Steroid signaling in plants and insects - Common themes, different pathways

Genes and Development

Volumn 16, Issue 24, 2002, Pages 3113-3129

  Thummel, Carl S ^a   Chory, Joanne ^b  

^a UNIVERSITY OF UTAH SCHOOL OF MEDICINE   (United States)

^b HOWARD HUGHES MEDICAL INSTITUTE   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

BRASSINOSTEROID; ECDYSTEROID; PESTICIDE; STEROID;

ARABIDOPSIS; DROSOPHILA MELANOGASTER; GENE EXPRESSION REGULATION; HOMEOSTASIS; INSECT; NONHUMAN; PLANT; PRIORITY JOURNAL; REVIEW; SIGNAL TRANSDUCTION; STEROIDOGENESIS;

ANIMALS; GENE EXPRESSION REGULATION; INSECTS; PLANT PHYSIOLOGY; SIGNAL TRANSDUCTION; STEROIDS;

ARABIDOPSIS; DROSOPHILA MELANOGASTER; HEXAPODA; INSECTA; MELANOGASTER;

EID: 0037115466     PISSN: 08909369     EISSN: None     Source Type: Journal    
DOI: 10.1101/gad.1042102     Document Type: Review

References (166)

1. Andres, A.J. and Cherbas, P. 1992. Tissue-specific ecdysone reponses: Regulation of the Drosophila genes Eip28/29 and Eip40 during larval development. Development 116: 865-876.
2. Andres, A.J., Fletcher, J.C., Karim, F.D., and Thummel, C.S. 1993. Molecular analysis of the initiation of insect metamorphosis: A comparative study of Drosophila ecdysteroid-regulated transcription. Dev. Biol. 160: 388-404.
3. Arbeitman, M.N. and Hogness, D.S. 2000. Molecular chaperones activate the Drosophila ecdysone receptor, an RXR heterodimer. Cell 101: 67-77.
4. Ashburner, M. 1971. Induction of puffs in polytene chromosomes of in vitro cultured salivary glands of Drosophila melanogaster by ecdysone and ecdysone analogues. Nat. New Biol. 230: 222-224.
5. Ashburner, M., Chihara, C., Meltzer, P., and Richards, G. 1974. Temporal control of puffing activity in polytene chromosomes. Cold Spring Harb. Symp. Quant. Biol. 38: 655-662.
6. Ayer, S., Walker, N., Mosammaparast, M., Nelson, J.P., Shilo, B.Z., and Benyajati, C. 1993. Activation and repression of Drosophila alcohol dehydrogenase distal transcription by two steroid hormone receptor superfamily members binding to a common response element. Nucleic Acids Res. 21: 1619-1627.
7. Baehrecke, E.H. 2002. How death shapes life during development. Nat. Rev. Mol. Cell Biol. 3: 779-787.
8. Baehrecke, E.H. and Thummel, C.S. 1995. The Drosophila E93 gene from the 93F early puff displays stage- and tissue-specific regulation by 20-hydroxyecdysone. Dev. Biol. 171: 85-97.
9. Baker, K.D., Warren, J.T., Thummel, C.S., Gilbert, L.I., and Mangelsdorf, D.J. 2000. Transcriptional activation of the Drosophila ecdysone receptor by insect and plant ecdysteroids. Insect Biochem. Mol. Biol. 30: 1037-1043.
10. Bayer, C.A., Holley, B., and Fristrom, J.W. 1996. A switch in Broad-complex zinc-finger isoform expression is regulated posttranscriptionally during the metamorphosis of Drosophila imaginal discs. Dev. Biol. 177: 1-14.
11. Bayer, C.A., von Kalm, L., and Fristrom, J.W. 1997. Relationships between protein isoforms and genetic functions demonstrate functional redundancy at the Broad-Complex during Drosophila metamorphosis. Dev. Biol. 187: 267-282.
12. Becker, H.J. 1959. Die puffs der speicheldrüsenchromosomen von Drosophila melanogaster. I. Beobachtungen zur verhalten des puffmusters in normalstamm und bei zwei mutanten, giant und lethal-giant-larvae. Chromosoma 10: 654-678.
13. Bender, M. 2003. Molecular mechanism of ecdysone action in insect development. In Encyclopedia of Hormones (eds. H. Henry and A. Norman), Academic Press, New York (In press).
14. Bender, M., Imam, F.B., Talbot, W.S., Ganetzky, B., and Hogness, D.S. 1997. Drosophila ecdysone receptor mutations reveal functional differences among receptor isoforms. Cell 91: 777-788.
15. Bergamasco, R. and Horn, D. 1980. The biological activities of ecdysteroids and ecdysteroid analogues. In Progress in Ecdysone Research (ed. J. Hoffman), pp. 299-324. Elsevier, New York.
16. Bialecki, M., Shilton, A., Fichtenberg, C., Segraves, W., and Thummel, C. 2002. Loss of the ecdysteroid-inducible E75A orphan nuclear receptor uncouples molting from metamorphosis in Drosophila. Dev. Cell 3: 209-220.
