20:803-10; PMID:20434337; DOI:10.1016/j. cub.2010.03.056.
147. Simon MA, Xu A, Ishikawa HO, Irvine KD. Modulation of Fat:Dachsous binding by the cadherin domain kinase four-jointed. Curr Biol 2010; 20:811-7; PMID:20434335; DOI:10.1016/j.cub.2010.04.016.
148. Matakatsu H, Blair SS. Separating the adhesive and signaling functions of the Fat and Dachsous protocadherins. Development 2006; 133:2315-24; PMID:16687445; DOI:10.1242/dev.02401.
149. Zeidler MP, Perrimon N, Strutt DI. Polarity determination in the Drosophila eye: a novel role for unpaired and JAK/STAT signaling. Genes Dev 1999; 13:1342-53; PMID:10346822; DOI:10.1101/gad.13.10.1342.
150. Fanto M, Clayton L, Meredith J, Hardiman K, Charroux B, Kerridge S, et al. The tumor-suppressor and cell adhesion molecule Fat controls planar polarity via physical interactions with Atrophin, a transcriptional co-repressor. Development 2003; 130:763-74; PMID:12506006; DOI:10.1242/dev.00304.
151. Matakatsu H, Blair SS. Interactions between Fat and Dachsous and the regulation of planar cell polarity in the Drosophila wing. Development 2004; 131:3785-94; PMID:15240556; DOI:10.1242/dev.01254.
152. Axelrod JD. Progress and challenges in understanding planar cell polarity signaling. Semin Cell Dev Biol 2009; 20:964-71; PMID:19665570; DOI:10.1016/j. semcdb.2009.08.001.
153. Simon MA. Planar cell polarity in the Drosophila eye is directed by graded Four-jointed and Dachsous expression. Development 2004; 131:6175-84; PMID:15548581; DOI:10.1242/dev.01550.
154. Sopko R, McNeill H. The skinny on Fat: an enormous cadherin that regulates cell adhesion, tissue growth and planar cell polarity. Curr Opin Cell Biol 2009; 21:717-23; PMID:19679459; DOI:10.1016/j.ceb.2009.07.001.
155. Wang WJ, Liu J, Adler PN. The frizzled gene of Drosophila encodes a membrane protein with an odd number of transmembrane domains. Mech Dev 1994; 45:127-37; PMID:8199049; DOI:10.1016/0925-4773(94)90026-4.
156. Adler PN, Lee H. Frizzled signaling and cell-cell interactions in planar polarity. Curr Opin Cell Biol 2001; 13:635-40; PMID:11544034; DOI:10.1016/S0955-0674(00)00263-5.
157. Lee H, Adler PN. The function of the frizzled pathway in the Drosophila wing is dependent on inturned and fuzzy. Genetics 2002; 160:1535-47; PMID:11973308.
158. Shulman JM, Perrimon N, Axelrod JD. Frizzled signaling and the developmental control of cell polarity. Trends Genet 1998; 14:452-8; PMID:9825673; DOI:10.1016/S0168-9525(98)01584-4.
159. Strutt D. Frizzled signalling and cell polarisation in Drosophila and vertebrates. Development 2003; 130:4501-13; PMID:12925579; DOI:10.1242/ dev.00695.
160. Adler PN, Vinson C, Park WJ, Conover S, Klein L. Molecular structure of frizzled, a Drosophila tissue polarity gene. Genetics 1990; 126:401-16; PMID:2174014.
161. Djiane A, Riou J, Umbhauer M, Boucaut J, Shi D. Role of frizzled 7 in the regulation of convergent extension movements during gastrulation in Xenopus laevis. Development 2000; 127:3091-100; PMID:10862746.
162. Jones SE, Jomary C, Grist J, Stewart HJ, Neal MJ. Modulated expression of secreted frizzledrelated proteins in human retinal degeneration. Neuroreport 2000; 11:3963-7; PMID:11192610; DOI:10.1097/00001756-200012180-00012.
163. Medina A, Reintsch W, Steinbeisser H. Xenopus frizzled 7 can act in canonical and non-canonical Wnt signaling pathways: implications on early patterning and morphogenesis. Mech Dev 2000; 92:227-37; PMID:10727861; DOI:10.1016/S0925-4773(00)00240-9.
