Gonadal ecdysteroidogenesis in arthropoda: Occurrence and regulation

Annual Review of Entomology

Volumn 54, Issue , 2009, Pages 105-125

  Brown, Mark R ^a   Sieglaff, Douglas H ^b   Rees, Huw H ^c  

^a UNIVERSITY OF GEORGIA   (United States)

^b UNIVERSITY OF CALIFORNIA   (United States)

^c UNIVERSITY OF LIVERPOOL   (United Kingdom)

Author keywords

Hormone; Ovary; Reproduction; Testes

Indexed keywords

ARTHROPODA; CRUSTACEA; HEXAPODA; INVERTEBRATA; IXODIDA; VERTEBRATA;

ECDYSTEROID;

ANIMAL; ARTHROPOD; BIOSYNTHESIS; GONAD; HISTOLOGY; METABOLISM; REVIEW;

ANIMALS; ARTHROPODS; ECDYSTEROIDS; GONADS;

EID: 60549107098     PISSN: 00664170     EISSN: None     Source Type: Book Series    
DOI: 10.1146/annurev.ento.53.103106.093334     Document Type: Article

References (166)

1. Adams TS, Gerst JW, Masler EP. 1997. Regulation of ovarian ecdysteroid production in the housefly, Musca domestica. Arch. Insect Biochem. Physiol. 35:135-48
2. Adams TS, Li QJ. 1998. Ecdysteroidostatin from the house fly, Musca domestica. Arch. Insect Biochem. Physiol. 38:166-76
3. Attardo GM, Hansen IA, Raikhel AS. 2005. Nutritional regulation of vitellogenesis in mosquitoes: implications for anautogeny. Insect Biochem. Mol. Biol. 35:661-75
4. Badisco L, Claeys I, Van Loy T, VanHiel M, Franssens V, et al. 2007. Neuroparsins, a family of conserved arthropod neuropeptides. Gen. Comp. Endocrinol. 153:64-71
5. Behmer ST, Nes WD. 2003. Insect sterol nutrition and physiology: a global overview. In Advances in Insect Physiology, ed. SJ Simpson, pp. 1-72. Amsterdam: Elsevier
6. Blais C, Dauphin-Villemant C, Kovganko N, Girault JP, Descoins C Jr, Lafont R. 1996. Evidence for the involvement of 3-oxo-delta 4 intermediates in ecdysteroid biosynthesis. Biochem. J. 320:413-19
7. Block DS, Bejarano AC, Thomas Chandler G. 2003. Ecdysteroid concentrations through various lifestages of the meiobenthic harpacticoid copepod, Amphiascus tenuiremis and the benthic estuarine amphipod, Leptocheirus plumulosus. Gen. Comp. Endocrinol. 132:151-60
8. Broué F, Liere P, Kenyon C, Baulieu EE. 2007. A steroid hormone that extends the lifespan of Caenorhabditis elegans. Aging Cell 6:87-94
9. Brown MR, Clark KD, Gulia M, Zhao Z, Garczynski SF, et al. 2008. An insulin-like peptide regulates egg maturation and metabolism in the mosquito Aedes aegypti. Proc. Natl. Acad. Sci. USA 105:5716-21
10. Brown MR, Graf R, Swiderek K. 1995. Structure and function of mosquito gonadotropins. In Molecular Mechanisms of Insect Metamorphosis and Diapause, ed. A Suzuki, H Kataoka, S Matsumoto, pp. 229-38. Tokyo: Ind. Publ. Consult.
11. Brown MR, Graf R, Swiderek KM, Fendley D, Stracker TH, et al. 1998. Identification of a steroidogenic neurohormone in female mosquitoes. J. Biol. Chem. 273:3967-71
12. Chang ES, Kaufman WR. 2005. Endocrinology of Crustacea and Chelicerata. See Ref. 36, 3:805-42
13. Chavez VM, Marques G, Delbecque JP, Kobayashi K, Hollingsworth M, et al. 2000. The Drosophila disembodied gene controls late embryonic morphogenesis and codes for a cytochrome P450 enzyme that regulates embryonic ecdysone levels. Development 127:4115-26
14. Chen JH, Turner PC, Rees HH. 1999. Molecular cloning and characterization of hemolymph 3-dehydroecdysone 3beta-reductase from the cotton leafworm, Spodoptera littoralis. A new member of the third superfamily of oxidoreductases. J. Biol. Chem. 274:10551-56
15. Chino H, Gilbert LI. 1971. The uptake and transport of cholesterol by haemolymph lipoproteins. Insect Biochem. 1:337-47
16. Conn DB. 2000. Atlas of Invertebrate Reproduction and Development. New York:Wiley. 300 pp.
