Phytoecdysteroids: Distribution, Structural Diversity, Biosynthesis, Activity, and Crosstalk with Phytohormones

In the paper

“A total of 207 new PEs have been identified from 17 plant families: Amaranthaceae, Asteraceae, Blechnaceae, Caryophyllaceae, Commelinaceae, Dioscoreaceae, Gleicheniaceae, Lamiaceae, Liliaceae, Limnanthaceae, Lygodiaceae, Malvaceae, Menispermaceae, Polypodiaceae, Polyporaceae, Rhodomelaceace, and Taxaceae (Table 1).”

In the paper

“Additionally, PEs are found in algae, fungi, ferns, gymnosperms, and angiosperms.”

What the sources say

• Not checked yetcites [4] Smagghe, G. (2009). Phytoecdysteroids: Diversity, biosynthesis and distribution. Ecdysone: Structures and Functions, Springer. source
• Not checked yetcites [7] Yusupova, U.Y., Ramazonov, N.S., Syrov, V.N., and Sagdullaev, S.S. (2022). Phytoecdysteroids. Phytoecdysteroids: Properties, Biological Activity and Applications, Springer. source
• Not checked yetcites [18] Strnad, 2016, Plant ecdysteroids: Plant sterols with intriguing distributions, biological effects and relations to plant hormones, Planta source
• Not checked yetcites [23] Adler, 1995, Biosynthesis and distribution of insect-molting hormones in plants—A review, Lipids source
• Not checked yetcites [24] Adler, 1999, Occurrence, biosynthesis, and putative role of ecdysteroids in plants, Crit. Rev. Biochem. Mol. source

In the paper

“More than 500 different PEs are found in over 100 terrestrial plants.”

What the sources say

• Not checked yetcites [4] Smagghe, G. (2009). Phytoecdysteroids: Diversity, biosynthesis and distribution. Ecdysone: Structures and Functions, Springer. source
• Not checked yetcites [7] Yusupova, U.Y., Ramazonov, N.S., Syrov, V.N., and Sagdullaev, S.S. (2022). Phytoecdysteroids. Phytoecdysteroids: Properties, Biological Activity and Applications, Springer. source
• Not checked yetcites [18] Strnad, 2016, Plant ecdysteroids: Plant sterols with intriguing distributions, biological effects and relations to plant hormones, Planta source
• Not checked yetcites [23] Adler, 1995, Biosynthesis and distribution of insect-molting hormones in plants—A review, Lipids source
• Not checked yetcites [24] Adler, 1999, Occurrence, biosynthesis, and putative role of ecdysteroids in plants, Crit. Rev. Biochem. Mol. source

In the paper

“Thus, the plant produces PEs mainly from MVA via cholesterol in the mevalonate pathway of the plant cell, using acetyl-CoA as a precursor.”

What the sources say

• Not checked yetcites [13] Bakrim, 2008, Ecdysteroids in spinach (Spinacia oleracea L.): Biosynthesis, transport and regulation of levels, Plant Physiol. Biochem. source

In the paper

“Accordingly, this may suggest that the highest concentrations of PEs are present in tissues and organs that play an important role in plant growth, development, and survival, or in reproductive organs [8].”

What the sources say

• Not checked yetcites [8] Das, 2021, The phytochemical, biological, and medicinal attributes of phytoecdysteroids: An updated review, Acta Pharm. Sin. B source

In the paper

“In annual plants, PEs are primarily found in organs such as young tissue, leaves, flowers, anthers, and seeds, whereas they are less frequently present in roots and stems.”

What the sources say

• Not checked yetcites [8] Das, 2021, The phytochemical, biological, and medicinal attributes of phytoecdysteroids: An updated review, Acta Pharm. Sin. B source

In the paper

“Furthermore, these allelochemicals can alter gene expression profiles in plants via DNA binding [18,19].”

What the sources say

• Not checked yetcites [18] Strnad, 2016, Plant ecdysteroids: Plant sterols with intriguing distributions, biological effects and relations to plant hormones, Planta source
• Not checked yetcites [19] Tarkowská, D., Krampolová, E., and Strnad, M. (2020). Plant triterpenoid crosstalk: The interaction of brassinosteroids and phytoecdysteroids in Lepidium sativum. Plants, 9. source

In the paper

“In Chenopodium album, PE concentrations are the highest in anthers, young leaves, and seeds [20].”

What the sources say

• Not checked yetcites [20] Menken, 1992, Phytoecdysteroids and insect-plant relationships in the Chenopodiaceae, Proceedings of the 8th International Symposium on Insect-Plant Relationships. Series Entomologica

In the paper

“In plants, PEs effectively induce biological activities and provide defense against insects, nematodes, and several environmental stresses.”

What the sources say

• Not checked yetcites [18] Strnad, 2016, Plant ecdysteroids: Plant sterols with intriguing distributions, biological effects and relations to plant hormones, Planta source
• Not checked yetcites [19] Tarkowská, D., Krampolová, E., and Strnad, M. (2020). Plant triterpenoid crosstalk: The interaction of brassinosteroids and phytoecdysteroids in Lepidium sativum. Plants, 9. source

In the paper

“In annual plants, PEs are translocated from seeds to developing organs such as the shoots and other growing regions.”

What the sources say

• Not checked yetcites [18] Strnad, 2016, Plant ecdysteroids: Plant sterols with intriguing distributions, biological effects and relations to plant hormones, Planta source

In the paper

“PEs are found in more than 100 terrestrial fern, gymnosperm, and angiosperm plant families [4,7,18,23,24].”

