Cited by 532Neurobiology and Insect Physiology ResearchSteroid Chemistry and BiochemistryLipid metabolism and biosynthesis

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Control and Biochemical Nature of the Ecdysteroidogenic Pathway

Gilbert, Lawrence I., Rybczynski, Robert, Warren, James T.·2002·Annual Review of Entomology

DOI 10.1146/annurev.ento.47.091201.145302

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Abstract

Molting is elicited by a critical titer of ecdysteroids that includes the principal molting hormone, 20-hydroxyecdysone (20E), and ecdysone (E), which is the precursor of 20E but also has morphogenetic roles of its own. The prothoracic glands are the predominate source of ecdysteroids, and the rate of synthesis of these polyhydroxylated sterols is critical for molting and metamorphosis. This review concerns three aspects of ecdysteroidogenesis: (a) how the brain neuropeptide prothoracicotropic hormone (PTTH) initiates a transductory cascade in cells of the prothoracic gland, which results in an increased rate of ecdysteroid biosynthesis (upregulation); (b) how the concentrations of 20E in the hemolymph feed back on the prothoracic gland to decrease rates of ecdysteroidogenesis (downregulation); and (c) how the prothoracic gland cells convert cholesterol to the precursor of E and then 20E, a series of reactions only now being understood because of the use of a combination of classical biochemistry and molecular genetics.