17. Billas, I.M., Moulinier, L., Rochel, N., and Moras, D. 2001. Crystal structure of the ligand-binding domain of the Ultraspiracle protein USP, the ortholog of retinoid X receptors in insects. J. Biol. Chem. 276: 7465-7474.
18. Bishop, G.J. and Koncz, C. 2002. Brassinosteroids and plant steroid hormone signaling. Plant Cell 14: 97-110.
19. Bishop, G.J. and Yokota, T. 2001. Plants steroid hormones, brassinosteroids: Current highlights of molecular aspects on their synthesis/metabolism, transport, perception and response. Plant Cell Physiol. 42: 114-120.
20. Bishop, G.J., Harrison, K., and Jones, J.D. 1996. The tomato Dwarf gene isolated by heterologous transposon tagging encodes the first member of a new cytochrome P450 family. Plant Cell 8: 959-969.
21. Bishop, G.J., Nomura, T., Yokota, T., Harrison, K., Noguchi, T., Fujioka, S., Takatsuto, S., Jones, J.D., and Kamiya, Y. 1999. The tomato DWARF enzyme catalyses C-6 oxidation in brassinosteroid biosynthesis. Proc. Natl Acad. Sci. 96: 1761-1766.
22. Blais, C., Dauphin-Villemant, C., Kovganko, N., Girault, J.P., Descoins Jr., C., and Lafont, R. 1996. Evidence for the involvement of 3-xo-δ 4 intermediates in ecdysteroid biosynthesis. Biochem. J. 320: 413-419.
23. Burtis, K.C., Thummel, C.S., Jones, C.W., Karim, F.D., and Hogness, D.S. 1990. The Drosophila 74EF early puff contains E74, a complex ecdysone-inducible gene that encodes two ets-related proteins. Cell 61: 85-99.
24. Cabrera y Poch, H.L., Peto, C.A., and Chory, J. 1993. A mutation in the Arabidopsis DET3 gene uncouples photoregulated leaf development from gene expression and chloroplast biogenesis. Plant J. 4: 671-682.
25. Cadigan, K.M. and Nusse, R. 1997. Wnt signaling: A common theme in animal development. Genes & Dev. 11: 3286-3305.
26. Champlin, D.T. and Truman, J.W. 1998. Ecdysteroid control of cell proliferation during optic lobe neurogenesis in the moth Manduca sexta. Development 125: 269-277.
27. -. 2000. Ecdysteroid coordinates optic lobe neurogenesis via a nitric oxide signaling pathway. Development 127: 3543-3551.
28. Chavez, V.M., Marques, G., Delbecque, J.P., Kobayashi, K., Hollingsworth, M., Burr, J., Natzle, J.E., and O'Connor, M.B. 2000. The Drosophila disembodied gene controls late embryonic morphogenesis and codes for a cytochrome P450 enzyme that regulates embryonic ecdysone levels. Development 127: 4115-4126.
29. Cherbas, L., Yonge, C., Cherbas, P., and Williams, C. 1980. The morphological response of Kc-H cells to ecdysteroids: Hormonal specificity. Willhelm Roux's Arch. Dev. Biol. 189: 1-15.
30. Choe, S., Dilkes, B.P., Fujioka, S., Takatsuto, S., Sakurai, A., and Feldmann, K.A. 1998a. The DWF4 gene of Arabidopsis encodes a cytochrome P450 that mediates multiple 22α-hydroxylation steps in brassinosteroid biosynthesis. Plant Cell 10: 231-243.
31. Choe, S., Fujioka, S., Tanaka, A., Tissier, C., Rossa, A., Noguchi, T., Takatsuto, S., Yoshidab, S., Taxa, F., and Feldmann, K. 1998b. Molecular cloning of Arabidopsis brassinosteroid biosynthetic genes DWF4, DWF5 and DWF7. In Abstracts of 9th International Conference on Arabidopsis Research pp. 555, University of Wisconsin-Madison, Madison, WI.
32. Choe, S., Dilkes, B.P., Gregory, B.D., Ross, A.S., Yuan, H., Noguchi, T., Fujioka, S., Takatsuto, S., Tanaka, A., Yoshida, S., et al. 1999a. The Arabidopsis dwarf1 mutant is defective in the conversion of 24-methylenecholesterol to campesterol in brassinosteroid biosynthesis. Plant Physiol. 119: 897-907.
33. Choe, S., Noguchi, T., Fujioka, S., Takatsuto, S., Tissier, C.P., Gregory, B.D., Ross, A.S., Tanaka, A., Yoshida, S., Tax, F.E., et al. 1999b. The Arabidopsis dwf7/ste1 mutant is defective in the δ7 sterol C-5 desaturation step leading to brassinosteroid biosynthesis. Plant Cell 11: 207-221.
34. Choe, S., Tanaka, A., Noguchi, T., Fujioka, S., Takatsuto, S., Ross, A.S., Tax, F.E., Yoshida, S., and Feldmann, K.A. 2000. Lesions in the sterol δ reductase gene of Arabidopsis cause dwarfism due to a block in brassinosteroid biosynthesis. Plant J. 21: 431-443.
35. Choe, S., Fujioka, S., Noguchi, T., Takatsuto, S., Yoshida, S., and Feldmann, K.A. 2001. Overexpression of DWARF4 in the brassinosteroid biosynthetic pathway results in increased vegetative growth and seed yield in Arabidopsis. Plant J. 26: 573-582.