What the sources say

• Not checked yetcites [4] Smagghe, G. (2009). Phytoecdysteroids: Diversity, biosynthesis and distribution. Ecdysone: Structures and Functions, Springer. source
• Not checked yetcites [7] Yusupova, U.Y., Ramazonov, N.S., Syrov, V.N., and Sagdullaev, S.S. (2022). Phytoecdysteroids. Phytoecdysteroids: Properties, Biological Activity and Applications, Springer. source
• Not checked yetcites [18] Strnad, 2016, Plant ecdysteroids: Plant sterols with intriguing distributions, biological effects and relations to plant hormones, Planta source
• Not checked yetcites [23] Adler, 1995, Biosynthesis and distribution of insect-molting hormones in plants—A review, Lipids source
• Not checked yetcites [24] Adler, 1999, Occurrence, biosynthesis, and putative role of ecdysteroids in plants, Crit. Rev. Biochem. Mol. source

In the paper

“In several perennial plants, PEs seasonal cycling takes place between underground and aerial parts.”

What the sources say

• Not checked yetcites [18] Strnad, 2016, Plant ecdysteroids: Plant sterols with intriguing distributions, biological effects and relations to plant hormones, Planta source

In the paper

“In Serratula coronata leaves and Leuzea carthamoides roots, 20-HE content reaches 1.5% (relative to the dry weight of the plant).”

What the sources say

• Not checked yetcites [28] Savchenko, R.G., Veskina, N.A., Odinokov, V.N., Benkovskaya, G.V., and Parfenova, L.V. (2022). Ecdysteroids: Isolation, chemical transformations, and biological activity. Phytochem. Rev., 1–42. source

In the paper

“Genus Silene has large amounts of diverse ECs, i.e., ca. 1–2% of the plant’s dry weight [30].”

What the sources say

• Not checked yetcites [30] Thiem, 2017, Ecdysteroids: Production in plant in vitro cultures, Phytochem. Rev. source

In the paper

“In autumn, PEs are mobilized from the shoot to underground parts, increasing PE levels in the roots [18].”

What the sources say

• Not checked yetcites [18] Strnad, 2016, Plant ecdysteroids: Plant sterols with intriguing distributions, biological effects and relations to plant hormones, Planta source

In the paper

“In particular, during the spring season, the highest concentration of PEs is present in newly developing parts such as shoots.”

What the sources say

• Not checked yetcites [18] Strnad, 2016, Plant ecdysteroids: Plant sterols with intriguing distributions, biological effects and relations to plant hormones, Planta source

In the paper

“In spinach, cotyledons and the first true leaves contain more PEs than the later leaves [21,22].”

What the sources say

• Not checked yetcites [21] Grebenok, 1993, Ecdysteroid biosynthesis during the ontogeny of spinach leaves, Phytochemistry source
• Not checked yetcites [22] Grebenok, 1994, Biosynthesis of ecdysone and ecdysone phosphates in spinach, Phytochemistry source

In the paper

“About 200 PEs are found in different plant genera, such as Polypodium, Ajuga, Silene, and Podocarpus [7,8].”

What the sources say

• Not checked yetcites [7] Yusupova, U.Y., Ramazonov, N.S., Syrov, V.N., and Sagdullaev, S.S. (2022). Phytoecdysteroids. Phytoecdysteroids: Properties, Biological Activity and Applications, Springer. source
• Not checked yetcites [8] Das, 2021, The phytochemical, biological, and medicinal attributes of phytoecdysteroids: An updated review, Acta Pharm. Sin. B source

In the paper

“A high concentration of ECs (4–5%) is found in medicinal plants such as Achyranthes and Cyathula, which are used in Chinese medicine [28].”

What the sources say

• Not checked yetcites [28] Savchenko, R.G., Veskina, N.A., Odinokov, V.N., Benkovskaya, G.V., and Parfenova, L.V. (2022). Ecdysteroids: Isolation, chemical transformations, and biological activity. Phytochem. Rev., 1–42. source

In the paper

“The taxons that are used in traditional medicines such as Leuzea carthamoides, Ajuga turkestanica, and several Pfaffia species have abundant ECs in their roots.”

What the sources say

• Not checked yetcites [7] Yusupova, U.Y., Ramazonov, N.S., Syrov, V.N., and Sagdullaev, S.S. (2022). Phytoecdysteroids. Phytoecdysteroids: Properties, Biological Activity and Applications, Springer. source
• Not checked yetcites [8] Das, 2021, The phytochemical, biological, and medicinal attributes of phytoecdysteroids: An updated review, Acta Pharm. Sin. B source
• Not checked yetcites [30] Thiem, 2017, Ecdysteroids: Production in plant in vitro cultures, Phytochem. Rev. source
• Not checked yetcites [31] Crouzet, 2009, Ecdysteroids from Cyanotis longifolia Benth. (Commelinaceae), Arch. Insect Biochem. Physiol. source
• Not checked yetcites [32] Dinan, 2009, The Karlson Lecture. Phytoecdysteroids: What use are they?, Arch. Insect Biochem. Physiol. source
• Not checked yetcites [33] Dinan, 2006, Effects and applications of arthropod steroid hormones (ecdysteroids) in mammals, J. Endocrinol. source
• Not checked yetcites [34] Gomes, 2010, Concentration of β-ecdysone (20E) in susceptible and resistant accessions of Pfaffia glomerata infected with Meloidogyne incognita and histological characterisation of resistance, Nematology source
• Not checked yetcites [35] Tan, 2011, Isolation and analysis of a new phytoecdysteroid from Cyanotis arachnoidea C. B. Clarke, Chin. J. Chromatogr.