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Ecdysteroid titers and expression of Halloween genes and ecdysteroid receptor in relation to overwintering and the long larval phase in the seabuckthorn carpenterworm, H olcocerus hippophaecolus
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“The active form of ecdysone, 20-hydroxyecdysone (20E), is the master regulator of insect development and reproduction (Gilbert et al., 2002; Gilbert & Warren, 2005).”
Background
Expression of β-adrenergic-like Octopamine Receptors duringDrosophilaDevelopment
Ohhara, Y., Kayashima, Y., Hayashi, Y., Kobayashi, S., & Yamakawa‐Kobayashi, K. (2012) · ZOOLOGICAL SCIENCE · doi:10.2108/zsj.29.83 · Cited by 22
“5N, O) because the ring gland contains the prothoracic gland (PG), which is known to be a site of ecdysteroid production (Gilbert et al., 2002; Marchal et al., 2010), and because octopamine facilitates the production of ecdysteroid in vitro cultures of Bombyx PG (Hirashima et al.”
Background
Biosynthesis of Triterpenoid Natural Products
Hu, D., Gao, H., & Yao, X. (2020) · Elsevier eBooks · doi:10.1016/b978-0-12-409547-2.14678-5 · Cited by 22
Venom gland components of the ectoparasitoid wasp, Anisopteromalus calandrae.
Perkin, L., Friesen, K. S., Flinn, P. W., & Oppert, B. (2015) · PubMed Central · Cited by 22
RNA interference‐mediated knockdown of the Halloween gene Spookiest (CYP307B1) impedes adult eclosion in the western tarnished plant bug, Lygus hesperus
Ekert, E. V., Wang, M., Miao, Y., Brent, C. S., & Hull, J. J. (2016) · Insect Molecular Biology · doi:10.1111/imb.12242 · Cited by 21
“…ecdysteroid biosynthesis must rely on dietary supplied cholesterol (obtained directly from prey by carnivorous insects) or plant-derived phytosterols, which must first undergo dealkylation to cholesterol in midgut cells (Gilbert et al., 2002; Gilbert & Warren, 2005; Canavoso et al., 2001).”
BackgroundInfluential
Knockdown of the Halloween Genes spook, shadow and shade Influences Oocyte Development, Egg Shape, Oviposition and Hatching in the Desert Locust
Schellens, S., Lenaerts, C., Baca, M. D. R. P., Cools, D., Peeters, P., Marchal, E., & Broeck, J. V. (2022) · International Journal of Molecular Sciences · doi:10.3390/ijms23169232 · Cited by 23
“Already in the intestine of the insects, plant-derived phytosterols will be dealkylated to cholesterol [2].”
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Calcium and cAMP directly modulate the speed of the Drosophila circadian clock
Palacios-Muñoz, A., & Ewer, J. (2018) · PLoS Genetics · doi:10.1371/journal.pgen.1007433 · Cited by 21
Incomplete sterols and hopanoids pathways in ciliates: Gene loss and acquisition during evolution as a source of biosynthetic genes
Tomazic, M. L., Poklépovich, T., Nudel, C. B., & Nusblat, A. D. (2014) · Molecular Phylogenetics and Evolution · doi:10.1016/j.ympev.2014.01.026 · Cited by 21
“, 2008), (ii) serve as precursors of bile salts and a number of different steroid hormones implicated in growth, cell cycle and development in mammals, of brassinosteroids in plants (Benveniste, 2004), of antheridiol and oogoniol in aquatic fungi (Brunt and Silver, 1991), of ecdysteroids in arthropods (Gilbert et al.,…”
Background
Coordinating morphology with behavior during development: an integrative approach from a fly perspective
Carvalho, M., & Mirth, C. K. (2015) · Frontiers in Ecology and Evolution · doi:10.3389/fevo.2015.00005 · Cited by 21
Nitric oxide directly regulates gene expression during Drosophila development: need some gas to drive into metamorphosis?: Figure 1.Preprint
Yamanaka, N., & O’Connor, M. B. (2011) · Genes & Development · doi:10.1101/gad.2080411 · Cited by 21
“Ecdysone biosynthesis (ecdysteroidogenesis) in the PG is mediated by multiple enzymes, each of which is engaged in a specific step of sequential hydroxylation and oxidation reactions (Gilbert et al. 2002).”
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The Integrative Physiology of Hormone Signaling: Insights from Insect Models
Koyama, T., Saeed, U., Rewitz, K., & Halberg, K. A. (2025) · Physiology · doi:10.1152/physiol.00030.2024 · Cited by 22
Sterol metabolism in the filasterean Capsaspora owczarzaki has features that resemble both fungi and animals
Najle, S. R., Molina, M. C., Ruiz‐Trillo, I., & Uttaro, A. D. (2016) · Open Biology · doi:10.1098/rsob.160029 · Cited by 20
“As signalling, sterols act directly as precursors of diverse hormones and bioactive metabolites, including steroids, bile salts and vitamin D in vertebrates [1], ecdysteroids in arthropods [2], dafachronic acids in nematodes [3], brassinosteroids in plants, and antheridiol and oogoniol in fungi [4].”