36. Choi, Y.-H., Fujioka, S., Harada, A., Yokota, T., Takatsuto, S., and Sakurai, A. 1996. A brassinolide biosynthetic pathway via 6-deoxocastasterone. Phytochemistry 43: 593-596.
37. Choi, Y.-H., Fujioka, S., Nomura, T., Harada, A., Yokota, T., Takatsuto, S., and Sakurai, A. 1997. An alternative brassinolide biosynthetic pathway via late C6-oxidation. Phytochemistry 44: 609-613.
38. Chory, J., Nagpal, P., and Peto, C.A. 1991. Phenotypic and genetic analysis of det2, a new mutant that affects light-regulated seedling development in Arabidopsis. Plant Cell 3: 445-459.
39. Chory, J., Reinecke, D., Sim, S., Washburn, T., and Brenner, M. 1994. A role for cytokinins in de-etiolation in Arabodipsis: Det mutants may have an altered response to cytokinins. Plant Physiol. 104: 339-347.
40. Clayton, G.M., Peak-Chew, S.Y., Evans, R.M., and Schwabe, J.W. 2001. The structure of the ultraspiracle ligand-binding domain reveals a nuclear receptor locked in an inactive conformation. Proc. Natl. Acad. Sci. 98: 1549-1554.
41. Clever, U. and Karlson, P. 1960. Induktion von puff-veränderungen in den speicheldrüsenchromosomen von chironomus tentans durch ecdyson. Exp. Cell. Res. 20: 623-626.
42. Clouse, S.D. 1996. Molecular genetic studies confirm the role of brassinosteroids in plant growth and development. Plant J. 10: 1-8.
43. Clouse, S. and Sasse, J. 1998. Brassinosteroids: Essential regulators of plant growth and development. Annu. Rev. Plant Physiol. Plant Mol. Biol. 49: 427-451.
44. Clouse, S. and Zurek, D. 1991. Molecular analysis of brassinolide action in plant growth and development. In Brassinosteroids: Chemistry, bioactivity and Applications (eds. H. Cutler, T. Yokota, and G. Adams), pp. 122-140. American Chemical Society, Washington, DC.
45. Clouse, S.D., Langford, M., and McMorris, T.C. 1996. A brassinosteroid-insensitive mutant in Arabidopsis thaliana exhibits multiple defects in growth and development. Plant Physiol. 111: 671-678.
46. Cutler, H.G., Yokota, T., and Adam, G., eds. 1991. Brassinosteroids. ACS Symposium Series pp. 474-496. American Chemical Society, Washington, DC.
47. D'Avino, P.P. and Thummel, C.S. 1998. crooked legs encodes a family of zinc finger proteins required for leg morphogenesis and ecdysone-regulated gene expression during Drosophila metamorphosis. Development 125: 1733-1745.
48. Descoins Jr., C. and Marion-Poll, F. 1999. Electrophysiological responses of gustatory sensilla of Mamestra brassicae larvae to three ecdysteroids: Ecdysone, 20-hydroxyecdysone and ponasterone A. J. Insect Phys. 45: 871-876.
49. DiBello, P.R., Withers, D.A., Bayer, C.A., Fristrom, J.W., and Guild, G.M. 1991. The Drosophila Broad-Complex encodes a family of related proteins containing zinc fingers. Genetics 129: 385-397.
50. Dinan, L. 1998. A strategy towards the elucidation of the contribution made by phytoecdysteroids to the deterrence of invertebrate predators on plants. Russ. J. Plant Phys. 45: 296-305.
51. -. 2001. Phytoecdysteroids: Biological aspects. Phytochemistry 57: 325-339.
52. Dinan, L., Savchenko, T., and Whiting, P. 2001. On the distribution of phytoecdysteroids in plants. Cell. Mol. Life Sci. 58: 1121-1132.
53. +-ATPase: A universal proton pump of eukaryotes. Biochem. J. 324: 697-712.
54. Fisk, G.J. and Thummel, C.S. 1995. Isolation, regulation, and DNA-binding properties of three Drosophila nuclear hormone receptor superfamily members. Proc. Natl. Acad. Sci. 92: 10604-10608.
55. FlyBase. 1999. The FlyBase database of the Drosophila genome projects and community literature. Nucleic Acids Res. 27: 85-88 http://flybase.bio.indiana.edu.
56. Friedrichsen, D. and Chory, J. 2001. Steroid signaling in plants: From the cell surface to the nucleus. Bioessays 23: 1028-1036.
57. Friedrichsen, D.M., Joazeiro, C.A., Li, J., Hunter, T., and Chory, J. 2000. Brassinosteroid-insensitive-1 is a ubiquitously expressed leucine-rich repeat receptor serine/threonine kinase. Plant Physiol. 123: 1247-1256.
58. Friedrichsen, D., Nemhauser, J., Muramitsu, T., Maloof, J., Alonso, J., Ecker, J., Furuya, M., and Chory, J. 2002. Three redundant brassinosteroid early response genes encode putative bHLH transcription factors required for normal growth. Genetics (in press).