In the paper

“In some cases, the oxo group may be located at different carbon positions, such as C-2, C-12, C-17, C-20, or C-22, along with the required C-6 position.”

What the sources say

• Not checked yetcites [4] Smagghe, G. (2009). Phytoecdysteroids: Diversity, biosynthesis and distribution. Ecdysone: Structures and Functions, Springer. source
• Not checked yetcites [8] Das, 2021, The phytochemical, biological, and medicinal attributes of phytoecdysteroids: An updated review, Acta Pharm. Sin. B source
• Not checked yetcites [15] 2015, Ecdysteroids in plants and their pharmacological effects in vertebrates and humans, Studies in Natural Products Chemistry source
• Not checked yetcites [32] Dinan, 2009, The Karlson Lecture. Phytoecdysteroids: What use are they?, Arch. Insect Biochem. Physiol. source

In the paper

“On the other hand, plants produce ECs via mevalonic acid (MVA) and cholesterol [21,22,115,116].”

What the sources say

• Not checked yetcites [21] Grebenok, 1993, Ecdysteroid biosynthesis during the ontogeny of spinach leaves, Phytochemistry source
• Not checked yetcites [22] Grebenok, 1994, Biosynthesis of ecdysone and ecdysone phosphates in spinach, Phytochemistry source
• Not checked yetcites [115] Gilbert, 2002, Control and biochemical nature of the ecdysteroidogenic pathway, Annu. Rev. Entomol. source
• Not checked yetcites [116] Guo, 1995, Developmental regulation of sterol biosynthesis in Zea mays, Lipids source

In the paper

“C27-sterols are needed for EC biosynthesis, so it has been estimated that several biosynthetic pathways are required in arthropods and insects that permit them to metabolize the C28- and C29-sterols in plants.”

In the paper

“The biosynthesis of PEs is based on the cytosolic sterol pathway.”

In the paper

“In Polypodium vulgare and Taxas baccata, a labeled feeding experiment produced labeled E and 20-HE.”

In the paper

“After applying hot water treatment to prothalli, a complete release of ECs leads to the de novo synthesis of ECs.”

What the sources say

• Not checked yetcites [104] Simon, 2011, Ecdysteroids from Polypodium vulgare L., Steroids source
• Not checked yetcites [147] Reixach, 1997, Phytoecdysteroid overproduction in Polypodium vulgare prothalli, Phytochemistry source
• Not checked yetcites [148] Reixach, 1996, Biosynthesis of ecdysteroids in in vitro prothalli cultures of Polypodium vulgare, Phytochemistry source

In the paper

“Chenopods and other plant families also contain Δ7 sterol as PEs precursor.”

In the paper

“Notably, this treatment converted 25-hydroxycholesterol into ECs [104,147,148].”

What the sources say

• Not checked yetcites [104] Simon, 2011, Ecdysteroids from Polypodium vulgare L., Steroids source
• Not checked yetcites [147] Reixach, 1997, Phytoecdysteroid overproduction in Polypodium vulgare prothalli, Phytochemistry source
• Not checked yetcites [148] Reixach, 1996, Biosynthesis of ecdysteroids in in vitro prothalli cultures of Polypodium vulgare, Phytochemistry source

In the paper

“Oxidation of ecdysone to 20-HE takes place via ecdysone 20-monooxygenase; the Δ7 sterol has a reduced chain of C-24; this was also shown in Polypodium vulgare.”

In the paper

“The location of 20-HE (radiolabeled) biosynthesis is indicated by mobilization of hydrogen from 3α- and 4β-positions to C-4 and C-5 via concomitant 1,2-Wagner–Meerwein hydride transportation from 4β- to 5β- and from 3α- to 4α-positions [8].”

What the sources say

• Not checked yetcites [8] Das, 2021, The phytochemical, biological, and medicinal attributes of phytoecdysteroids: An updated review, Acta Pharm. Sin. B source

In the paper

“The same mechanism (no EC phosphates formed) has been reported in Polypodium vulgare prothalli, where it actively produces a certain concentration of 20-HE.”

What the sources say

• Not checked yetcites [104] Simon, 2011, Ecdysteroids from Polypodium vulgare L., Steroids source
• Not checked yetcites [147] Reixach, 1997, Phytoecdysteroid overproduction in Polypodium vulgare prothalli, Phytochemistry source
• Not checked yetcites [148] Reixach, 1996, Biosynthesis of ecdysteroids in in vitro prothalli cultures of Polypodium vulgare, Phytochemistry source

In the paper

“However, the precursor of PE biosynthesis is a reduced sidechain at C-24 of Δ7 sterol [18].”

What the sources say

• Not checked yetcites [18] Strnad, 2016, Plant ecdysteroids: Plant sterols with intriguing distributions, biological effects and relations to plant hormones, Planta source

In the paper

“Several plants contain C-24 alkylated sterols, which biosynthesize C-24-alkylated PEs.”

In the paper

“Thus, PEs are synthesized from MVA pathways and cholesterol in higher plants; other pathways are still unclear and need to be explored.”

In the paper

“In Taxus shoots, EC biosynthesis alters with age [4].”

What the sources say

• Not checked yetcites [4] Smagghe, G. (2009). Phytoecdysteroids: Diversity, biosynthesis and distribution. Ecdysone: Structures and Functions, Springer. source

In the paper

“Furthermore, saponins can bind to water-insoluble sterol complexes, therefore abolishing them from existing as EC precursors [154].”

What the sources say

• Not checked yetcites [154] Oleszek, W., and Marston, A. (2000). Chemo-ecological role of spirostanol saponins in the interaction between plants and insects. Saponins in Food, Feedstuffs and Medicinal Plant, Springer. source

In the paper

“PEs provide plants tolerance against several soil nematodes and insects; thus, PE biosynthesis increases during mechanical wounding or pathogenic attack.”