Effect of Dietary Cholesterol on Growth Performance, Cholesterol Deposition, and Lipid Metabolism in Adult Chinese Mitten Crab (Eriocheir sinensis)
Guo, H., Jiang, G., Dai, Y., Huang, Y., Wang, X., Xiao, K., Wang, M.-M., Abasubong, K. P., Xu, Y., Zhang, D.-D., et al. (2022) · Aquaculture Nutrition · doi:10.1155/2022/2012958 · Cited by 20
“The deficiency of CHO in the feed may be one of the reasons for these problems [8, 24].”
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Growth inhibition and differences in protein profiles in azadirachtin‐treated Drosophila melanogaster larvae
Wang, H., Lai, D., Yuan, M., & Xu, H. (2014) · Electrophoresis · doi:10.1002/elps.201300318 · Cited by 20
Changes in ecdysteroid levels and expression patterns of ecdysteroid-responsive factors and neuropeptide hormones during the embryogenesis of the blue crab, Callinectes sapidus
Techa, S., Alvarez, J. V., & Chung, J. S. (2014) · General and Comparative Endocrinology · doi:10.1016/j.ygcen.2014.07.017 · Cited by 20
“Upregulation of EcR and RXR by ecdysteroids that occurs in prothoracic gland of M. sexta further modulates this gland’s activity for ecdysteroid synthsis (Gilbert et al., 2002, 1997; Song and Gilbert, 1998).”
Background
The receptor tyrosine kinase torso regulates ecdysone homeostasis to control developmental timing in Bombyx mori
Zhang, Z., Liu, X., Yu, Y., Fangying, Y., & Li, K. (2020) · Insect Science · doi:10.1111/1744-7917.12879 · Cited by 20
“Steroid hormones are the primary endocrine molecules that regulate the developmental transition (Gilbert et al., 2002).”
Background
Transcriptome of a specialized extra‐embryonic cell, teratocyte, and its host immunosuppressive role revealed by ex vivo RNA interference
Ali, M. R., Lim, J., & Kim, Y. (2014) · Insect Molecular Biology · doi:10.1111/imb.12132 · Cited by 20
Evolution of the thyroid hormone, retinoic acid, ecdysone and liver X receptors
Ollikainen, N., Chandsawangbhuwana, C., & Baker, M. E. (2006) · Integrative and Comparative Biology · doi:10.1093/icb/icl035 · Cited by 20
Temporal changes in DNA synthesis of prothoracic gland cells during larval development and their correlation with ecdysteroidogenic activity in the silkworm, Bombyx mori
Gu, S., & Chow, Y. (2005) · Journal of Experimental Zoology Part A Comparative Experimental Biology · doi:10.1002/jez.a.142 · Cited by 20
Induction of diapause by clock proteins period and timeless via changes in PTTH and ecdysteroid titer in the sugar beet moth, Scrobipalpa ocellatella (Lepidoptera: Gelechiidae)
Ahmadi, F., Mikani, A., & Moharramipour, S. (2021) · Archives of Insect Biochemistry and Physiology · doi:10.1002/arch.21790 · Cited by 19
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“This in turn leads to the secretion of α-ecdysone from the prothoracic gland (PG) (Warren et al., 2002; Gilbert et al., 2002; Gilbert, 2004).”
Background
Fackel interacts with gibberellic acid signaling and vernalization to mediate flowering in Arabidopsis
Huang, B., Qian, P., Gao, N., Shen, J., & Hou, S. (2017) · Planta · doi:10.1007/s00425-017-2652-5 · Cited by 19
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Sandlund, L., Kongshaug, H., Horsberg, T. E., Male, R., Nilsen, F., & Dalvin, S. T. (2018) · PLoS ONE · doi:10.1371/journal.pone.0191995 · Cited by 19
A comprehensive study of the changes in ecdysteroid levels during the feeding phase of fifth instar larvae of the silkworm, Bombyx mori (Lepidoptera: Bombycidae)
Keshan, B., Thounaojam, B., & Kh., S. D. (2015) · European Journal of Entomology · doi:10.14411/eje.2015.088 · Cited by 19
Identification and expression patterns of Halloween genes encoding cytochrome P450s involved in ecdysteroid biosynthesis in the cotton bollwormHelicoverpa armigera
Zheng, J., Tian, K., Yuan, Y., Li, M., & Qiu, X. (2016) · Bulletin of Entomological Research · doi:10.1017/s0007485316000663 · Cited by 19
“…that insects synthesize ecdysone in the prothoracic gland (PG) fromdietary cholesterol or phytosterol, and the synthesized ecdysone (E) is secreted in the hemolymph, and then converted into 20E in various peripheral tissues predominantly in midgut (MG) or fat body (FB) (Gilbert et al., 2002).”
Background
EcR expression in the prothoracicotropic hormone‐producing neurosecretory cells of the Bombyx mori brain
Hossain, M., Shimizu, S., Fujiwara, H., Sakurai, S., & Iwami, M. (2006) · FEBS Journal · doi:10.1111/j.1742-4658.2006.05398.x · Cited by 19
Bombyx mori prothoracicostatic peptide inhibits ecdysteroidogenesis in vivo
Liu, X., Tanaka, Y., Song, Q., Xu, B., & Hua, Y. (2004) · Archives of Insect Biochemistry and Physiology · doi:10.1002/arch.20005 · Cited by 19