17. Darvas B, Szekacs A, Fonagy A, Szesi M, Toth I. 1997. Progesterone in Periplaneta americana and Neobellieria bullata adults from procuticle phase until first progeny production. Gen. Comp. Endocrinol. 107:450-60
18. Dauphin-Villemant C, Bocking D, Blais C, Toullec JY, Lafont R. 1997. Involvement of a 3betahydroxysteroid dehydrogenase activity in ecdysteroid biosynthesis. Mol. Cell. Endocrinol. 128:139-49
19. Davies TG, Dinan LN, Lockley WJ, Rees HH, Goodwin TW. 1981. Formation of the A/B cis ring junction of ecdysteroids in the locust, Schistocerca gregaria. Biochem. J. 194:53-62
20. Delbecque JP, Weidner K, Hoffmann JA. 1990. Alternative sites for ecdysteroid production in insects. Invertebr. Reprod. Dev. 18:29-42
21. De Loof A, Baggerman G, Breuer M, Claeys I, Cerstiaens A, et al. 2001. Gonadotropins in insects: an overview. Arch. Insect Biochem. Physiol. 47:129-38
22. Descamps M, Charlet M, Leu B. 1990. Conversion of 2,22,25- trideoxyecdysone (5B-ketodiol) by body parts and gonads of Lithobius forflcatus L. (Myriapoda, Chilopoda). Invertebr. Reprod. Dev. 18:111
23. Diehl PA, Connat JL, Dotson E. 1986. Chemistry, function and metabolism of tick ecdysteroids. In Morphology, Physiology, and Behavioral Biology of Ticks, ed. JR Sauer, JA Hair, pp. 165-93. Chichester, UK: Horwood
24. Dinan LN. 1997. Ecdysteroids in adults and eggs of the house cricket, Acheta domesticus (Orthoptera: Gryllidae). J. Comp. Biochem. Physiol. B 116: 129-35
25. Dinan LN, Rees HH. 1981. Incorporation in vivo of [4-14C]-cholesterol into the conjugated ecdysteroids in ovaries and eggs of Schistocerca americana gregaria. Insect Biochem. 11:255-65
26. 2-1,2]-7-dehydrocholesterol. Conversion to ecdysone in follicle cells of Locusta (Insects). Tetrahedron 46:5305-16
27. Entchev EV, Kurzchalia TV. 2005. Requirement of sterols in the life cycle of the nematode Caenorhabditis elegans. Semin. Cell Dev. Biol. 16:175
28. Escobedo G, Roberts CW, Carrero JC, Morales-Montor J. 2005. Parasite regulation by host hormones: an old mechanism of host exploitation? Trends Parasitol. 21:588-93
29. Fluegel ML, Parker TJ, Pallanck LJ. 2006. Mutations of a Drosophila NPC1 gene confer sterol and ecdysone metabolic defects. Genetics 172:185-96
30. Freeman MR, Dobritsa A, Gaines P, Segraves WA, Carlson JR. 1999. The dare gene: steroid hormone production, olfactory behavior, and neural degeneration in Drosophila. Development 126:4591-602
31. Fujimoto Y, Hiramoto M, Kakinuma K, Ikekawa N. 1989. Elimination of C-6-hydrogen during the formation of ecdysteroids from cholesterol in Locusta migratoria ovaries. Steroids 53:477-85
32. Fullbright G, Lacy ER, Bullesbach EE. 1997. The prothoracicotropic hormone bombyxin has specific receptors on insect ovarian cells. Eur. J. Biochem. 245:774-80
33. Gaziova I, Bonnette PC, Henrich VC, Jindra M. 2004. Cell-autonomous roles of the ecdysoneless gene in Drosophila development and oogenesis. Development 131:2715-25
34. Gerisch B, Rottiers V, Li D, Motola DL, Cummins CL, et al. 2007. A bile acid-like steroid modulates Caenorhabditis elegans lifespan through nuclear receptor signaling. Proc. Natl. Acad. Sci. USA 104:5014-19
35. Gilbert LI. 2004. Halloween genes encode P450 enzymes that mediate steroid hormone biosynthesis in Drosophila melanogaster. Mol. Cell. Endocrinol. 215:1-10
36. Gilbert LI, Gill S, Iatrou K, eds. 2005. Comprehensive Molecular Insect Science, Vol. 3. Amsterdam: Elsevier
37. Gilbert LI, Rybczynski R, Warren JT. 2002. Control and biochemical nature of the ecdysteroidogenic pathway. Annu. Rev. Entomol. 47:883-916
38. Glass H, Emmerich H, Spindler K.-D. 1978. Immunohistochemical localisation of ecdysteroids in the follicle epithelium of locust oocytes. Cell Tissue Res. 194:237-44