What the sources say

• Not checked yetcites [21] Grebenok, 1993, Ecdysteroid biosynthesis during the ontogeny of spinach leaves, Phytochemistry source
• Not checked yetcites [22] Grebenok, 1994, Biosynthesis of ecdysone and ecdysone phosphates in spinach, Phytochemistry source
• Not checked yetcites [149] Lamhamdi, 2013, Protective role of a methanolic extract of spinach (Spinacia oleracea L.) against Pb toxicity in wheat (Triticum aestivum L.) seedlings: Beneficial effects for a plant of a nutraceutical used with animals, Environ. Sci. Pollut. Res. source
• Not checked yetcites [150] Schmelz, 1998, Damage-induced accumulation of phytoecdysteroids in spinach: A rapid root response involving the octadecanoic acid pathway, J. Chem. Ecol. source
• Not checked yetcites [151] Schmelz, 1999, Insect-induced synthesis of phytoecdysteroids in spinach, Spinacia oleracea, J. Chem. Ecol. source

In the paper

“The feedback mechanism inhibits EC biosynthesis in non-accumulating species of plants.”

In the paper

“Thus, it has been concluded that every plant synthesizes ECs [152].”

What the sources say

• Not checked yetcites [152] Devarenne, 1995, Biosynthesis of ecdysteroids in Zea mays, Phytochemistry source

In the paper

“Another possible inhibition mechanism was demonstrated in Allium porrum (Liliaceae), also known as EC-negative plants [153].”

What the sources say

• Not checked yetcites [153] Arnault, 1986, Dietary effects of phytoecdysones in the leek-moth, Acrolepiopsis assectella Zell. (Lepidoptera: Acrolepiidae), J. Chem. Ecol. source

In the paper

“Meanwhile, it has been demonstrated that JA signaling induces PE and EC biosynthesis during a pathogenic attack [18].”

What the sources say

• Not checked yetcites [18] Strnad, 2016, Plant ecdysteroids: Plant sterols with intriguing distributions, biological effects and relations to plant hormones, Planta source

In the paper

“For the first time in Zea mays, a detectable concentration of ECs was not produced.”

What the sources say

• Not checked yetcites [152] Devarenne, 1995, Biosynthesis of ecdysteroids in Zea mays, Phytochemistry source

In the paper

“In Spinacia oleracea roots, it has been found that PE production increases during pathogenic attack [21,22,149,150,151].”

What the sources say

• Not checked yetcites [21] Grebenok, 1993, Ecdysteroid biosynthesis during the ontogeny of spinach leaves, Phytochemistry source
• Not checked yetcites [22] Grebenok, 1994, Biosynthesis of ecdysone and ecdysone phosphates in spinach, Phytochemistry source
• Not checked yetcites [149] Lamhamdi, 2013, Protective role of a methanolic extract of spinach (Spinacia oleracea L.) against Pb toxicity in wheat (Triticum aestivum L.) seedlings: Beneficial effects for a plant of a nutraceutical used with animals, Environ. Sci. Pollut. Res. source
• Not checked yetcites [150] Schmelz, 1998, Damage-induced accumulation of phytoecdysteroids in spinach: A rapid root response involving the octadecanoic acid pathway, J. Chem. Ecol. source
• Not checked yetcites [151] Schmelz, 1999, Insect-induced synthesis of phytoecdysteroids in spinach, Spinacia oleracea, J. Chem. Ecol. source

In the paper

“Moreover, PEs also have wound-healing and regeneration properties in diabetic animals [155].”

What the sources say

• Not checked yetcites [155] Lafont, 2003, Practical uses for ecdysteroids in mammals including humans: And update, J. Insect Sci. source

In the paper

“PEs, e.g., ajugacetalsterone E, inhibit cell proliferation of cell lines as anticancer agents [83].”

What the sources say

• Not checked yetcites [83] Chen, 2018, Neo-clerodane diterpenes and phytoecdysteroids from Ajuga decumbens Thunb. and evaluation of their effects on cytotoxic, superoxide anion generation and elastase release in vitro, Fitoterapia source

In the paper

“PEs isolated from Ajuga forrestii display cytotoxic activity against human lung cancer cells, hepatoma cells, breast cancer cells, and carcinoma cells [28].”

What the sources say

• Not checked yetcites [28] Savchenko, R.G., Veskina, N.A., Odinokov, V.N., Benkovskaya, G.V., and Parfenova, L.V. (2022). Ecdysteroids: Isolation, chemical transformations, and biological activity. Phytochem. Rev., 1–42. source

In the paper

“PEs display antimicrobial properties against multi-resistant strains such as Escherichia coli, Staphylococcus aureus, Candida albicans, and Klebsiella pneumoniae [60].”

What the sources say

• Not checked yetcites [60] Aliouche, 2018, Isolation, antioxidant and antimicrobial activities of ecdysteroids from Serratula cichoracea, Curr. Bioact. Compd. source

In the paper

“In quinoa-seed extract, PEs have adaptogenic activity on postmenopausal syndrome [164].”

What the sources say

• Not checked yetcites [164] Ehrhardt, 2011, The effects of 20-hydroxyecdysone and 17β-estradiol on the skin of ovariectomized rats, Menopause source

In the paper

“ECs reduce the adverse effects of stress by inducing the release of glucocorticoids in mice [163].”

What the sources say

• Not checked yetcites [163] Dai, 2017, Beta-ecdysone protects mouse osteoblasts from glucocorticoid-induced apoptosis in vitro, Planta Med. source

In the paper

“PEs isolated from Vitex doniana act as an antidepressant.”