59. Fristrom, D. and Fristrom, J.W. 1993. The metamorphic development of the adult epidermis. In The Development of Drosophila Melanogaster (eds. M. Bate and A. Martinez Arias), pp. 843-897. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, NY.
60. Fujioka, S., Li, J., Choi, Y.H., Seto, H., Takatsuto, S., Noguchi, T., Watanabe, T., Kuriyama, H., Yokota, T., Chory, J., et al. 1997. The Arabidopsis deetiolated2 mutant is blocked early in brassinosteroid biosynthesis. Plant Cell 9: 1951-1962.
61. Gates, J. and Thummel, C.S. 2000. An enhancer trap screen for ecdysone-inducible genes required for Drosophila adult leg morphogenesis. Genetics 156: 1765-1776.
62. Gilbert, L.I., Rybczynski, R., and Warren, J.T. 2002. Control and biochemical nature of the ecdysteroidogenic pathway. Annu. Rev. Entomol. 47: 883-916.
63. Gotwals, P.J. and Fristrom, J.W. 1991. Three neighboring genes interact with the Broad-Complex and the Stubble-stubbloid locus to affect imaginal disc morphogenesis in Drosophila. Genetics 127: 747-759.
64. Grieneisen, M. 1994. Recent advances in our knowledge of ecdysteroid biosynthesis in insects and crustaceans. Insect Biochem. Mol. Biol. 24: 115-132.
65. Grieneisen, M.L., Warren, J.T., and Gilbert, L.I. 1993. Early steps in ecdysteroid biosynthesis: Evidence for the involvement of cytochrome P-450 enzymes. Insect Biochem. Mol. Biol. 23: 13-23.
66. He, Z., Wang, Z.Y., Li, J., Zhu, Q., Lamb, C., Ronald, P., and Chory, J. 2000. Perception of brassinosteroids by the extracellular domain of the receptor kinase BRI1. Science 288: 2360-2363.
67. He, J.X., Gendron, J.M., Yang, Y., Li, J., and Wang, Z.Y. 2002. The GSK3-like kinase BIN2 phosphorylates and destabilizes BZR1, a positive regulator of the brassinosteroid signaling pathway in Arabidopsis. Proc. Natl. Acad. Sci. 99: 10185-10190.
68. Hewes, R.S., Schaefer, A.M., and Taghert, P.H. 2000. The cryptocephal gene (ATF4) encodes multiple basic-leucine zipper proteins controlling molting and metamorphosis in Drosophila. Genetics 155: 1711-1723.
69. Ho, M.N., Hill, K.J., Lindorfer, M.A., and Stevens, T.H. 1993. Isolation of vacuolar membrane H(+)-ATPase-deficient yeast mutants; the VMA5 and VMA 4 genes are essential for assembly and activity of the vacuolar H(+)-ATPase. J. Biol. Chem. 268: 221-227.
70. Hock, T., Cottrill, T., Keegan, J., and Garza, D. 2000. The E23 early gene of Drosophila encodes an ecdysone-inducible ATP-binding cassette transporter capable of repressing ecdysone-mediated gene activation. Proc. Natl. Acad. Sci. 97: 9519-9524.
71. Horner, M.A. and Thummel, C.S. 1997. Mutations in the DHR39 orphan receptor gene have no effect on viability. Drosoph. Inf. Serv. 80: 35-37.
72. Horner, M., Chen, T., and Thummel, C.S. 1995. Ecdysone regulation and DNA binding properties of Drosophila nuclear hormone receptor superfamily members. Dev. Biol. 168: 490-502.
73. Huelsken, J. and Birchmeier, W. 2001. New aspects of Wnt signaling pathways in higher vertebrates. Curr. Opin. Genet. Dev. 11: 547-553.
74. Hurban, P. and Thummel, C.S. 1993. Isolation and characterization of fifteen ecdysone-inducible Drosophila genes reveal unexpected complexities in ecdysone regulation. Mol. Cell Biol 13: 7101-7111.
75. Ikebawa, N. and Zhao, Y.-J. 1981. Application of 24-epibrassinolide in agriculture. In Brassinosteroids; Chemistry, Bioactivity and Applications (eds. H. Cutler, T. Yokota, and G. Adams), pp. 280-291. ACS Symposium Series, American Chemical Society, Washington, DC.
76. Imai, S., Toyosato, T., Sakai, M., Sato, Y., Fujioka, S., Murata, E., and Goto, M. 1969. Screening results of plants for phytoecdysones. Chem. Pharm. Bull. (Tokyo) 17: 335-339.
77. Iwasaki, T. and Shibaoka, H. 1991. Brassinosteroids act as regulators of tracheary-element differentiation in isolated Zinnia mesophyll cells. Plant Cell Physiol. 32: 1007-1014.
78. Jiang, C., Baehrecke, E.H., and Thummel, C.S. 1997. Steroid regulated programmed cell death during Drosophila metamorphosis. Development 124: 4673-4683.
79. Jochova, J., Zakeri, Z., and Lockshin, R.A. 1997. Rearrangement of the tubulin and actin cytoskeleton during programmed cell death in Drosophila salivary glands. Cell Death Differ. 4: 140-149.