39. Goltzene F, Lagueux M, Charlet M, Hoffmann JA. 1978. The follicle cell epithelium of maturing ovaries of Locusta migratoria: a new biosynthetic tissue for ecdysone. Hoppe Seylers Z Physiol. Chem. 359:1427-34
40. Graf R, Neuenschwander S, Brown MR, Ackermann U. 1997. Insulin-mediated secretion of ecdysteroids from mosquito ovaries and molecular cloning of the insulin receptor homologue from ovaries of bloodfed Aedes aegypti. Insect Mol. Biol. 6:151-63
41. Greenwood DR, Dinan LN, Rees HH. 1984. Mechanism of hydroxylation at C-2 during the biosynthesis of ecdysone in ovaries of the locust, Schistocerca gregaria. Biochem. J. 217:783-89
42. Grieneisen ML. 1994. Recent advances in our knowledge of ecdysteroid biosynthesis in insects and crustaceans. Insect Biochem. Mol. Biol. 24:115-32
43. Grieneisen ML, Warren JT, Gilbert LI. 1993. Early steps in ecdysteroid biosynthesis: evidence for the involvement of cytochrome P-450 enzymes. Insect Biochem. Mol. Biol. 23:13-23
44. Grimaldi DA, Engel MS. 2004. Evolution of the Insects. Cambridge, UK: Cambridge Univ. Press. 778 pp.
45. Gupta AP. 1990. Morphogenetic hormones and their glands in arthropods: evolutionary aspects. See Ref. 46, pp. 1-34
46. Gupta AP, ed. 1990. Morphogenetic Hormones of Arthropods: Discoveries, Syntheses, Metabolism, Evolution, Modes of Action, and Techniques, Vol. 1. New Brunswick, NJ: Rutgers Univ. Press
47. Hagedorn HH. 1985. The role of ecdysteroids in reproduction. See Ref. 73, 8:205-62
48. Hagedorn HH. 1989. Physiological roles of hemolymph ecdysteroids in the adult insect. See Ref. 77, pp. 279-89
49. Hagedorn HH, O'Connor JD, Fuchs MS, Sage B, Schlaeger DA, Bohm MK. 1975. The ovary as a source of alpha-ecdysone in an adult mosquito. Proc. Natl. Acad. Sci. USA 72:3255-59
50. Hatle JD, Juliano SA, Borst DW. 2003. Hemolymph ecdysteroids do not affect vitellogenesis in the lubber grasshopper. Arch. Insect Biochem. Physiol. 52:45-57
51. Hetru C, Lagueux M, Bang L, Hoffmann JA. 1978. Adult ovaries of Locusta migratoria contain the sequence of biosynthetic intermediates for ecdysone. Life Sci. 22:2141-54
52. Hetru C, Lagueux M, Lachaise F, Hoffman JA. 1978. Adult ovaries of Locusta migratoria and ovaries of Carcinus maenas during vitellogenesis contain the sequence of biosynthetic intermediates for ecdysone. In Comparative Endocrinology, ed. PJ Gaillard, HH Boer, pp. 515-16. Amsterdam: Elsevier/North-Holland Biomed.
53. Hetru CC, Kappler C, Hoffmann JA, Nearn R, Bang L, Horn DH. 1982. The biosynthetic pathway of ecdysone: studies with vitellogenic ovaries of Locusta migratoria (Orthoptera). Mol. Cell. Endocrinol. 26:51-80
54. Hoffmann JA, Lagueux M. 1985. Endocrine aspects of embryonic development in insects. See Ref. 73, 1:435-60
55. Hoffmann JA, Weidner K, Seidel M. 1992. Sites of ecdysteroid biosynthesis in female adults of Gryllus bimaculatus (Ensifera, Gryllidae). J. Comp. Physiol. B 162:731-39
56. Hoffmann KH. 1982. Free ecdysteroids in adult male crickets, Gryllus bimaculatus. Physiol. Entomol. 7:269-79
57. Hoffmann KH, Dettner K, Tomaschko KH. 2006. Chemical signals in insects and other arthropods: from molecular structure to physiological functions. Physiol. Biochem. Zool. 79:344-56
58. Horike N, Takemori H, Nonaka Y, Sonobe H, Okamoto M. 2000. Molecular cloning of NADPHcytochrome P450 oxidoreductase from silkworm eggs: its involvement in 20-hydroxyecdysone biosynthesis during embryonic development. Eur. J. Biochem. 267:6914-20
59. Huang X, Suyama K, Buchanan J, Zhu AJ, Scott MP. 2005. A Drosophila model of the Niemann-Pick type C lysosome storage disease: dnpc1a is required for molting and sterol homeostasis. Development 132:5115-24