What the sources say

• Not checked yetcites [163] Dai, 2017, Beta-ecdysone protects mouse osteoblasts from glucocorticoid-induced apoptosis in vitro, Planta Med. source

In the paper

“It was found that mechanistically low doses of PEs and cisplatin affect cells biochemical dynamics, altering DNA and protein biosynthesis in the thymus, liver, pancreas, spleen, and adrenals of tumor-bearing mice.”

What the sources say

• Not checked yetcites [161] Konovalova, 2002, Ecdysterone modulates antitumor activity of cytostatics and biosynthesis of macromolecules in tumor-bearing mice, Biol. Bull. Russ. Acad. Sci. source

In the paper

“Calonysterone has a stronger effect on Akt (a serine/threonine-specific protein kinase) phosphorylation in mammalian skeletal muscle cells [162].”

What the sources say

• Not checked yetcites [162] Issaadi, 2017, Centrifugal partition chromatography in the isolation of minor ecdysteroids from Cyanotis arachnoidea, J. Chromatogr. B source

In the paper

“PEs exhibit their toxicity against mouse T-cell lymphoma cells and their subcell line transfected with retrovirus overexpressing the ABCB1 efflux pump.”

What the sources say

• Not checked yetcites [121] Martins, 2015, Ecdysteroids sensitize MDR and non-MDR cancer cell lines to doxorubicin, paclitaxel, and vincristine but tend to protect them from cisplatin, BioMed Res. Int. source
• Not checked yetcites [160] Martins, 2012, Significant activity of ecdysteroids on the resistance to doxorubicin in mammalian cancer cells expressing the human ABCB1 transporter, J. Med. Chem. source

In the paper

“ECs and their analogs enhance protein biosynthesis by interacting with insulin signal transduction in insects [165].”

What the sources say

• Not checked yetcites [165] Cai, 2016, 20-hydroxyecdysone activates Forkhead box O to promote proteolysis during Helicoverpa armigera molting, Development

In the paper

“Moreover, high doses of PEs and cisplatin affect metastatic activity [161].”

What the sources say

• Not checked yetcites [161] Konovalova, 2002, Ecdysterone modulates antitumor activity of cytostatics and biosynthesis of macromolecules in tumor-bearing mice, Biol. Bull. Russ. Acad. Sci. source

In the paper

“The dry weight of Leuzea carthamoides contains many PEs relative to the number of phytohormones [169].”

What the sources say

• Not checked yetcites [169] Koudela, 1995, Stimulation of growth and development in Japanase quails after oral administration of ecdysteroid-containing diet, Eur. J. Entomol.

In the paper

“ECs isolated from Chenopodium album have been used to evaluate their bioactivity on Lactuca sativa seeds; an aqueous solution of PEs ranging from 10−4 to 10−7 M was taken. However, the application of PEs did not significantly increase seed germination, root length, or shoot length of Lactuca sativa [170].”

What the sources say

• Not checked yetcites [170] DellaGreca, 2005, Structure elucidation and phytotoxicity of ecdysteroids from Chenopodium album, Chem. Biodivers. source

In the paper

“They induce glycogen accumulation in the heart and liver.”

In the paper

“Exogenous ecdysone or 20-HE (10−10–10−8 M) facilitates the growth and development of the alga Chlorella vulgaris [174,175,176,177].”

What the sources say

• Not checked yetcites [174] Smith, 1992, The effect of ecdysterone on the cyanobacterium Nostoc 6720, Z. Naturforsch. C source
• Not checked yetcites [175] Bajguz, 2004, Effects of ecdysteroids on Chlorella vulgaris, Physiol. Plant. source
• Not checked yetcites [176] Bajguz, 2004, Protective role of 20-hydroxyecdysone against lead stress in Chlorella vulgaris cultures, Phytochemistry source
• Not checked yetcites [177] Bajguz, 2001, Effect of ecdysone application on the growth and biochemical changes in Chlorella vulgaris cells, Plant Physiol. Biochem. source

In the paper

“They regulate blood sugar levels during alloxan-induced hyperglycemia.”

In the paper

“Thus, it was found that a 20-HE concentration of 10−4 M proved to be more beneficial than 10−5 M, as it induced seedling growth in different cotton varieties [172].”

What the sources say

• Not checked yetcites [172] Khaitov, 2013, Plant-derived 20-hydroxyecdysone alleviates salt stress in cotton (Gossypium hirsutum L.) seedlings, Asian Australas. J. Plant Sci. Biotechnol.

In the paper

“PEs also exhibit effects such as the ET ‘triple response’ in vegetable seeds [171].”

What the sources say

• Not checked yetcites [171] Bakrim, 2007, Effects of plant hormones and 20-hydroxyecdysone on tomato (Lycopersicum esculentum) seed germination and seedlings growth, Afr. J. Biotechnol. source

In the paper

“In warm-blooded species, PEs can regulate fat accumulation and fat-level reduction [167].”

What the sources say

• Not checked yetcites [167] Lin, M., Han, P., Li, Y., Wang, W., Lai, D., and Zhou, L. (2019). Quinoa secondary metabolites and their biological activities or functions. Molecules, 24. source

In the paper

“In spinach, several PE types are detected in seeds and shoots.”

What the sources say

• Not checked yetcites [180] Cheng, 2010, Variation in phytoecdysteroid accumulation in seeds and shoots of Spinacia oleracea L. accessions, HortScience source

In the paper

“PEs play a chief role in inducing photosynthesis [19].”