164. Winklbauer R, Medina A, Swain RK, Steinbeisser H. Frizzled-7 signalling controls tissue separation during Xenopus gastrulation. Nature 2001; 413:856-60; PMID:11677610; DOI:10.1038/35101621.
165. Yu H, Smallwood PM, Wang Y, Vidaltamayo R, Reed R, Nathans J. Frizzled 1 and frizzled 2 genes function in palate, ventricular septum and neural tube closure: general implications for tissue fusion processes. Development 2010; 137:3707-17; PMID:20940229; DOI:10.1242/dev.052001.
166. Axelrod JD, Miller JR, Shulman JM, Moon RT, Perrimon N. Differential recruitment of Dishevelled provides signaling specificity in the planar cell polarity and Wingless signaling pathways. Genes Dev 1998; 12:2610-22; PMID:9716412; DOI:10.1101/ gad.12.16.2610.
167. Klingensmith J, Nusse R, Perrimon N. The Drosophila segment polarity gene dishevelled encodes a novel protein required for response to the wingless signal. Genes Dev 1994; 8:118-30; PMID:8288125; DOI:10.1101/ gad.8.1.118.
168. Schwarz-Romond T, Fiedler M, Shibata N, Butler PJ, Kikuchi A, Higuchi Y, et al. The DIX domain of Dishevelled confers Wnt signaling by dynamic polymerization. Nat Struct Mol Biol 2007; 14:484-92; PMID:17529994; DOI:10.1038/nsmb1247.
169. Theisen H, Purcell J, Bennett M, Kansagara D, Syed A, Marsh JL. dishevelled is required during wingless signalling to establish both cell polarity and cell identity. Development 1994; 120:347-60; PMID:8149913.
170. Tomlinson A, Strapps WR, Heemskerk J. Linking Frizzled and Wnt signaling in Drosophila development. Development 1997; 124:4515-21; PMID:9409669.
171. Wong HC, Mao J, Nguyen JT, Srinivas S, Zhang W, Liu B, et al. Structural basis of the recognition of the dishevelled DEP domain in the Wnt signaling pathway. Nat Struct Biol 2000; 7:1178-84; PMID:11101902; DOI:10.1038/82047.
172. Copp AJ, Greene ND, Murdoch JN. Dishevelled: linking convergent extension with neural tube closure. Trends Neurosci 2003; 26:453-5; PMID:12948650; DOI:10.1016/S0166-2236(03)00212-1.
173. Klingensmith J, Yang Y, Axelrod JD, Beier DR, Perrimon N, Sussman DJ. Conservation of dishevelled structure and function between flies and mice: isolation and characterization of Dvl2. Mech Dev 1996; 58:15-26; PMID:8887313; DOI:10.1016/S0925-4773(96)00549-7.
174. Sokol SY. Analysis of Dishevelled signalling pathways during Xenopus development. Curr Biol 1996; 6:1456-67; PMID:8939601; DOI:10.1016/S0960- 9822(96)00750-6.
175. Sokol SY, Klingensmith J, Perrimon N, Itoh K. Dorsalizing and neuralizing properties of Xdsh, a maternally expressed Xenopus homolog of dishevelled. Development 1995; 121:3487; PMID:7588081.
176. Wallingford JB, Rowning BA, Vogeli KM, Rothbacher U, Fraser SE, Harland RM. Dishevelled controls cell polarity during Xenopus gastrulation. Nature 2000; 405:81-5; PMID:10811222; DOI:10.1038/35011077.
177. Etheridge SLRS, Li S, Hamblet NS, Lijam N, Tsang M, Greer J, et al. Murine dishevelled 3 functions in redundant pathways with dishevelled 1 and 2 in normal cardiac outflow tract, cochlea and neural tube development. PLoS Genet 2008; 4:1000259; PMID:19008950; DOI:10.1371/journal.pgen.1000259.
178. Tada M, Smith JC. Xwnt11 is a target of Xenopus Brachyury: regulation of gastrulation movements via Dishevelled, but not through the canonical Wnt pathway. Development 2000; 127:2227-38; PMID:10769246.