60. Huang X, Warren JT, Buchanan J, Gilbert LI, Scott MP. 2007. Drosophila Niemann-Pick type C-2 genes control sterol homeostasis and steroid biosynthesis: a model of human neurodegenerative disease. Development 134:3733-42
61. Huang X, Warren JT, Gilbert LI. 2008. New players in the regulation of ecdysone biosynthesis. J. Genet. Genomics 35:1-10
62. Jaenicke E, Foll R, Decker H. 1999. Spider hemocyanin binds ecdysone and 20-OH-ecdysone. J. Biol. Chem. 274:34267-71
63. Janer G, Porte C. 2007. Sex steroids and potential mechanisms of nongenomic endocrine disruption in invertebrates. Ecotoxicology 16:145-60
64. Jarvis TD, Earley GP, Rees HH. 1994. Ecdysteroid biosynthesis in larval testes of Spodoptera littoralis. Insect Biochem. Mol. Biol. 24:531-37
65. Jin X, Sun X, Song Q. 2005. woc gene mutation causes 20E-dependent alpha-tubulin detyrosination in Drosophila melanogaster. Arch. Insect Biochem. Physiol. 60:116-29
66. Jouni ZE, Takada N, Gazard J, Maekawa H, Wells MA, Tsuchida K. 2003. Transfer of cholesterol and diacylglycerol from lipophorin to Bombyx mori ovarioles in vitro: role of the lipid transfer particle. Insect Biochem. Mol. Biol. 33:145-53
67. Jouni ZE, Zamora J, Wells MA. 2002. Absorption and tissue distribution of cholesterol in Manduca sexta. Arch. Insect Biochem. Physiol. 49:167-75
68. Kappler C, Goltzene F, Lagueux M, Hetru C, Hoffman JA. 1986. Role of the follicle cells and the oocytes in ecdysone biosynthesis and esterification in vitellogenic females of Locusta migratoria. Int. J. Invertebr. Reprod. Dev. 9:17-34
69. Kappler C, Hetru C, Durst F, Hoffmann JA. 1989. Enzymes involved in ecdysone biosynthesis. See Ref. 77, pp. 161-66
70. Kappler C, Kabbouh M, Hetru C, Durst F, Hoffmann JA. 1988. Characterization of three hydroxylases involved in the final steps of biosynthesis of the steroid hormone ecdysone in Locusta migratoria (Insecta, Orthoptera). J. Steroid Biochem. 31:891-98
71. Kawooya JK, Law JH. 1988. Role of lipophorin in lipid transport to the insect egg. J. Biol. Chem. 263:8748-53
72. Kawooya JK, Osir EO, Law JH. 1988. Uptake of the major hemolymph lipoprotein and its transformation in the insect egg. J. Biol. Chem. 263:8740-47
73. Kerkut GA, Gilbert LI, eds. 1985. Comprehensive Insect Physiology, Biochemistry, and Pharmacology, Vols. 1, 7, 8. New York: Pergamon
74. Kiriishi S, Rountree DB, Sakurai S, Gilbert LI. 1990. Prothoracic gland synthesis of 3-dehydroecdysone and its hemolymph 3 beta-reductase mediated conversion to ecdysone in representative insects. Experientia 46:716-21
75. Kolmer M, Roos C, Tirronen M, Myohanen S, Alho H. 1994. Tissue-specific expression of the diazepambinding inhibitor in Drosophila melanogaster: cloning, structure, and localization of the gene. Mol. Cell Biol. 14:6983-95
76. Koolman J. 1982. Ecdysone metabolism. Insect Biochem. 12:225-50
77. Koolman J, ed. 1989. Ecdysone: From Chemistry to Mode of Action. New York: Thieme Med. Publ.
78. Krebs KC, Lan Q. 2003. Isolation and expression of a sterol carrier protein-2 gene from the yellow fever mosquito, Aedes aegypti. Insect Mol. Biol. 12:51-60
79. Lachaise F. 1990. Synthesis, metabolism, and effects on molting ecdysteroids in Crustacea, Chelicerata, and Myriapoda. See Ref. 46, pp. 275-323
80. Lafont R, Dauphin-Villemant C, Warren JT, Rees HH. 2005. Ecdysteroid chemistry and biochemistry. See Ref. 36, 3:125-95
81. Lafont R, Mathieu M. 2007. Steroids in aquatic invertebrates. Ecotoxicology 16:109-30
82. Lagueux M, Hirn M, Hoffmann JA. 1977. Ecdysone during ovarian development in Locusta migratoria. J. Insect Physiol. 23:109-19