What the sources say

• Not checked yetcites [19] Tarkowská, D., Krampolová, E., and Strnad, M. (2020). Plant triterpenoid crosstalk: The interaction of brassinosteroids and phytoecdysteroids in Lepidium sativum. Plants, 9. source

In the paper

“PEs positively regulate photosynthesis in plants.”

In the paper

“In Limnanthes alba, PE production and accumulation are associated with flowering.”

What the sources say

• Not checked yetcites [145] Dinan, 2002, Phytoecdysteroid levels and distribution during development in Limnanthes alba Hartw. ex Benth. (Limnanthaceae), Z. Naturforsch. C source

In the paper

“PEs produce allelochemicals, which are chemicals released by plants into the soil that hamper the growth and development of neighboring plants, fungi, microbes, or insects.”

In the paper

“PEs are polar steroidal secondary metabolites that display a defensive role against biotic (e.g., insects and, nematodes) and abiotic (e.g., heavy metals, radiation, and salinity) stress by increasing the growth, development, and biochemical responses that reduce the lethality of stress (Figure 5).”

In the paper

“PEs exhibit antistress activity in animals, algae, and plants [173].”

What the sources say

• Not checked yetcites [173] Lamhamdi, 2016, 20-Hydroxyecdysone protects wheat seedlings (Triticum aestivum L.) against lead stress, Plant Physiol. Biochem. source

In the paper

“Additionally, PE application induces RuBP production [18].”

What the sources say

• Not checked yetcites [18] Strnad, 2016, Plant ecdysteroids: Plant sterols with intriguing distributions, biological effects and relations to plant hormones, Planta source

In the paper

“They can be used in non-chemical weed management [189].”

What the sources say

• Not checked yetcites [189] Ruiz, 2013, Quinoa biodiversity and sustainability for food security under climate change. A review, Agron. Sustain. Dev. source

In the paper

“Lychnis flos-cuculi (ragged robin) contains PEs that provide plants with high medicinal value.”

What the sources say

• Not checked yetcites [185] Kikowska, 2019, Various in vitro systems of Ragged Robin (Lychnis flos-cuculi L.): A new potential source of phytoecdysteroids?, Plant Cell Tissue Organ Cult. source

In the paper

“Asparagus dumosus extract containing PEs displays antifungal and antibacterial properties [186].”

What the sources say

• Not checked yetcites [186] Ahmad, 1996, An antimicrobial ecdysone from Asparagus dumosus, Fitoterapia

In the paper

“In Spinacia oleracea, root damage and root herbivory by larvae of the dark-winged fungus gnat (Bradysia impatiens) increases the accumulation of PEs.”

What the sources say

• Not checked yetcites [193] Schmelz, 2002, Interactions between Spinacia oleracea and Bradysia impatiens: A role for phytoecdysteroids, Arch. Insect Biochem. Physiol. source

In the paper

“20-HE and its analog ponasterone A inhibit ecdysis of pink bollworm Pectinophora gossypiella [192].”

What the sources say

• Not checked yetcites [192] Hedin, 1983, Isolation of phytoecdysones as insect ecdysis inhibitors and feeding deterrents, Plant Resistance to Insects source

In the paper

“PEs play a crucial role in enhancing immunity against plant-parasitic nematodes.”

What the sources say

• Not checked yetcites [190] Soriano, 2004, Phytoecdysteroids: A novel defense against plant-parasitic nematodes, J. Chem. Ecol. source

In the paper

“It has been reported that PEs play a protective role in Spinacia oleracea against four classes of nematodes: Heterodera avenae (cereal cyst nematode), Heterodera schachtii (sugarbeet cyst nematode), Pratylenchus neglectus (root-lesion nematode), and Meloidogyne javanica (root-knot nematode).”

What the sources say

• Not checked yetcites [190] Soriano, 2004, Phytoecdysteroids: A novel defense against plant-parasitic nematodes, J. Chem. Ecol. source

In the paper

“Thus, it can be concluded that wounding increases root 20-HE accumulation as the result of induced de novo 20-HE synthesis in the root [151].”

What the sources say

• Not checked yetcites [151] Schmelz, 1999, Insect-induced synthesis of phytoecdysteroids in spinach, Spinacia oleracea, J. Chem. Ecol. source

In the paper

“Induction of PEs along with JA reduces the toxic effect of Bradysia impatiens on plants.”

What the sources say

• Not checked yetcites [193] Schmelz, 2002, Interactions between Spinacia oleracea and Bradysia impatiens: A role for phytoecdysteroids, Arch. Insect Biochem. Physiol. source

In the paper

“In the bracken fern (Pteridium aquilinum), PEs play a potent defensive role in enhancing immunity against phytophagous insect attacks [194].”

What the sources say

• Not checked yetcites [194] Jones, 1978, Role of phytoecdysteroids in bracken fern, Pteridium aquilinum (L.) Kuhn as a defense against phytophagous insect attack, J. Chem. Ecol. source

In the paper

“In Silene claviformis, the sum of PEs exhibits potent stress-protective activity [201].”

What the sources say

• Not checked yetcites [201] Ramazonov, 2020, Anti stress activity of phytoecdysteroids isolated from aerial part of Silene claviformis, Chem. Sci. Int. J. source

In the paper

“The addition of 2,4-dichlorophenoxy acetic acid (2,4-D), which is synthetic auxin in the cultures of a plant transformed by Agrobacterium rhizogenes, displays a dose-dependent inhibitory effect on PE concentration.”

What the sources say

• Not checked yetcites [203] Bajaj, Y.P.S. (1996). Serratula tinctoria L. (Dyer’s Savory): In vitro culture and the production of ecdysteroids and other secondary metabolites. Medicinal and Aromatic Plants IX, Springer.