179. Sussman DJKJ, Salinas P, Adams PS, Nusse R, Perrimon N. Isolation and characterization of a mouse homolog of the Drosophila segment polarity gene dishevelled. Dev Biol 1994; 166:73-86; PMID:7958461; DOI:10.1006/dbio.1994.1297.
180. Shimada Y, Usui T, Yanagawa S, Takeichi M, Uemura T. Asymmetric colocalization of Flamingo, a seven- pass transmembrane cadherin and Dishevelled in planar cell polarization. Curr Biol 2001; 11:859-63; PMID:11516647; DOI:10.1016/S0960- 9822(01)00233-0.
181. Curtin JA, Quint E, Tsipouri V, Arkell RM, Cattanach B, Copp AJ, et al. Mutation of Celsr1 disrupts planar polarity of inner ear hair cells and causes severe neural tube defects in the mouse. Curr Biol 2003; 13:1129-33; PMID:12842012; DOI:10.1016/S0960- 9822(03)00374-9.
182. Formstone CJ, Little PF. The flamingo-related mouse Celsr family (Celsr1-3) genes exhibit distinct patterns of expression during embryonic development. Mech Dev 2001; 109:91-4; PMID:11677057; DOI:10.1016/ S0925-4773(01)00515-9.
183. Davies A, Formstone C, Mason I, Lewis J. Planar polarity of haircells in the chick inner ear is correlated with polarized distribution of c-flamingo-1 protein. Dev Dyn 2005; 233:998-1005; PMID:15830377; DOI:10.1002/dvdy.20376.
184. Carreira-Barbosa F, Kajita M, Morel V, Wada H, Okamoto H, Martinez, et al. Flamingo regulates epiboly and convergence/extension movements through cell cohesive and signalling functions during zebrafish gastrulation. Development 2009; 136:383-92; PMID:19091770; DOI:10.1242/dev.026542.
185. Dabdoub A, Montcouquiol M, Kelley MW. Planar cell polarity in the vertebrate inner ear. In: Mlodzik M, Ed. Planar cell polarization during development. San Diego: Elsevier 2005; 107-30. (Pubitemid 44725681)
186. Moeller H, Jenny A, Schaeffer HJ, Schwarz-Romond T, Mlodzik M, Hammerschmidt M, et al. Diversin regulates heart formation and gastrulation movements in development. Proc Natl Acad Sci USA 2006; 103:15900-5; PMID:17032765; DOI:10.1073/ pnas.0603808103.
187. Schwarz-Romond T, Asbrand C, Bakkers J, Kuhl M, Schaeffer HJ, Huelsken J, et al. The ankyrin repeat protein Diversin recruits Casein kinase Iepsilon to the beta-catenin degradation complex and acts in both canonical Wnt and Wnt/JNK signaling. Genes Dev 2002; 16:2073-84; PMID:12183362; DOI:10.1101/ gad.230402.
188. Watanabe D, Saijoh Y, Nonaka S, Sasaki G, Ikawa Y, Yokoyama T, et al. The left-right determinant Inversin is a component of node monocilia and other 9 + 0 cilia. Development 2003; 130:1725-34; PMID:12642479; DOI:10.1242/dev.00407.
189. Morgan D, Turnpenny L, Goodship J, Dai W, Majumder K, Matthews L, et al. Inversin, a novel gene in the vertebrate left-right axis pathway, is partially deleted in the inv mouse. Nat Genet 1998; 20:149-56; PMID:9771707; DOI:10.1038/2450.
190. Otto EA, Schermer B, Obara T, O'Toole JF, Hiller KS, Mueller AM, et al. Mutations in INVS encoding inversin cause nephronophthisis type 2, linking renal cystic disease to the function of primary cilia and leftright axis determination. Nat Genet 2003; 34:413-20; PMID:12872123; DOI:10.1038/ng1217.
191. Simons M, Gloy J, Ganner A, Bullerkotte A, Bashkurov M, Kronig C, et al. Inversin, the gene product mutated in nephronophthisis type II, functions as a molecular switch between Wnt signaling pathways. Nat Genet 2005; 37:537-43; PMID:15852005; DOI:10.1038/ ng1552.