83. Lagueux M, Hoffmann JA, Goltzené F, Kappler C, Tsoupras G, et al. 1984. Ecdysteroids in ovaries and embryos of Locusta migratoria. In Biosynthesis, Metabolism and Mode of Action of Invertebrate Hormones, ed. JA Hoffmann, M Porchet, pp. 168-80. Berlin: Springer-Verlag
84. Lanot R, Cledon P. 1989. Ecdysteroids and meiotic reinitiation in Palaemon serratus (Crustacea: Decapoda: Natantia) and in Locusta migratoria (Insecta: Orthoptera). A comparative study. Invertebr. Reprod. Dev. 16:169-75
85. Lanot R, Dorn A, Gunster B, Thiebold J, Lagueux M, Hoffmann JA. 1989. Functions of ecdysteroids in oocyte maturation and embryonic development of insects. See Ref. 77, pp. 262-70
86. Lanot R, Thiebold J, Costet-Corio MF, Benveniste P, Hoffmann JA. 1988. Further experimental evidence for the involvement of ecdysone in the control of meiotic reinitiation in oocytes of Locusta migratoria(Insecta, Orthoptera). Dev. Biol. 126:212-14
87. Laudet V, Bonneton F. 2005. Evolution of nuclear hormone receptors in insects. See Ref. 36, 3:287-318
88. Lea AO, Brown MR. 1990. Neuropeptides of mosquitoes. In Molecular Insect Science, ed. JH Law, pp. 181-88. New York: Plenum
89. Lea AO, Van Handel E. 1982. A neurosecretory hormone-releasing factor from ovaries of mosquito fed blood. J. Insect Physiol. 28:503-8
90. Leubert VF, Eibisch H, Kroschwitz H, Scheffel H. 1982. Ecdysteroid-biosynthese durch das ovar von Lithobius forflcatus (L.) (Chilopoda). Zool. J. Physiol. 86:465-76
91. Loeb M, Woods CW, Brandt EP, Borkovec AB. 1982. Larval testes of the tobacco budworm: a new source of insect ecdysteroids. Science 218:896-98
92. Loeb MJ. 1986. Ecdysteroids in testis sheaths of Heliothis virescens larvae: an immunocytological study. Arch. Insect Biochem. Physiol. 3:173-80
93. Loeb MJ, De Loof A, Gelman DB, Hakim RS, Jaffe H, et al. 2001. Testis ecdysiotropin, an insect gonadotropin that induces synthesis of ecdysteroid. Arch. Insect Biochem. Physiol. 47:181-88
94. Lomas LO, Rees HH. 1998. Endocrine regulation of development and reproduction in acarines. In Recent Advances in Arthropod Endocrinology, ed. GM Coast, SG Webster, pp. 91-124. Cambridge, UK: Cambridge Univ. Press
95. Lorenz MW, Lorenz JI, Treiblmayr K, Hoffmann KH. 1998. In vivo effects of allatostatins in crickets, Gryllus bimaculatus (Ensifera: Gryllidae). Arch. Insect Biochem. Physiol. 38:32-43
96. Maniere G, Rondot I, Bullesbach EE, Gautron F, Vanhems E, Delbecque JP. 2004. Control of ovarian steroidogenesis by insulin-like peptides in the blowfly (Phormia regina). J. Endocrinol. 181:147-56
97. Maniere G, Vanhems E, Delbecque JP. 2000. Cyclic AMP-dependent and independent stimulation of ovarian steroidogenesis by brain factors in the blowfly, Phormia regina. Mol. Cell. Endocrinol. 168:31-40
98. Maniere G, Vanhems E, Gautron F, Delbecque JP. 2002. Calcium inhibits ovarian steroidogenesis in the blowfly Phormia regina. J. Endocrinol. 173:533-44
99. Maniere G, Vanhems E, Gautron F, Delbecque JP. 2003. Inhibition of ovarian steroidogenesis by cyclic GMP in a fly. J. Endocrinol. 177:35-44
100. Meister MF, Dimarcq JL, Kappler C, Hetru C, Lagueux M, et al. 1985. Conversion of a radiolabelled ecdysone precursor, 2,22,25-trideoxyecdysone, by embryonic and larval tissues of Locusta migratoria. Mol. Cell. Endocrinol. 41:27-44
101. Mizuno T, Watanabe K, Ohnishi E. 1981. Developmental changes of ecdysteroids in the eggs of the silkworm, Bombyx mori. Dev. Growth Differ. 23:543-52
102. Neubueser D, Warren JT, Gilbert LI, Cohen SM. 2005. molting defective is required for ecdysone biosynthesis. Dev. Biol. 280:362-72
103. Niwa R, Matsuda T, Yoshiyama T, Namiki T, Mita K, et al. 2004. CYP306A1, a cytochrome P450 enzyme, is essential for ecdysteroid biosynthesis in the prothoracic glands of Bombyx and Drosophila. J. Biol. Chem. 279:35942-49