Sex and stripping: The key to the intimate relationship between Wolbachia and host?
Negri, I., Pellecchia, M., Grève, P., Daffonchio, D., Bandi, C., & Alma, A. (2010) · Communicative & Integrative Biology · doi:10.4161/cib.3.2.10520 · Cited by 18
Effect of elevated CO2 on interactions between the host plant Phaseolus vulgaris and the invasive western flower thrips, Frankliniella occidentalis
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The polyembryonic wasp Copidosoma floridanum produces two castes by differentially parceling the germ line to daughter embryos during embryo proliferation
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“Like many Lepidoptera, growth and molting of the host larva (T. ni) is a highly synchronized process regulated by the interactions between nutrition and hormones (Nijhout 2003; Baehrecke et al. 1993; Gilbert et al. 2002; Corley et al. 2005).”
Background
Identification of Neuropeptides and Their Receptors in the Ectoparasitoid, Habrobracon hebetor
Yu, K., Xiong, S., Xu, G., Ye, X., Yao, H., Wang, F., Fang, Q., Song, Q., & Ye, G. (2020) · Frontiers in Physiology · doi:10.3389/fphys.2020.575655 · Cited by 18
“PTTH is secreted by the brain’s neurosecretory cells and released into hemolymph at a specific time and in particular developmental stages (Yamanaka et al., 2010).”
BackgroundInfluential
Nuclear receptors linking physiology and germline stem cells in Drosophila
Finger, D. S., Whitehead, K. M., Phipps, D. N., & Ables, E. T. (2021) · Vitamins and hormones · doi:10.1016/bs.vh.2020.12.008 · Cited by 17
Neuropeptidergic control of neurosteroids biosynthesis
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“Ecdysteroids are the insect steroid hormones that are synthesized and secreted by the prothoracic glands upon stimulation by the brain neurohormone, prothoracicotropic hormone (PTTH) [1–3].”
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Sterol Characteristics in Silkworm Brain and Various Tissues Characterized by Precise Sterol Profiling Using LC-MS/MS
Takeshima, M., Ogihara, M. H., & Kataoka, H. (2019) · International Journal of Molecular Sciences · doi:10.3390/ijms20194840 · Cited by 17
“In animals, the main sterol is cholesterol (Chl), which is a necessary component of cell membranes [1], a material of steroid hormones [2], and a ligand for the Hedgehog signaling process [3,4].”
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Roles of and cross-talk between ecdysteroid and sesquiterpenoid pathways in embryogenesis of branchiopod crustacean Daphnia magna
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An ecdysone receptor from the pentatomomorphan, Nezara viridula, shows similar affinities for moulting hormones makisterone A and 20-hydroxyecdysone
Tohidi‐Esfahani, D., Graham, L. D., Hannan, G. N., Simpson, A. M., & Hill, R. J. (2010) · Insect Biochemistry and Molecular Biology · doi:10.1016/j.ibmb.2010.10.002 · Cited by 17
Characterization of candidate intermediates in the Black Box of the ecdysone biosynthetic pathway in Drosophila melanogaster: Evaluation of molting activities on ecdysteroid-defective larvae
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Authors: Gilbert, L. I., Rybczynski, R., & Warren, J. T.
Affiliations: University of North Carolina at Chapel Hill (US)
Venue: Annual Review of Entomology
Publisher: Annual Reviews
Published: 2002-01
Volume: 47
Issue: 1
Pages: 883-916
ISSN: 0066-4170 (ISSN-L)
1545-4487
DOI: 10.1146/annurev.ento.47.091201.145302
OpenAlex: W2157228103
Cited by: 532
References: 185
Citations by year: 2026: 15 · 2025: 20 · 2024: 16 · 2023: 14 · 2022: 14 · 2021: 21 · 2020: 32 · 2019: 20 · 2018: 11 · 2017: 13 · 2016: 20 · 2015: 21 · 2014: 31 · 2013: 28 · 2012: 26
FWCI: 2.40
Type: journal article
Language: en
Open access: closed
Locations: Annual Review of Entomology (publishedVersion)
PubMed (publishedVersion)
Topics: Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neurobiology and Insect Physiology Research
Physical Sciences › Chemistry › Organic Chemistry › Steroid Chemistry and Biochemistry
Life Sciences › Biochemistry, Genetics and Molecular Biology › Biochemistry › Lipid metabolism and biosynthesis
Keywords: ecdysteroid biosynthesis, 20-hydroxyecdysone, ecdysone, prothoracic gland, prothoracicotropic hormone, molting, metamorphosis, cholesterol conversion, steroidogenesis, insect endocrinology
MeSH terms: Animals, Insecta, Down-Regulation, Ecdysteroids

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Control and Biochemical Nature of the Ecdysteroidogenic Pathway

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