192. Lee OK, Frese KK, James JS, Chadda D, Chen ZH, Javier RT, et al. Discs-Large and Strabismus are functionally linked to plasma membrane formation. Nat Cell Biol 2003; 5:987-93; PMID:14562058; DOI:10.1038/ncb1055.
193. Rawls AS, Wolff T. Strabismus requires Flamingo and Prickle function to regulate tissue polarity in the Drosophila eye. Development 2003; 130:1877-87; PMID:12642492; DOI:10.1242/dev.00411.
194. Darken RS, Scola AM, Rakeman AS, Das G, Mlodzik M, Wilson PA. The planar polarity gene strabismus regulates convergent extension movements in Xenopus. EMBO J 2002; 21; In press; PMID:11867525; DOI:10.1093/emboj/21.5.976.
195. Gong Y, Mo C, Fraser SE. Planar cell polarity signalling controls cell division orientation during zebrafish gastrulation. Nature 2004; 430:689-93; PMID:15254551; DOI:10.1038/nature02796.
196. Goto T, Keller R. The planar cell polarity gene strabismus regulates convergence and extension and neural fold closure in Xenopus. Dev Biol 2002; 247:165-81; PMID:12074560; DOI:10.1006/dbio.2002.0673.
197. Henderson DJ, Phillips HM, Chaudhry B. Vang-like 2 and non-canonical Wnt signaling in outflow tract development. Trends Cardiovasc Med 2006; 16:38-45; PMID:16473760; DOI:10.1016/j.tcm.2005.11.005.
198. Jessen JR, Topczewski J, Bingham S, Sepich DS, Marlow F, Chandrasekhar A, et al. Zebrafish trilobite indentifies new roles for Strabismus in gastrulation and neuronal movements. Nat Cell Biol 2002; 4:610-5; PMID:12105418.
199. Kibar Z, Vogan K, Groulx N, Justice MJ, Underhill DA, Gros P. Ltap, a mammalian homolog of Drosophila Strabismus/Van Gogh, is altered in the mouse neural tube mutant Loop-tail. Nat Genet 2001; 28:251-2; PMID:11431695; DOI:10.1038/90081.
200. Montcouquiol M, Rachel RA, Lanford PJ, Copeland NG, Jenkins NA, Kelley MW. Identification of Vangl2 and Scrb1 as planar polarity genes in mammals. Nature 2003; 423:173-7; PMID:12724779; DOI:10.1038/ nature01618.
201. Borovina A, Superina S, Voskas D, Ciruna B. Vangl2 directs the posterior tilting and asymmetric localization of motile primary cilia. Nat Cell Biol 2010; 12:407-12; PMID:20305649; DOI:10.1038/ncb2042.
202. Kibar Z, Bosoi CM, Kooistra M, Salem S, Finnell RH, De Marco P, et al. Novel mutations in VANGL1 in neural tube defects. Hum Mutat 2009; 30:706-15; PMID:19319979; DOI:10.1002/humu.21026.
203. Kibar Z, Torban E, McDearmid JR, Reynolds A, Berghout J, Mathieu M, et al. Mutations in VANGL1 associated with neural-tube defects. N Engl J Med 2007; 356:1432-7; PMID:17409324; DOI:10.1056/ NEJMoa060651.
204. Murdoch JN, Doudney K, Paternotte C, Copp AJ, Stanier P. Severe neural tube defects in the loop-tail mouse result from mutation of Lpp1, a novel gene involved in floor plate specification. Hum Mol Genet 2001; 10:2593-601; PMID:11709546; DOI:10.1093/ hmg/10.22.2593.
205. Phillips HM, Hildreth V, Peat JD, Murdoch JN, Kobayashi K, Chaudhry B, et al. Non-cell-autonomous roles for the planar cell polarity gene Vangl2 in development of the coronary circulation. Circ Res 2008; 102:615-23; PMID:18174466; DOI:10.1161/ CIRCRESAHA.107.160861.