In the paper

“However, Pfaffia glomerata moderates salinity by increasing the production of 20-HE, salicylic acid, osmolytes, and antioxidants.”

What the sources say

• Not checked yetcites [198] Felipe, 2019, Salinity-induced modifications on growth, physiology and 20-hydroxyecdysone levels in Brazilian-ginseng [Pfaffia glomerata (Spreng.) Pedersen], Plant Physiol. Biochem. source

In the paper

“A high-productivity putative mutant that produces a high level of 20-HE has been used to propagate PEs on a large scale for commercial use [184].”

What the sources say

• Not checked yetcites [184] Kapare, 2017, Impact of gamma irradiation induced variation in cell growth and phytoecdysteroid production in Sesuvium portulacastrum, J. Plant Growth Regul. source

In the paper

“In cell suspension culture from Serratula tinctoria, free and esterified PEs have been found, including 20-HE, polypodine B, and 20-HE-3-acetate [202].”

What the sources say

• Not checked yetcites [202] Chapuis, 1993, Sterol and ecdysteroid profiles of Serratula tinctoria (L.): Plant and cell cultures producing steroids, Insect Biochem. Mol. Biol. source

In the paper

“PEs alleviate salt stress in wheat seedlings by increasing the activity of antioxidants such as catalase, peroxidase, and SOD and the contents of ascorbic acid and glutathione, which reduce ROS and MDA content and promote wheat tolerance to salinity [200].”

What the sources say

• Not checked yetcites [200] Li, 2018, 20-hydroxyecdysone protects wheat seedlings from salt stress, Arch. Biol. Sci. source

In the paper

“In plants, the callus of dedifferentiated cells can increase and grow to a large extent in a disorganized manner, similar to human cancerous cells; thus, iPR showed pronounced effects in plants in the re-differentiation of cells to form adventitious buds [206].”

What the sources say

• Not checked yetcites [206] Tanimoto, 1982, Effects of cytokinin and anticytokinin on the initial stage of adventitious bud differentiation in the epidermis of Torenia stem segments, Plant Cell Physiol.

In the paper

“Therefore, the anti-aging bioactivity of CKs reduces age-associated deaths in adults by reducing development during larval and pupal stages [211].”

What the sources say

• Not checked yetcites [211] Sharma, 1995, Plant growth hormone kinetin delays aging, prolongs the lifespan, and slows down development of the fruitfly Zaprionus paravittiger, Biochem. Biophys. Res. Commun. source

In the paper

“In Locusta migratoria (migratory locust), 22-N6-(isopentenyl) adenosine monophosphoric ester of ecdysone was found in high concentration in newly laid eggs [207].”

What the sources say

• Not checked yetcites [207] Tsoupras, 1983, A cytokinin (isopentenyl-adenosyl-mononucleotide) linked to ecdysone in newly laid eggs of Locusta migratoria, Science source

In the paper

“Additionally, 6-furfurylaminopurine delays senescence and aging (increased lifespan) in several plants and Zaprionus paravittiger (fruit flies).”

What the sources say

• Not checked yetcites [211] Sharma, 1995, Plant growth hormone kinetin delays aging, prolongs the lifespan, and slows down development of the fruitfly Zaprionus paravittiger, Biochem. Biophys. Res. Commun. source

In the paper

“Among them, only a single compound shows a PE-synergistic relationship, albeit at higher doses and with weaker activity (2000-fold) than 20-HE [214].”

What the sources say

• Not checked yetcites [214] Voigt, 2001, The ecdysteroid agonist/antagonist and brassinosteroid-like activities of synthetic brassinosteroid/ecdysteroid hybrid molecules, Cell. Mol. Life Sci. source

In the paper

“ECs display weak or no bioactivity inside the bioassay of BR-responsive plants.”

What the sources say

• Not checked yetcites [205] Machackova, 1995, Comparison between the effects of 20-hydroxyecdysone and phytohormones on growth and development in plants, Eur. J. Entomol.

In the paper

“In Oryza sativa lamina inclination bioassay and Drosophila melanogaster BII cell bioassay, synthetic analogs of BRs and ECs are made in labs to detect their synergistic or antagonist relationships.”

What the sources say

• Not checked yetcites [214] Voigt, 2001, The ecdysteroid agonist/antagonist and brassinosteroid-like activities of synthetic brassinosteroid/ecdysteroid hybrid molecules, Cell. Mol. Life Sci. source

In the paper

“Jasmonic acid (JA) and its derivatives play a potent role in enhancing plant tolerance against various biotic and abiotic stresses.”

What the sources say

• Not checked yetcites [218] Wasternack, 2013, Jasmonates: Biosynthesis, perception, signal transduction and action in plant stress response, growth and development. An update to the 2007 review in Annals of Botany, Ann. Bot. source

In the paper

“JA rapidly accumulates during plant mechanical wounding or after an attack by pathogens, e.g., herbivores or insects.”

What the sources say

• Not checked yetcites [218] Wasternack, 2013, Jasmonates: Biosynthesis, perception, signal transduction and action in plant stress response, growth and development. An update to the 2007 review in Annals of Botany, Ann. Bot. source

In the paper

“However, after 2 days, induction of PE concentration was identified in shoots.”

What the sources say

• Not checked yetcites [150] Schmelz, 1998, Damage-induced accumulation of phytoecdysteroids in spinach: A rapid root response involving the octadecanoic acid pathway, J. Chem. Ecol. source
• Not checked yetcites [193] Schmelz, 2002, Interactions between Spinacia oleracea and Bradysia impatiens: A role for phytoecdysteroids, Arch. Insect Biochem. Physiol. source

In the paper

“Further, the exogenous application of BL changes the PE profile in younger leaves.”