80. Jones, C. and Firn, R. 1978. A role of phytoecdysteroids in bracken fern, Pteridium aquilinum (L.) Kuhn, as a defence against phytophagous insect attack. J. Chem. Ecol. 4: 117-138.
81. Jones, G., Wozniak, M., Chu, Y., Dhar, S., and Jones, D. 2001. Juvenile hormone III-dependent conformational changes of the nuclear receptor ultraspiracle. Insect Biochem. Mol. Biol. 32: 33-49.
82. Jürgens, G., Wieschaus, E., Nüsslein-Volhard, C., and Kluding, H. 1984. Mutations affecting the pattern of the larval cuticle in Drosophila melanogaster. II. Zygotic loci on the third chromosome. Roux's Arch. Dev. Biol. 193: 283-295.
83. Kang, J.G., Yun, J., Kim, D.H., Chung, K.S., Fujioka, S., Kim, J.I., Dae, H.W., Yoshida, S., Takatsuto, S., Song, P.S., et al. 2001. Light and brassinosteroid signals are integrated via a darkinduced small G protein in etiolated seedling growth. Cell 105: 625-636.
84. Karim, F.D. and Thummel, C.S. 1992. Temporal coordination of regulatory gene expression by the steroid hormone ecdysone. EMBO J. 11: 4083-4093.
85. Kauschmann, A., Jessop, A., Koncz, C., Szekeres, M., Willmitzer, L., and Altmann, T. 1996. Genetic evidence for an essential role of brassinosteroids in plant development. Plant J. 9: 701-713.
86. Klahre, U., Noguchi, T., Fujioka, S., Takatsuto, S., Yokota, T., Nomura, T., Yoshida, S., and Chua, N.H. 1998. The Arabidopsis DIMINUTO/DWARF1 gene encodes a protein involved in steroid synthesis. Plant Cell 10: 1677-1690.
87. Koelle, M.R., Talbot, W.S., Segraves, W.A., Bender, M.T., Cherbas, P., and Hogness, D.S. 1991. The Drosophila EcR gene encodes an ecdysone receptor, a new member of the steroid receptor superfamily. Cell 67: 59-77.
88. Koelle, M.R., Segraves, W.A., and Hogness, D.S. 1992. DHR3: A Drosophila steroid receptor homolog. Proc. Natl. Acad. Sci. 89: 6167-6171.
89. Koka, C.V., Cerny, R.E., Gardner, R.G., Noguchi, T., Fujioka, S., Takatsuto, S., Yoshida, S., and Clouse, S.D. 2000. A putative role for the tomato genes DUMPY and CURL-3 in brassinosteroid biosynthesis and response. Plant Physiol. 122: 85-98.
90. Lafont, R., Bouthier, A., and Wilson, I. 1991. Phytoecdysteroids: Structures, occurrence, biosynthesis and possible ecological significance. In Insect chemical ecology (ed. I. Hardy), pp. 197-214. Academica Prague, Prague, Czechoslovakia.
91. Lam, G., Hall, B.L., Bender, M., and Thummel, C.S. 1999. DHR3 is required for the prepupal-pupal transition and differentiation of adult structures during Drosophila metamorphosis. Dev. Biol. 212: 204-216.
92. Lee, C.Y. and Baehrecke, E.H. 2001. Steroid regulation of autophagic programmed cell death during development. Development 128: 1443-1455.
93. Li, J. and Chory, J. 1997. A putative leucine-rich repeat receptor kinase involved in brassinosteroid signal transduction. Cell 90: 929-938.
94. Li, J. and Nam, K.H. 2002. Regulation of brassinosteroid signaling by a GSK3/SHAGGY-like kinase. Science 295: 1299-1301.
95. Li, J., Nagpal, P., Vitart, V., McMorris, T.C., and Chory, J. 1996. A role for brassinosteroids in light-dependent development of Arabidopsis. Science 272: 398-401.
96. Li, J., Biswas, M.G., Chao, A., Russell, D.W., and Chory, J. 1997. Conservation of function between mammalian and plant steroid 5α-reductases. Proc. Natl. Acad. Sci. 94: 3554-3559.
97. Li, J., Lease, K.A., Tax, F.E., and Walker, J.C. 2001a. BRS1, a serine carboxypeptidase, regulates BRI1 signaling in Arabidopsis thaliana. Proc. Natl. Acad. Sci. 98: 5916-5921.
98. Li, J., Nam, K.H., Vafeados, D., and Chory, J. 2001b. Bin2, a new brassinosteroid-insensitive locus in Arabidopsis. Plant Physiol. 127: 14-22.
99. Li, J., Wen, J., Lease, K.A., Doke, J.T., Tax, F.E., and Walker, J.C. 2002. BAK1, an Arabidopsis LRR receptor-like protein kinase, interacts with BRI1 and modulates brassinosteroid signaling. Cell 110: 213-222.
100. Mandava, N.B. 1988. Plant growth-promoting brassinosteroids. Ann. Rev. Plant Physiol. Plant Mol. Biol. 39: 23-52.
101. Mathur, J., Molnar, G., Fujioka, S., Takatsuto, S., Sakurai, A., Yokota, T., Adam, G., Voigt, B., Nagy, F., Maas, C., et al. 1998. Transcription of the Arabidopsis CPD gene, encoding a steroidogenic cytochrome P450, is negatively controlled by brassinosteroids. Plant J. 14: 593-602.