206. Phillips HM, Murdoch JN, Chaudhry B, Copp AJ, Henderson DJ. Vangl2 acts via RhoA signaling to regulate polarized cell movements during development of the proximal outflow tract. Circ Res 2005; 96:292-9; PMID:15637299; DOI:10.1161/01.RES.0000154912.08695.88
207. Phillips HM, Rhee HJ, Murdoch JN, Hildreth V, Peat JD, Anderson RH, et al. Disruption of planar cell polarity signaling results in congenital heart defects and cardiomyopathy attributable to early cardiomyocyte disorganization. Circ Res 2007; 101:137-45; PMID:17556662; DOI:10.1161/ CIRCRESAHA.106.142406.
208. Reynolds A, McDearmid JR, Lachance S, Marco PD, Merello E, Capra V, et al. VANGL1 rare variants associated with neural tube defects affect convergent extension in zebrafish. Mech Dev 2010; 127:385-92; PMID:20043994; DOI:10.1016/j.mod.2009.12.002.
209. Ybot-Gonzalez P, Savery D, Gerrelli D, Signore M, Mitchell CE, Faux CH, et al. Convergent extension, planar-cell-polarity signalling and initiation of mouse neural tube closure. Development 2007; 134:789-99; PMID:17229766; DOI:10.1242/dev.000380.
210. Marlow F, Zwartkruis F, Malicki J, Neuhauss SCF, Abbas L, Weaver M, et al. Functional interactions of genes mediating convergent extension, knypek and trilobite, during the partitioning of the eye primordium in zebrafish. Dev Biol 1998; 203:382-93; PMID:9808788; DOI:10.1006/dbio.1998.9032.
211. Zhang YLM. Left-right asymmetry in the chick embryo requires core planar cell polarity protein Vangl2. Genesis 2009; 47:719-28; PMID:19621439; DOI:10.1002/dvg.20551.
212. Song H, Hu J, Chen W, Elliott G, Andre P, Gao B, et al. Planar cell polarity breaks bilateral symmetry by controlling ciliary positioning. Nature 2010; 466:378- 82; PMID:20562861; DOI:10.1038/nature09129.
213. Mlodzik M. Spiny legs and prickled bodies: new insights and complexities in planar polarity establishment. Bioessays 2000; 22:311-5; PMID:10723028; DOI:10.1002/(SICI)1521- 1878(200004)22:4〈311::AID-BIES1〉3.0.CO;2-J.
214. Carreira-Barbosa F, Concha ML, Takeuchi M, Ueno N, Wilson SW, Tada M. Prickle 1 regulates cell movements during gastrulation and neuronal migration in zebrafish. Development 2003; 130:4037-46; PMID:12874125; DOI:10.1242/dev.00567.
215. Veeman MT, Slusarski DC, Kaykas A, Louie SH, Moon RT. Zebrafish prickle, a modulator of noncanonical wnt/fz signaling, regulates gastrulation movements. Curr Biol 2003; 13:680-5; PMID:12699626; DOI:10.1016/S0960-9822(03)00240-9.
216. Wallingford JB, Goto T, Keller R, Harland RM. Cloning and expression of Xenopus Prickle, an orthologue of a Drosophila planar cell polarity gene. Mech Dev 2002; 116:183-6; PMID:12128221; DOI:10.1016/S0925-4773(02)00133-8.
217. Takeuchi M, Nakabayashi J, Sakaguchi T, Yamamoto TS, Takahashi H, Takeda H, et al. The prickle-Related Gene in Vertebrates Is Essential for Gastrulation Cell Movements. Curr Biol 2003; 13:674-9; PMID:12699625; DOI:10.1016/S0960- 9822(03)00245-8.
218. Lawrence PA, Struhl G, Casal J. Do the protocadherins Fat and Dachsous link up to determine both planar cell polarity and the dimensions of organs? Nat Cell Biol 2008; 10:1379-82; PMID:19043429; DOI:10.1038/ ncb1208-379.
219. Halbleib JM, Nelson WJ. Cadherins in development: cell adhesion, sorting and tissue morphogenesis. Genes Dev 2006; 20:3199-214; PMID:17158740; DOI:10.1101/gad.1486806.
220. Sopko R, Silva E, Clayton L, Gardano L, Barrios- Rodiles M, Wrana J, et al. Phosphorylation of the tumor suppressor fat is regulated by its ligand Dachsous and the kinase discs overgrown. Curr Biol 2009; 19:1112-7; PMID:19540118; DOI:10.1016/j.cub.2009.05.049.