104. Ogihara K, Horigane M, Nakajima Y, Moribayashi A, Taylor D. 2007. Ecdysteroid hormone titer and its relationship to vitellogenesis in the soft tick, Ornithodoros moubata (Acari: Argasidae). Gen. Comp. Endocrinol. 150:371-80
105. Ohnishi E, Mizuno T, Chatani F, Ikekawa N, Sakurai S. 1977. 2-Deoxy-alpha-ecdysone from ovaries and eggs of the silkworm, Bombyx mori. Science 197:66-67
106. Ohnishi E, Mizuno T, Ikekawa N, Ikeda T. 1981. Accumulation of 2-deoxy-ecdysteroids in ovaries of the silkworm, Bombyx mori. Insect Biochem. 11:155-59
107. Ono H, Rewitz KF, Shinoda T, Itoyama K, Petryk A, et al. 2006. spook and spookier code for stage-specific components of the ecdysone biosynthetic pathway in Diptera. Dev. Biol. 298:555-70
108. Parker KL, Rice DA, Lala DS, Ikeda Y, Luo X, et al. 2002. Steroidogenic factor 1: an essential mediator of endocrine development. Recent Prog. Horm. Res. 57:19-36
109. Parker KL, Schimmmer BP. 1997. Steroidogenic factor 1: a key determinant of endocrine development and function. Endocr. Rev. 18:361-77
110. Parvy JP, Blais C, Bernard F, Warren JT, Petryk A, et al. 2005. A role for BFTZ-F1 in regulating ecdysteroid titers during postembryonic development in Drosophila melanogaster. Dev. Biol. 282:84-94
111. Petryk A, Warren JT, Marques G, Jarcho MP, Gilbert LI, et al. 2003. Shade is the Drosophila P450 enzyme that mediates the hydroxylation of ecdysone to the steroid insect molting hormone 20-hydroxyecdysone. Proc. Natl. Acad. Sci. USA 100:13773-78
112. Poretsky L, Cataldo N, Rosenwaks Z, Guidice L. 1999. The insulin-related ovarian regulatory system in health and disease. Endocr. Rev. 200:535-82
113. Pourie G, Trabalon M. 2003. The role of 20-hydroxyecdysone on the control of spider vitellogenesis. Gen. Comp. Endocrinol. 131:250-57
114. Raikhel AS, Brown MR, Belles X. 2005. Hormonal control of reproductive processes. See Ref. 36, 3:432-91
115. Redfern CPF. 1989. Ecdysiosynthetic tissues. See Ref. 77, pp. 182-87
116. Rees HH. 1985. Biosynthesis of ecdysone. See Ref. 73, 7:249-93
117. Rees HH. 1989. Pathways of biosynthesis of ecdysone. See Ref. 77, pp. 152-60
118. Rees HH. 1995. Ecdysteroid biosynthesis and inactivation in relation to function. Eur. J. Entomol. 92:9-39
119. Rees HH. 2003. Ecdysteroidogenic pathway. In Encyclopedia of Hormones, ed. HL Henry, AWNorman, pp. 460-66. San Diego: Academic
120. Rees HH. 2004. Hormonal control of tick development and reproduction. Parasitology 129:S127-S43
121. Rees HH, Isaac RE. 1984. Biosynthesis of ovarian ecdysteroid phosphates and their metabolic fate during embryogenesis in Schistocerca gregaria. In Biosynthesis, Metabolism and Mode of Action of Invertebrate Hormones, ed. JA Hoffmann, M Porchet, pp. 181-95. Berlin: Springer-Verlag
122. Rewitz KF, O'Connor MB, Gilbert LI. 2007. Molecular evolution of the insect Halloween family of cytochrome P450s: phylogeny, gene organization and functional conservation. Insect Biochem. Mol. Biol. 37:741-53
123. Rewitz KF, Rybczynski R, Warren JT, Gilbert LI. 2006. Identification, characterization and developmental expression of Halloween genes encoding P450 enzymes mediating ecdysone biosynthesis in the tobacco hornworm, Manduca sexta. Insect Biochem. Mol. Biol. 36:188-99
124. Riehle MA, Brown MR. 1999. Insulin stimulates ecdysteroid production through a conserved signaling cascade in the mosquito Aedes aegypti. Insect Biochem. Mol. Biol. 29:855-60
125. Riehle MA, Brown MR. 2002. Insulin receptor expression during development and a reproductive cycle in the ovary of the mosquito Aedes aegypti. Cell Tissue Res. 308:409-20