102. Meyerowitz, E.M. 2002. Plants compared to animals: The broadest comparative study of development. Science 295: 1482-1485.
103. Millar, A., Straume, M., Chory, J., Chua, N.-H., and Kay, S. 1995. The regulation or circadian period by phototransduction pathways in Arabidopsis. Science 267: 1163-1166.
104. Mussig, C., Fischer, S., and Altmann, T. 2002. Brassinosteroidregulated gene expression. Plant Physiol. 129: 1241-1251.
105. Nam, K.H. and Li, J. 2002. BRI1/BAK1, a receptor kinase pair mediating brassinosteroid signaling. Cell 110: 203-212.
106. Neff, M.M., Nguyen, S.M., Malancharuvil, E.J., Fujioka, S., Noguchi, T., Seto, H., Tsubuki, M., Honda, T., Takatsuto, S., Yoshida, S., et al. 1999. BAS1: A gene regulating brassinosteroid levels and light responsiveness in Arabidopsis. Proc. Natl. Acad. Sci. 96: 15316-15323.
107. Noguchi, T., Fujioka, S., Choe, S., Takatsuto, S., Yoshida, S., Yuan, H., Feldmann, K.A., and Tax, F.E. 1999. Brassinosteroid-insensitive dwarf mutants of Arabidopsis accumulate brassinosteroids. Plant Physiol. 121: 743-752.
108. Noguchi, T., Fujioka, S., Choe, S., Takatsuto, S., Tax, F.E., Yoshida, S., and Feldmann, K.A. 2000. Biosynthetic pathways of brassinolide in Arabidopsis. Plant Physiol. 124: 201-209.
109. Nomura, T., Kitasaka, Y., Takatsuto, S., Reid, J.B., Fukami, M., and Yokota, T. 1999. Brassinosteroid/sterol synthesis and plant growth as affected by 1ka and 1kb mutations of pea. Plant Physiol. 119: 1517-1526.
110. Nüsslein-Volhard, C., Wieschaus, E., and Kluding, H. 1984. Mutations affecting the pattern of the larval cuticle in Drosophila melanogaster. I. Zygotic loci on the second chromosome. Roux's Arch. Dev. Biol. 183: 267-282.
111. Oh, M.-H., Romanow, W., Smith, R., Zamski, E., Sasse, J., and Clouse, S. 1998. Soybean BRU1 encodes a functional xyloglucan endotransglycosylase that is highly expressed in inner epicotyl tissues during brassinosteroid-promoted elongation. Plant Cell Physiol. 39.
112. Oh, M.H., Ray, W.K., Huber, S.C., Asara, J.M., Gage, D.A., and Clouse, S.D. 2000. Recombinant brassinosteroid insensitive 1 receptor-like kinase autophosphorylates on serine and threonine residues and phosphorylates a conserved peptide motif in vitro. Plant Physiol. 124: 751-766.
113. Ohno, C.K. and Petkovich, M. 1993. FTZ-F1 β, a novel member of the Drosophila nuclear receptor family. Mech. Dev. 40: 13-24.
114. Pecasse, F., Beck, Y., Ruiz, C., and Richards, G. 2000. Kruppelhomolog, a stage-specific modulator of the prepupal ecdysone response, is essential for Drosophila metamorphosis. Dev. Biol. 221: 53-67.
115. Perez-Perez, J.M., Ponce, M.R., and Micol, J.L. 2002. The UCU1 Arabidopsis gene encodes a SHAGGY/GSK3-like kinase required for cell expansion along the proximodistal axis. Dev. Biol. 242: 161-173.
116. Rees, H. 1995. Ecdysteroid biosynthesis and inactivation in relation to function. Eur. J. Entomol. 92: 9-39.
117. Renault, N., King-Jones, K., and Lehmann, M. 2001. Downregulation of the tissue-specific transcription factor Fork head by Broad-Complex mediates a stage-specific hormone response. Development 128: 3729-3737.
118. Restifo, L.L. and Wilson, T.G. 1998. A juvenile hormone agonist reveals distinct developmental pathways mediated by ecdysone-inducible Broad Complex transcription factors. Dev. Genet. 22: 141-159.
119. Richards, G. 1976. Sequential gene activation by ecdysone in polytene chromosomes of Drosophila melanogaster. IV. The mid-prepupal period. Dev. Biol. 54: 256-263.
120. -. 1981. The radioimmune assay of ecdysteroid titres in Drosophila melanogaster. Mol. Cell. Endocrinol. 21: 181-197.
121. -. 1997. The ecdysone regulatory cascades in Drosophila. Adv. in Dev. Biol. 5: 81-135.
122. Riddiford, L.M. 1993. Hormones and Drosophila development. In The Development of Drosophila Melanogaster (eds. M. Bate and A. Martinez-Arias), pp. 899-939. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, NY.
123. Riddiford, L. 1996. Molecular aspects of juvenile hormone action in insect metamorphosis. In Metamorphosis: Postembryonic Reprogramming of Gene Expression in Amphibian and Insect Cells (eds. L. Gilbert, J. Tata, and B. Atkinson), pp. 223-251. Academic Press, New York.