221. Saburi S, Hester I, Fischer E, Pontoglio M, Eremina V, Gessler M, et al. Loss of Fat4 disrupts PCP signaling and oriented cell division and leads to cystic kidney disease. Nat Genet 2008; 40:1010-5; PMID:18604206; DOI:10.1038/ng.179.
222. Speck O, Hughes SC, Noren NK, Kulikauskas RM, Fehon RG. Moesin functions antagonistically to the Rho pathway to maintain epithelial integrity. Nature 2003; 421:83-7; PMID:12511959; DOI:10.1038/ nature01295.
223. Eaton S, Auvinen P, Luo L, Jan YN, Simons K. CDC42 and Rac1 control different actin-dependent processes in the Drosophila wing disc epithelium. J Cell Biol 1995; 131:151-64; PMID:7559772; DOI:10.1083/ jcb.131.1.151.
224. Habas R, Dawid IB, He X. Coactivation of Rac and Rho by Wnt/Frizzled signaling is required for vertebrate gastrulation. Genes Dev 2003; 17:295-309; PMID:12533515; DOI:10.1101/gad.1022203.
225. Tahinci E, Symes K. Distinct functions of Rho and Rac are required for convergent extension during Xenopus gastrulation. Dev Biol 2003; 259:318- 35; PMID:12871704; DOI:10.1016/S0012-1606(03)00206-9.
226. Marlow F, Topczewski J, Sepich D, Solnica-Krezel L. Zebrafish rho kinase 2 acts downstream of wnt11 to mediate cell polarity and effective convergence and extension movements. Curr Biol 2002; 12:876-84; PMID:12062050; DOI:10.1016/S0960- 9822(02)00864-3.
227. Habas R, Kato Y, He X. Wnt/Frizzled activation of rho regulates vertebrate gastrulation and requires a novel formin homology protein Daam1. Cell 2001; 107:843-54; PMID:11779461; DOI:10.1016/S0092- 8674(01)00614-6.
228. Liu W, Sato A, Khadka D, Bharti R, Diaz H, Runnels LW, et al. Mechanism of activation of the Formin protein Daam1. Proc Natl Acad Sci USA 2008; 105:210-5; PMID:18162551; DOI:10.1073/pnas.0707277105.
229. Su Y, Treisman JE, Skolnik EY. The Drosophila Ste20- related kinase misshapen is required for embryonic dorsal closure and acts through a JNK MAPK module on an evolutionary conserved signaling pathway. Genes Dev 1998; 12:2371-80; PMID:9694801; DOI:10.1101/gad.12.15.2371.
230. Treisman JE, Ito N, Rubin GM. misshapen encodes a protein kinase involved in cell shape control in Drosophila. Gene 1997; 186:119-25; PMID:9047354; DOI:10.1016/S0378-1119(96)00694-4.
231. Yamanaka H, Moriguchi T, Masuyama N, Kusakabe M, Hanafusa H, Takada R, et al. JNK functions in the non-canonical Wnt pathway to regulate convergent extension movements in vertebrates. EMBO Rep 2002; 3:69-75; PMID:11751577; DOI:10.1093/ embo-reports/kvf008.
232. Couso JP, Martinez Arias A. Notch is required for Wingless signaling in Drosophila. Cell 1994; 79:259-72; PMID:7954794; DOI:10.1016/0092- 8674(94)90195-3.
233. Diaz-Benjumea FJ, Cohen SM. Serrate signals through Notch to establish a Wingless-dependent organizer at the dorsal/ventral compartment boundary of the Drosophila wing. Development 1995; 121:4215-25; PMID:8575321.
234. Domingos PM, Mlodzik M, Mendes CS, Brown S, Steller H, Mollereau B. Spalt transcription factors are required for R3/R4 specification and establishment of planar cell polarity in the Drosophila eye. Development 2004; 131:5695-702; PMID:15509769; DOI:10.1242/dev.01443.
235. Dominguez M, de Celis JF. A dorsal/ventral boundary established by Notch controls growth and polarity in the Drosophila eye. Nature 1998; 396:276-8; PMID:9834035; DOI:10.1038/24402.
