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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“E-mail address: s.g.webster@bangor.ac.uk (S.G. Webster). of prothoracicotropic hormone (PTTH)1 in stimulating ecdysteroid secretion by the prothoracic glands (reviews by Gilbert et al., 1996, 2002), a series of neuropeptide hormones are involved in the insect ecdysis programme.”
Background
PTTH-stimulated ERK phosphorylation in prothoracic glands of the silkworm, Bombyx mori: Role of Ca2+/calmodulin and receptor tyrosine kinase
Gu, S., Lin, J., & Lin, P. (2009) · Journal of Insect Physiology · doi:10.1016/j.jinsphys.2009.09.008 · Cited by 49
Drosophila is an inclusive model for human diseases, growth and development
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The sterol carrier protein 2/3-oxoacyl-CoA thiolase (SCPx) is involved in cholesterol uptake in the midgut of Spodoptera litura: gene cloning, expression, localization and functional analyses
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“Insects utilize phytols, such as β-sitosterol, campesterol and stigmasterol, and synthesize ecdysteroids (molting hormone) in the prothoracic glands [3].”
BackgroundInfluential
Juvenile hormone synthesis, metabolism, and resulting haemolymph titre in Heliothis virescens larvae parasitized by Toxoneuron nigriceps
Li, S., Falabella, P., Kuriachan, I., Vinson, S. B., Borst, D. W., Malva, C., & Pennacchio, F. (2003) · Journal of Insect Physiology · doi:10.1016/s0022-1910(03)00185-9 · Cited by 49
Body-Size Regulation: Combining Genetics and Physiology
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Protein tyrosine phosphatases of Toxoneuron nigriceps bracovirus as potential disrupters of host prothoracic gland function
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A cuticle protein gene in the honeybee: Expression during development and in relation to the ecdysteroid titer
Soares, M. P. M., Elias-Neto, M., Simões, Z. L. P., & Bitondi, M. M. G. (2007) · Insect Biochemistry and Molecular Biology · doi:10.1016/j.ibmb.2007.07.014 · Cited by 48
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Gilboa, L. (2015) · Current Opinion in Genetics & Development · doi:10.1016/j.gde.2015.01.005 · Cited by 50
“Current Opinion in Genetics & Development 2015, 32:31–36 prothoracic gland [34].”
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Microarray and RNAi Analysis of P450s in Anopheles gambiae Male and Female Steroidogenic Tissues: CYP307A1 Is Required for Ecdysteroid Synthesis
Pondeville, E., David, J., Crilat, E., Maria, A., Jacques, J., Ranson, H., Bourgouin, C., & Dauphin‐Villemant, C. (2013) · PLoS ONE · doi:10.1371/journal.pone.0079861 · Cited by 47
“The black box, and more particularly the oxidative steps, is thought to involve one or more P450 enzymes that still remain uncharacterized [6,18].”
Background
Prothoracicotropic hormone stimulated extracellular signal-regulated kinase (ERK) activity: the changing roles of Ca2+- and cAMP-dependent mechanisms in the insect prothoracic glands during metamorphosis
Rybczynski, R., & Gilbert, L. I. (2003) · Molecular and Cellular Endocrinology · doi:10.1016/s0303-7207(03)00090-x · Cited by 47
Balancing energy expenditure and storage with growth and biosynthesis during Drosophila development
Gillette, C. M., Tennessen, J. M., & Reis, T. (2021) · Developmental Biology · doi:10.1016/j.ydbio.2021.01.019 · Cited by 43
Interaction of proteins involved in ecdysone and juvenile hormone signal transduction
Bitra, K., & Palli, S. R. (2008) · Archives of Insect Biochemistry and Physiology · doi:10.1002/arch.20281 · Cited by 46
Neuropeptide signaling mechanisms in crustacean and insect molting glands
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“The divergence in amino acid sequences, particularly those in polar regions located at the surface of the native protein, is probably the basis for PTTH specificity, as a PTTH from one species has no effect on the PG of another species (see Gilbert et al. 2002 for review).”
BackgroundInfluential
Differential Regulation of Ecdysteroidogenic P450 Gene Expression in the Silkworm,Bombyx mori
Yamanaka, N., Honda, N., OSATO, N., Niwa, R., Mizoguchi, A., & Kataoka, H. (2007) · Bioscience Biotechnology and Biochemistry · doi:10.1271/bbb.70420 · Cited by 48
CDK8-Cyclin C Mediates Nutritional Regulation of Developmental Transitions through the Ecdysone Receptor in Drosophila
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Simultaneous quantification of individual intermediate steroids in silkworm ecdysone biosynthesis by liquid chromatography–tandem mass spectrometry with multiple reaction monitoring
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Morphological, Molecular, and Hormonal Basis of Limb Regeneration across Pancrustacea: Table 1
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“The prothoracic gland in insects produces ecdysone from cholesterol via a series of biochemical reactions and ecdysone is converted into 20E in the peripheral tissues (Gilbert et al. 2002).”