124. Riddiford, L.M., Cherbas, P., and Truman, J.W. 2000. Ecdysone receptors and their biological actions. Vitam. Horm. 60: 173.
125. Roberts, K. and McCann, M.C. 2000. Xylogenesis: The birth of a corpse. Curr. Opin. Plant. Biol. 3: 517-522.
126. Robertson, C.W. 1936. The metamorphosis of Drosophila melanogaster, including an accurately timed account of the principal morphological changes. J. Morphol. 59: 351-399.
127. Rouleau, M., Marsolais, F., Richard, M., Nicolle, L., Voigt, B., Adam, G., and Varin, L. 1999. Inactivation of brassinosteroid biological activity by a salicylate-inducible steroid sulfotransferase from Brassica napus. J. Biol. Chem. 274: 20925-20930.
128. Russell, S. and Ashburner, M. 1996. Ecdysone-regulated chromosome puffing in Drosophila melanogaster. In Metamorphosis. Postembryonic Reprogramming of Gene Expression in Amphibian and Insect Cells (eds. B.G. Atkinson, L.I. Gilbert, and J.R. Tata), pp. 109-144. Academic Press, New York.
129. Russell, D.W. and Wilson, J.D. 1994. Steroid 5a-reductase: Two genes/two enzymes. Annu. Rev. Biochem. 63: 25-61.
130. Russell, S.R., Heimbeck, G., Goddard, C.M., Carpenter, A.T., and Ashburner, M. 1996. The Drosophila Eip78C gene is not vital but has a role in regulating chromosome puffs. Genetics 144: 159-170.
131. Sakurai, A. and Fujioka, S. 1997. Studies on biosynthesis of brassinosteroids. Biosci. Biotechnol. Biochem. 61: 757-762.
132. Schmelz, E., Grebenok, R., Galbraith, D., and Bowers, W. 1998. Damage-induced accumulation of phytoecdysteroids in spinach: A rapid root response involving the octadecanoic acid pathway. J. Chem. Ecol. 24: 339-360.
133. Schmelz, E., Grebenok, R., Galbraith, D., and Bowers, W. 1999. Insect-induced synthesis of phyecdysteroids in spinach, Spinacia oleracea. J. Chem. Ecol. 25: 1739-1757.
134. Schumacher, K., Vafeados, D., McCarthy, M., Sze, H., Wilkins, T., and Chory, J. 1999. The Arabidopsis det3 mutant reveals a central role for the vacuolar H(+)-ATPase in plant growth and development. Genes & Dev. 13: 3259-3270.
135. Segraves, W. 1998. Ecdysone response in Drosophila. In Hormones and Growth Factors in Development and Neoplasia (eds. R. Dickson and D. Salomon), pp. 45-78. John Wiley & Sons, New York.
136. Segraves, W.A. and Hogness, D.S. 1990. The E75 ecdysone-in-ducible gene responsible for the 75B early puff in Drosophila encodes two new members of the steroid receptor superfamily. Genes & Dev. 4: 204-219.
137. Sharpe, C., Lawrence, N., and Martinez Arias, A. 2001. Wnt signalling: A theme with nuclear variations. Bioessays 23: 311-318.
138. Siegmund, T. and Lehmann, M. 2002. The Drosophila Pipsqueak protein defines a new family of helix-turn-helix DNA-binding proteins. Dev. Genes Evol. 212: 152-157.
139. Song, Q. and Gilbert, L.I. 1998. Alterations in ultraspiracle (USP) content and phosphorylation state accompany feedback regulation of ecdysone synthesis in the insect prothoracic gland. Insect Biochem. Mol. Biol. 28: 849-860.
140. Stone, B.L. and Thummel, C.S. 1993. The Drosophila 78C early late puff contains E78, an ecdysone-inducible gene that encodes a novel member of the nuclear hormone receptor superfamily. Cell 75: 307-320.
141. Strott, C.A. 1996. Steroid sulfotransferases. Endocr. Rev. 17: 670-697.
142. Szekeres, M., Nemeth, K., Koncz-Kalman, Z., Mathur, J., Kauschmann, A., Altmann, T., Redei, G.P., Nagy, F., Schell, J., and Koncz, C. 1996. Brassinosteroids rescue the deficiency of CYP90, a cytochrome P450, controlling cell elongation and deetiolation in Arabidopsis. Cell 85: 171-182.
143. Tanaka, Y., Asaoka, K., and Takeda, S. 1994. Different feeding and gustatory responses to ecdysone and 20-hydroxyecdysone by larvae of the silkworm, Bombyxmori. J. Chem. Ecol. 20: 125-133.
144. The Arabidopsis Genome Initiative. 2000. Analysis of the genome sequence of the flowering plant Arabidopsis thaliana. Nature 408: 796-815.
145. Thomas, H.E., Stunnenberg, H.G., and Stewart, A.F. 1993. Heterodimerization of the Drosophila ecdysone receptor with retinoid X receptor and ultraspiracle. Nature 362: 471-475.
146. Thummel, C.S. 1995. From embryogenesis to metamorphosis: The regulation and function of Drosophila nuclear receptor superfamily members. Cell 83: 871-877.