126. Riehle MA, Brown MR. 2003. Molecular analysis of the serine/threonine kinase Akt and its expression in the mosquito Aedes aegypti. Insect Mol. Biol. 12:225-32
127. Robinson-White A, Stratakis CA. 2002. Protein kinase A signaling: "cross talk" with other pathways in endocrine cells. Ann. N. Y. Acad. Sci. 968:256-70
128. Roth GE, Gierl MS, Vollborn L, Meise M, Lintermann R, Korge G. 2004. The Drosophila gene Start1: A putative cholesterol transporter and key regulator of ecdysteroid synthesis. Proc. Natl. Acad. Sci. USA 101:1601-6
129. Roy S, De J, Kundu S, Biswas A, Pramanik M, Ray AK. 2007. Estradiol-17beta: tracing its metabolic significance in female fat body of fifth instar larvae of silkworm, Bombyx mori L (race: Nistari). Life Sci. 80:446-53
130. Ruppert EE, Fox RS, Barnes RD. 2004. Invertebrate Zoology: A Functional Approach. Belmont, CA: Brooks/Cole-Thomson Learning. 963 pp.
131. Sakuri S. 2005. Feedback regulation of prothoracic gland activity. See Ref. 36, 3:409-31
132. Sall C, Tsoupras G, Kappler C, Lagueux M, Zachary D, et al. 1983. Fate of maternal conjugated ecdysteroids during embryonic development in Locusta migratoria. J. Insect Physiol. 29:491-507
133. Seifert G. 1990. Morphology, histology, and ultrastructure of ecdysial glands in Myriapoda. See Ref. 46, pp. 307-41
134. Seto-Young D, Zajac J, Liu H-C, Rosenwaks Z, Poretsky L. 2003. The role of mitogen-activated protein kinase in insulin and insulin-like growth factor I (IGF-I) signaling cascades for progesterone and IGFbinding protein-1 production in human granulosa cells. J. Clin. Endocrinol. Metab. 88:3385-91
135. Shapiro JP. 1983. Ovarian cyclic AMP and response to a brain hormone from the mosquito Aedes aegypti. Insect Biochem. 13:273-79
136. Sieglaff DH, Adams Duncan K, Brown MR. 2005. Expression of genes encoding proteins involved in ecdysteroidogenesis in the female mosquito, Aedes aegypti. Insect Biochem. Mol. Biol. 35:471-90
137. Slinger AJ, Isaac RE. 1988. Ecdysteroid titers during embryogenesis of the cockroach, Periplaneta americana. J. Insect Physiol. 34:1119-25
138. Smith SL. 1985. Regulation of ecdysteroid titer: synthesis. See Ref. 73, 7:295-341
139. Smith SL, Mitchell MJ. 1986. Ecdysone 20-monoxygenase systems in a larval and adult dipteran: an overview of their biochemistry, physiology and pharmacology. Insect Biochem. 16:49-55
140. Snyder MJ, Feyereisen R. 1993. A diazepam binding inhibitor (DBI) homolog from the tobacco hornworm, Manduca sexta. Mol. Cell. Endocrinol. 94:R1-4
141. Snyder MJ, Van Antwerpen R. 1998. Evidence for a diazepam-binding inhibitor (DBI) benzodiazepine receptor-like mechanism in ecdysteroidogenesis by the insect prothoracic gland. Cell Tissue Res. 294:161-68
142. Sonobe H, Ohira T, Ieki K, Maeda S, Ito Y, et al. 2006. Purification, kinetic characterization, and molecular cloning of a novel enzyme, ecdysteroid 22-kinase. J. Biol. Chem. 281:29513-24
143. Sonobe H, Yamada R. 2004. Ecdysteroids during early embryonic development in silkworm Bombyx mori: metabolism and functions. Zool. Sci. 21:503-16
144. Srivastava DP, Yu EJ, Kennedy K, Chatwin H, Reale V, et al. 2005. Rapid, nongenomic responses to ecdysteroids and catecholamines mediated by a novel Drosophila G-protein-coupled receptor. J. Neurosci. 25:6145-55
145. Subramoniam T. 2000. Crustacean ecdysteroids in reproduction and embryogenesis. J. Comp. Biochem. Physiol. C 125:135-56
146. Suzuki S, Yamasaki K, Fujita T, Mamiya Y, Sonobe H. 1996. Ovarian and hemolymph ecdysteroids in the terrestrial isopod Armadillidium vulgare (Malacostracan Crustacea). Gen. Comp. Endocrinol. 104:129-38