Background
Plant phloem sterol content: forms, putative functions, and implications for phloem-feeding insects
Behmer, S. T., Olszewski, N., Sebastiani, J., Palka, S., Sparacino, G., Sciarrno, E., & Grebenok, R. J. (2013) · Frontiers in Plant Science · doi:10.3389/fpls.2013.00370 · Cited by 43
“Insects require very small pools of cholesterol as the precursor to the steroid molting hormones (Gilbert et al., 2002), and in plants cholesterol is a close analog to 24-methylene cholesterol (Benveniste, 2002), which is an intermediate in the synthesis of campesterol, the direct precursor to…”
Background
The effect of silencing 20E biosynthesis relative genes by feeding bacterially expressed dsRNA on the larval development of Chilo suppressalis
Zhu, J. K., Dong, Y., Li, P., & Niu, C. (2016) · Scientific Reports · doi:10.1038/srep28697 · Cited by 43
Bombyx E75 isoforms display stage- and tissue-specific responses to 20-hydroxyecdysone
Li, K., Guo, E., Hossain, M. S., Li, Q., Cao, Y., Tian, L., Deng, X., & Li, S. (2015) · Scientific Reports · doi:10.1038/srep12114 · Cited by 45
Novel DNA-binding element within the C-terminal extension of the nuclear receptor DNA-binding domain
Jakób, M., Kolodziejczyk, R., Orłowski, M., Krzywda, S., Kowalska, A., Dutko-Gwóźdź, J., Gwóźdź, T., Kochman, M., Jaskólski, M., & Ożyhar, A. (2007) · Nucleic Acids Research · doi:10.1093/nar/gkm162 · Cited by 43
“Although 20E is thought to elicit most of the above-mentioned physiological responses, mounting evidence indicates, that alternate signaling pathways exist that are driven by ecdysteroids other than 20E, some of which are present at specific stages during development (13).”
Background
Genome-wide identification of whole ATP-binding cassette (ABC) transporters in the intertidal copepod Tigriopus japonicus
Jeong, C., Kim, B., Lee, J., & Rhee, J. (2014) · BMC Genomics · doi:10.1186/1471-2164-15-651 · Cited by 42
“japonicus, as ecdysone has a conserved role in crustaceans [53].”
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Wolbachia as an “Infectious” Extrinsic Factor Manipulating Host Signaling Pathways
Negri, I. (2012) · Frontiers in Endocrinology · doi:10.3389/fendo.2011.00115 · Cited by 42
“Then the EcR/USP complex activates the transcriptional processes underlying the cellular and morphogenetic molting cascade events (Gilbert et al., 2002).”
Endocrine interactions between plants and animals: Implications of exogenous hormone sources for the evolution of hormone signaling
Miller, A. E. M., & Heyland, A. (2009) · General and Comparative Endocrinology · doi:10.1016/j.ygcen.2009.09.016 · Cited by 42
“In adults it is synthesized in the ovaries, male testes and accessory glands, and possibly other tissues as well (Gilbert et al., 2002; Pondeville et al., 2008).”
Ecdysteroid-Dependent Expression of the Tweedle and Peroxidase Genes during Adult Cuticle Formation in the Honey Bee, Apis mellifera
Soares, M. P. M., Silva-Torres, F. A., Elias-Neto, M., Nunes, F. M. F., Simões, Z. L. P., & Bitondi, M. M. G. (2011) · PLoS ONE · doi:10.1371/journal.pone.0020513 · Cited by 42
“The coordination of cuticle renewal is known to be controlled by the ecdysteroid hormone 20-hydroxyecdysone [17,18,19]; its precursor, ecdysone, and other structurally related compounds are only active as molting hormones in a few organisms [20].”
Background
Studies on the black box: Incorporation of 3-oxo-7-dehydrocholesterol into ecdysteroids by Drosophila melanogaster and Manduca sexta
Warren, J. T., O’Connor, M. B., & Gilbert, L. I. (2009) · Insect Biochemistry and Molecular Biology · doi:10.1016/j.ibmb.2009.08.004 · Cited by 40
“sexta prothoracic glands, the irradiation of in vitro-loaded PSKB-ketal isomers increased ecdysteroid production 6-fold, almost as if the glands had been stimulated by the addition of PTTH (Gilbert et al., 2002).”