147. -. 1996. Flies on steroids - Drosophila metamorphosis and the mechanisms of steroid hormone action. Trends Genet. 12: 306-310.
148. von Kalm, L., Fristrom, D., and Fristrom, J. 1995. The making of a fly leg: A model for epithelial morphogenesis. Bioessays 17: 693-702.
149. Wang, Z.Y., Seto, H., Fujioka, S., Yoshida, S., and Chory, J. 2001. BRI1 is a critical component of a plasma-membrane receptor for plant steroids. Nature 410: 380-383.
150. Wang, Z.Y., Nakano, T., Gendron, J., He, J., Chen, M., Vafeados, D., Yang, Y., Fujioka, S., Yoshida, S., Asami, T., et al. 2002. Nuclear-localized BZR1 mediates brassinosteroid-induced growth and feedback suppression of brassinosteroid biosynthesis. Dev. Cell 2: 505-513.
151. Warren, J.T., Bachmann, J.S., Dai, J.D., and Gilbert, L.I. 1996. Differential incorporation of cholesterol and cholesterol derivatives into ecdysteroids by the larval ring glands and adult ovaries of Drosophila melanogaster: A putative explanation for the 1(3)ecd1 mutation. Insect Biochem. Mol. Biol. 26: 931-943.
152. Warren, J.T., Petryk, A., Marques, G., Jarcho, M., Parvy, J.P., Dauphin-Villemant, C., O'Connor, M.B., and Gilbert, L.I. 2002. Molecular and biochemical characterization of two P450 enzymes in the ecdysteroidogenic pathway of Drosophila melanogaster. Proc. Natl. Acad. Sci. 99: 11043-11048.
153. White, K.P., Hurban, P., Watanabe, T., and Hogness, D.S. 1997. Coordination of Drosophila metamorphosis by two ecdysone-induced nuclear receptors. Science 276: 114-117.
154. White, K.P., Rifkin, S.A., Hurban, P., and Hogness, D.S. 1999. Microarray analysis of Drosophila development during metamorphosis. Science 286: 2179-2184.
155. Wieschaus, E., Nüsslein-Volhard, C., and Jürgens, G. 1984. Mutations affecting the pattern of the larval cuticle in Drosophila melanogaster. III. Zygotic loci on the X-chromosome and fourth chromosome. Roux's Arch. Dev. Biol. 193: 296-307.
156. Winter, J., Bilbe, G., Richener, H., Sehringer, B., and Kayser, H. 1999. Cloning of a cDNA encoding a novel cytochrome P450 from the insect Locusta migratoria: CYP6H1, a putative ecdysone 20-h-ydroxylase. Biochem. Biophys. Res. Commun. 259: 305-310.
157. Woodgett, J.R. 2001. Judging a protein by more than its name: gsk-3. Sci. STKE 2001: RE12.
158. Xu, W., Purugganan, M.M., Polisensky, D.H., Antosiewicz, D.M., Fry, S.C., and Braam, J. 1995. Arabidopsis TCH4, regulated by hormones and the environment, encodes a xyloglucan endotransglycosylase. Plant Cell 7: 1555-1567.
159. Yamamoto, R., Fujioka, S., Demura, T., Takatsuto, S., Yoshida, S., and Fukuda, H. 2001. Brassinosteroid levels increase drastically prior to morphogenesis of tracheary elements. Plant Physiol. 125: 556-563.
160. Yamamuro, C., Ihara, Y., Wu, X., Noguchi, T., Fujioka, S., Takatsuto, S., Ashikari, M., Kitano, H., and Matsuoka, M. 2000. Loss of function of a rice brassinosteroid insensitive1 homolog prevents internode elongation and bending of the lamina joint. Plant Cell 12: 1591-1606.
161. Yao, T., Segraves, W.A., Oro, A.E., McKeown, M., and Evans, R.M. 1992. Drosophila ultraspiracle modulates ecdysone receptor function via heterodimer formation. Cell 71: 63-72.
162. Yao, T., Forman, B.M., Jiang, Z., Cherbas, L., Chen, J.D., McKeown, M., Cherbas, P., and Evans, R.M. 1993. Functional ecdysone receptor is the product of EcR and ultraspiracle genes. Nature 366: 476-479.
163. Yin, Y., Wang, Z.Y., Mora-Garcia, S., Li, J., Yoshida, S., Asami, T., and Chory, J. 2002. BES1 accumulates in the nucleus in response to brassinosteroids to regulate gene expression and promote stem elongation. Cell 109: 181-191.
164. Zhou, X. and Riddiford, L.M. 2002. Broad specifies pupal development and mediates the "status quo" action of juvenile hormone on the pupal-adult transformation in Drosophila and Manduca. Development 129: 2259-2269.
165. Zurek, D.M. and Clouse, S.D. 1994. Molecular cloning and characterization of a brassinosteroid-regulated gene from elongating soybean (Glycine max L.) epicotyls. Plant Physiol. 104: 161-170.
166. Zurek, D.M., Rayle, D.L., McMorris, T.C., and Clouse, S.D. 1994. Investigation of gene expression, growth kinetics, and wall extensibility during brassinosteriod-regulated stem elongation. Plant Physiol. 104: 505-513.