147. Svoboda JA, Feldlaufer MF. 1991. Neutral sterol metabolism in insects. Lipids 26:614-18
148. Swevers L, Lambert JG, De Loof A. 1991. Synthesis and metabolism of vertebrate-type steroids by tissues of insects: a critical evaluation. Experientia 47:687-98
149. Takeuchi H, Chen JH, Jenkins JR, Bun-Ya M, Turner PC, Rees HH. 2004. Characterization of a sterol carrier protein 2/3-oxoacyl-CoA thiolase from the cotton leafworm (Spodoptera littoralis): a lepidopteran mechanism closer to that in mammals than that in dipterans. Biochem. J. 382:93-100
150. Tatar M, Kopelman A, Epstein D, Tu M-P, Yin C-M, Garofalo RS. 2001. A mutant Drosophila insulin receptor homolog that extends life-span and impairs neuroendocrine function. Science 292:107-10
151. Telford MJ, Bourlat SJ, Economou A, Papillon D, Rota-Stabelli O. 2008. The evolution of the Ecdysozoa. Philos. Trans. R. Soc. London Ser. B 363:1529-37
152. Thompson MJ, Weirich GF, Svoboda JA. 1990. Metabolism of insect molting hormones: bioconversion and titer regulation. See Ref. 46, pp. 326-60
153. Tu M-P, Yin C-M, Tatar M. 2002. Impaired ovarian ecdysone synthesis of Drosophila melanogaster insulin receptor mutants. Aging Cell 1:158-60
154. Vafopoulou X, Steel CGH. 2005. Testis ecdysiotropic peptides in Rhodnius prolixus: biological activity and distribution in the nervous system and testis. J. Insect Physiol. 51:1227
155. Warren JT, Bachmann JS, Dai JD, Gilbert LI. 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 l(3)ecd1 mutation. Insect Biochem. Mol. Biol. 26:931-43
156. Warren JT, Gilbert LI. 1996. Metabolism in vitro of cholesterol and 25-hydroxycholesterol by the larval prothoracic glands of Manduca sexta. Insect Biochem. Mol. Biol. 26:917-29
157. Warren JT, Petryk A, Marques G, Jarcho M, Parvy JP, et al. 2002. Molecular and biochemical characterization of two P450 enzymes in the ecdysteroidogenic pathway of Drosophila melanogaster. Proc. Natl. Acad. Sci. USA 99:11043-48
158. Warren JT, Petryk A, Marques G, Parvy JP, Shinoda T, et al. 2004. Phantom encodes the 25-hydroxylase of Drosophila melanogaster and Bombyx mori: a P450 enzyme critical in ecdysone biosynthesis. Insect Biochem. Mol. Biol. 34:991-1010
159. Watanabe K, Hull JJ, Niimi T, Imai K, Matsumoto S, et al. 2007. FXPRL-amide peptides induce ecdysteroidogenesis through a G-protein coupled receptor expressed in the prothoracic gland of Bombyx mori. Mol. Cell. Endocrinol. 273:51-58
160. Watson CS, Alyea RA, Jeng YJ, Kochukov MY. 2007. Nongenomic actions of low concentration estrogens and xenoestrogens on multiple tissues. Mol. Cell. Endocrinol. 274:1-7
161. Wilder MN, Okumura T, Aida K. 1991. Accumulation of ovarian ecdysteroids in synchronization with gonadal development in the giant freshwater prawn, Macrobrachium rosenbergii. Zool. Sci. 8:919-27
162. Wood JR, Strauss JF III. 2002. Multiple signal transduction pathways regulate ovarian steroidogenesis. Rev. Endocr. Metab. Disord. 3:33-46
163. Yamada R, Sonobe H. 2003. Purification, kinetic characterization and molecular cloning of a novel enzyme ecdysteroid-phosphate phosphatase. J. Biol. Chem. 278:26365-73
164. Yamanaka N, Hua Y-J, Mizoguchi A, Watanabe K, Niwa R, et al. 2005. Identification of a novel prothoracicostatic hormone and its receptor in the silkworm Bombyx mori. J. Biol. Chem. 280;14684-90
165. Yoshiyama T, Namiki T, Mita K, Kataoka H, Niwa R. 2006. Neverland is an evolutionally conserved Rieske-domain protein that is essential for ecdysone synthesis and insect growth. Development 133:2565-74
166. Zhu XX, Gfeller H, Lanzrein B. 1983. Ecdysteroids during oogenesis in the oviviparous cockroach Nauphoeta cinerea. J. Insect Physiol. 29:225-35