BackgroundInfluential
Sterols are required for cell‐fate commitment and maintenance of the stomatal lineage in A rabidopsis
Qian, P., Han, B., Forestier, E. C. F., Hu, Z., Gao, N., Lu, W., Schaller, H., Li, J., & Hou, S. (2013) · The Plant Journal · doi:10.1111/tpj.12190 · Cited by 40
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The Putzig-NURF Nucleosome Remodeling Complex Is Required for Ecdysone Receptor Signaling and Innate Immunity in Drosophila melanogaster
Kugler, S. J., Gehring, E., Wallkamm, V., Krüger, V., & Nagel, A. C. (2011) · Genetics · doi:10.1534/genetics.111.127795 · Cited by 40
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Cholesterol derivatives induce dephosphorylation of the histone deacetylases Rpd3/HDAC1 to upregulate autophagy
Wu, W., Luo, M., Li, K., Dai, Y., Yi, H., Zhong, Y., Cao, Y., Tettamanti, G., & Tian, L. (2020) · Autophagy · doi:10.1080/15548627.2020.1725376 · Cited by 40
Two independent duplications forming the Cyp307a genes in Drosophila
Sztal, T. E., Chung, H., Gramzow, L., Daborn, P. J., Batterham, P., & Robin, C. (2007) · Insect Biochemistry and Molecular Biology · doi:10.1016/j.ibmb.2007.05.017 · Cited by 40
Protein kinase C modulates ecdysteroidogenesis in the prothoracic gland of the tobacco hornworm, Manduca sexta
Rybczynski, R., & Gilbert, L. I. (2006) · Molecular and Cellular Endocrinology · doi:10.1016/j.mce.2006.02.015 · Cited by 40
Ecdysone differentially regulates metamorphic timing relative to 20-hydroxyecdysone by antagonizing juvenile hormone in Drosophila melanogaster
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“A polyhydroxylated steroid, 20-hydroxyecdysone (20E), has been characterized as the principal molting hormone in insects (Gilbert et al., 2002).”
Finishing the egg
Berg, C. A., Sieber, M., & Sun, J. (2023) · Genetics · doi:10.1093/genetics/iyad183 · Cited by 41
Regulation of the C. elegans molt by pqn-47
Russel, S., Frand, A. R., & Ruvkun, G. (2011) · Developmental Biology · doi:10.1016/j.ydbio.2011.09.025 · Cited by 39
“where physiological and environmental cues that influence the period between molts are integrated (Gilbert et al., 2002).”
Background
microRNA-14 as an efficient suppressor to switch off ecdysone production after ecdysis in insects
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RNA interference‐mediated knockdown of the transcription factor Krüppel homologue 1 suppresses vitellogenesis in Chilo suppressalis
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“JH and 20E are secreted and synthesized by the corpora allata (CA) and prothoracic gland (PG), respectively (Röller et al., 1967; Gilbert et al., 2002).”
Background
Olfactory perception of herbicide butachlor by GOBP2 elicits ecdysone biosynthesis and detoxification enzyme responsible for chlorpyrifos tolerance in Spodoptera litura
Sun, Z., Wang, R., Du, Y., Gao, B., Gui, F., & Lǚ, K. (2021) · Environmental Pollution · doi:10.1016/j.envpol.2021.117409 · Cited by 38
Regulation of prothoracic gland ecdysteroidogenic activity leading to pupal metamorphosis
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“Although the molecular mechanism of feedback inhibition remain to be seen, it may be mediated by the heterodimeric nuclear receptor composed of the ecdysone receptor (EcR) and Ultraspiracle (USP) proteins (Gilbert et al., 2002).”
BackgroundMethod
Proteomic and Bioinformatic Analysis on Endocrine Organs of Domesticated Silkworm, Bombyx mori L. for a Comprehensive Understanding of Their Roles and Relations
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The POU Factor Ventral Veins Lacking/Drifter Directs the Timing of Metamorphosis through Ecdysteroid and Juvenile Hormone Signaling
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“enzymes necessary for catalyzing a series of reactions that ultimately convert cholesterol into E 115 [26,33-35].”
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Amino Acids and TOR Signaling Promote Prothoracic Gland Growth and the Initiation of Larval Molts in the Tobacco Hornworm Manduca sexta
Kemirembe, K., Liebmann, K., Bootes, A. M., Smith, W. A., & Suzuki, Y. (2012) · PLoS ONE · doi:10.1371/journal.pone.0044429 · Cited by 37
“In the hemolymph, 2-dehydroecdysone and 3-dehydroecdysone are rapidly converted to ecdysone, which is subsequently converted to 20E, a more active form of ecdysone that initiates a series of events associated with molting and eventual ecdysis of the larva [8,9,13,14].”
Background
BmStart1, a novel carotenoid-binding protein isoform from Bombyx mori, is orthologous to MLN64, a mammalian cholesterol transporter
Sakudoh, T., Tsuchida, K., & Kataoka, H. (2005) · Biochemical and Biophysical Research Communications · doi:10.1016/j.bbrc.2005.08.241 · Cited by 37
Analysis of ecdysteroidogenic activity of the prothoracic glands during the last larval instar of the silkworm,Bombyx mori
Gu, S., & Chow, Y. (2004) · Archives of Insect Biochemistry and Physiology · doi:10.1002/arch.20029 · Cited by 37
Endocrine and enzymatic shifts during insect diapause: a review of regulatory mechanisms
Izadi, H. (2025) · Frontiers in Physiology · doi:10.3389/fphys.2025.1544198 · Cited by 36
“These genes code for cytochrome P 450 enzymes in the ecdysteroidogenic pathway, which is responsible for the biosynthesis of ecdysone from cholesterol (Gilbert et al., 2002; Gu et al., 2021).”

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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