236. Fiehler RW, Wolff T. Nemo is required in a subset of photoreceptors to regulate the speed of ommatidial rotation. Dev Biol 2008; 313:533-44; PMID:18068152; DOI:10.1016/j.ydbio.2007.10.034.
237. Freeman M. Misexpression of the Drosophila argos gene, a secreted regulator of cell determination. Development 1994; 120:2297-304; PMID:7925030.
238. Freeman M, Klämbt C, Goodman CS, Rubin GM. The argos gene encodes a diffusible factor that regulates cell fate decisions in the Drosophila eye. Cell 1992; 69:963-75; PMID:1606617; DOI:10.1016/0092- 8674(92)90615-J.
239. Sawamoto K, Okabe M, Tanimura T, Mikoshiba K, Nishida Y, Okano H. The Drosophila secreted protein Argos regulates signal transduction in the Ras/MAPK pathway. Dev Biol 1996; 178:13-22; PMID:8812105; DOI:10.1006/dbio.1996.0194.
240. Sawamoto K, Okano H, Kobayakawa Y, Hayashi S, Mikoshiba K, Tanimura T. The function of argos in regulating cell fate decisions during Drosophila eye and wing vein development. Dev Biol 1994; 164:267-76; PMID:8026629; DOI:10.1006/dbio.1994.1197.
241. Bellaiche Y, Radovic A, Woods DF, Hough CD, Parmentier ML, O'Kane CJ, et al. The Partner of Inscuteable/Discs-large complex is required to establish planar polarity during asymmetric cell division in Drosophila. Cell 2001; 106:355-66; PMID:11509184; DOI:10.1016/S0092-8674(01)00444-5.
242. Collier S, Gubb D. Drosophila tissue polarity requires the cell-autonomous activity of the fuzzy gene, which encodes a novel transmembrane protein. Development 1997; 124:4029-37; PMID:9374400.
243. Park TJHS, Wallingford JB. Ciliogenesis defects in embryos lacking inturned or fuzzy function are associated with failure of planar cell polarity and Hedgehog signaling. Nat Genet 2006; 38:303-11; PMID:16493421; DOI:10.1038/ng1753.
244. Gray RS, Abitua PB, Wlodarczyk BJ, Szabo-Rogers HL, Blanchard OLI, Weiss GS, et al. The planar cell polarity effector Fuz is essential for targeted membrane trafficking, ciliogenesis and mouse embryonic development. Nat Cell Biol 2009; 11:1225-32; PMID:19767740; DOI:10.1038/ncb1966.
245. Heydeck W, Zeng H, Liu A. Planar cell polarity effector gene Fuzzy regulates cilia formation and Hedgehog signal transduction in mouse. Dev Dyn 2009; 238:3035- 42; PMID:19877275; DOI:10.1002/dvdy.22130.
246. Adler PN, Charlton J, Park WJ. The Drosophila tissue polarity gene inturned functions prior to wing hair morphogenesis in the regulation of hair polarity and number. Genetics 1994; 137:1-8; PMID:8056302.
247. Yun UJ, Kim SY, Liu J, Adler PN, Bae E, Kim J, et al. The inturned protein of Drosophila melanogaster is a cytoplasmic protein located at the cell periphery in wing cells. Dev Genet 1999; 25:297- 305; PMID:10570461; DOI:10.1002/(SICI)1520- 6408(1999)25:4〈297::AID-DVG3〉3.0.CO;2-L.
248. Zeng H, Hoover AN, Liu A. PCP effector gene Inturned is an important regulator of cilia formation and embryonic development in mammals. Dev Biol 2010; 339:418-28; PMID:20067783; DOI:10.1016/j. ydbio.2010.01.003.
249. Collier SLH, Burgess R, Adler P. The WD40 repeat protein fritz links cytoskeletal planar polarity to frizzled subcellular localization in the Drosophila epidermis. Genetics 2005; 169:2035-45; PMID:15654087; DOI:10.1534/genetics.104.033381.
250. Kim SKSA, Park TJ, Oh EC, Ghosh S, Gray RS, Lewis RA, et al. Planar cell polarity acts through septins to control collective cell movement and ciliogenesis. Science 2010; 329:1337-40; PMID:20671153; DOI:10.1126/science. 1191184.