What the sources say

• Not checked yetcites [217] Kamlar, 2015, The effect of exogenous 24-epibrassinolide on the ecdysteroid content in the leaves of Spinacia oleracea L., Steroids source

In the paper

“It has been found that mechanically damaged roots showed a higher accumulation of 20-HE, by about 3-fold, within 2 days.”

What the sources say

• Not checked yetcites [150] Schmelz, 1998, Damage-induced accumulation of phytoecdysteroids in spinach: A rapid root response involving the octadecanoic acid pathway, J. Chem. Ecol. source
• Not checked yetcites [193] Schmelz, 2002, Interactions between Spinacia oleracea and Bradysia impatiens: A role for phytoecdysteroids, Arch. Insect Biochem. Physiol. source

In the paper

“In hydroponically grown spinach, increases in PE concentrations have been reported after MeJA application.”

What the sources say

• Not checked yetcites [150] Schmelz, 1998, Damage-induced accumulation of phytoecdysteroids in spinach: A rapid root response involving the octadecanoic acid pathway, J. Chem. Ecol. source
• Not checked yetcites [193] Schmelz, 2002, Interactions between Spinacia oleracea and Bradysia impatiens: A role for phytoecdysteroids, Arch. Insect Biochem. Physiol. source

In the paper

“PEs and BRs participate in plant growth and developmental processes by modulating photosynthetic traits.”

What the sources say

• Not checked yetcites [19] Tarkowská, D., Krampolová, E., and Strnad, M. (2020). Plant triterpenoid crosstalk: The interaction of brassinosteroids and phytoecdysteroids in Lepidium sativum. Plants, 9. source
• Not checked yetcites [183] Yusupova, U.Y., Ramazonov, N.S., Syrov, V.N., and Sagdullaev, S.S. (2022). Biological activities of phytoecdisteroids. Phytoecdysteroids: Properties, Biological Activity and Applications, Springer. source

In the paper

“These responses validate the effective role of PEs in boosting plant immunity during a pathogenic attack [218].”

What the sources say

• Not checked yetcites [218] Wasternack, 2013, Jasmonates: Biosynthesis, perception, signal transduction and action in plant stress response, growth and development. An update to the 2007 review in Annals of Botany, Ann. Bot. source

In the paper

“Thus, it can be concluded that BR application leads to modulation of PE profiles inside plant tissues within hours [217].”

What the sources say

• Not checked yetcites [217] Kamlar, 2015, The effect of exogenous 24-epibrassinolide on the ecdysteroid content in the leaves of Spinacia oleracea L., Steroids source

In the paper

“Application of brassinolide (BL) (10−8 or 10−6 M) alters PE content to a certain degree, which largely depends on leaves’ developmental stages.”

What the sources say

• Not checked yetcites [217] Kamlar, 2015, The effect of exogenous 24-epibrassinolide on the ecdysteroid content in the leaves of Spinacia oleracea L., Steroids source

In the paper

“In Lycopersicum esculentum, 20-HE and GA3 mutually induce germination rate, shoot–root length, and proline content [171].”

What the sources say

• Not checked yetcites [171] Bakrim, 2007, Effects of plant hormones and 20-hydroxyecdysone on tomato (Lycopersicum esculentum) seed germination and seedlings growth, Afr. J. Biotechnol. source

In the paper

“Additionally, after ET production and its effect disappear, ecdysone has a tiny effect on ET biosynthesis and bioactivity [18].”

What the sources say

• Not checked yetcites [18] Strnad, 2016, Plant ecdysteroids: Plant sterols with intriguing distributions, biological effects and relations to plant hormones, Planta source

In the paper

“Nevertheless, shoot concentration was unchanged after Spodoptera exigua (insect) attack.”

What the sources say

• Not checked yetcites [150] Schmelz, 1998, Damage-induced accumulation of phytoecdysteroids in spinach: A rapid root response involving the octadecanoic acid pathway, J. Chem. Ecol. source
• Not checked yetcites [193] Schmelz, 2002, Interactions between Spinacia oleracea and Bradysia impatiens: A role for phytoecdysteroids, Arch. Insect Biochem. Physiol. source

In the paper

“Further experiments on spinach using the same inducible system found that, after an attack by Bradysia impatiens (dark-winged gnat fungus), MeJA application increased PE production about four- to seven-fold in roots.”

What the sources say

• Not checked yetcites [193] Schmelz, 2002, Interactions between Spinacia oleracea and Bradysia impatiens: A role for phytoecdysteroids, Arch. Insect Biochem. Physiol. source

In the paper

“Furthermore, 20-HE accumulation reduced larval establishment of dark-winged gnat fungus.”

What the sources say

• Not checked yetcites [193] Schmelz, 2002, Interactions between Spinacia oleracea and Bradysia impatiens: A role for phytoecdysteroids, Arch. Insect Biochem. Physiol. source

In the paper

“Thus, PEs are potently inducible against attacks by insects and herbivores [193].”

What the sources say

• Not checked yetcites [193] Schmelz, 2002, Interactions between Spinacia oleracea and Bradysia impatiens: A role for phytoecdysteroids, Arch. Insect Biochem. Physiol. source

In the paper

“In Chenopodium rubrum and dwarf Zea mays, ecdysone application results in slight ET production.”

What the sources say

• Not checked yetcites [18] Strnad, 2016, Plant ecdysteroids: Plant sterols with intriguing distributions, biological effects and relations to plant hormones, Planta source