Variation in phytoecdysteroid accumulation in seeds and shoots of Spinacia oleracea L. accessions
Open accessAbstract
Spinach ( Spinacea oleracea L.) is a valuable agricultural crop that accumulates phytoecdysteroids, polyhydroxylated triterpenoids, which may play a role in plant defense and have purported health benefits for human consumers. In this study, phytoecdysteroid accumulation was measured in seeds and shoots of 15 spinach accessions to determine whether phytoecdysteroid levels vary between spinach varieties and whether seed content could reliably predict relative levels in the edible foliage. Additionally, phytosterols, precursors to phytoecdysteroids, were examined to determine potential points of regulation of spinach phytoecdysteroid biosynthesis. Significant variations in phytoecdysteroid levels between accessions were observed ( P < 0.05), suggesting the potential for genetic manipulation through traditional breeding or genetic engineering to increase phytoecdysteroid levels in spinach. However, results suggest that estimation of phytoecdysteroid levels in shoots may not be achieved by measuring levels in the seeds. Levels of phytoecdysteroids in spinach ranged from 19.9 to 44.1 μg per shoot, 0.7 to 1.2 μg·mg −1 dry mass shoot, 3.2 to 9.6 μg per seed, and 0.5 to 1.1 μg·mg −1 seed. Several phytosterols connected to the phytoecdysteroid biosynthetic pathway were identified by gas chromatography–mass spectroscopy, predominantly spinasterol, 5-dihydroergosterol, and 22-dihydrospinasterol, which comprised 79.8%, 6.3%, and 4.6% of the total phytosterol content, respectively. Detection of the phytosterols cycloartenol and lanosterol in spinach suggests that spinach may also have dual biosynthetic pathways to phytosterols that contribute to the production of phytoecdysteroids.
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Written from the paper’s full text.
TL;DR
Phytoecdysteroid levels varied significantly among 15 spinach accessions in both seeds and shoots, but seed content did not reliably predict shoot content. Phytosterol profiling identified spinasterol as the dominant sterol and detected cycloartenol and lanosterol, suggesting possible dual phytosterol biosynthetic pathways in spinach.
Key findings
- Significant variation in phytoecdysteroid accumulation was observed among accessions (P < 0.05): shoots ranged from 19.9 to 44.1 mg per shoot and 0.7 to 1.2 mg·mg⁻¹ dry mass shoot; seeds ranged from 3.2 to 9.6 mg per seed and 0.5 to 1.1 mg·mg⁻¹ seed.
- Seed and shoot phytoecdysteroid levels showed moderate correlations (per mass of seeds vs. per dry mass shoots: r = 0.52, P = 0.04, n = 15; per seed vs. per dry mass shoots: r = 0.58, P = 0.02, n = 15), but these were driven by an outlier and became non-significant when it was removed.
- Phytosterols in accession NSL 92513 were predominantly spinasterol (79.8% of total phytosterols), followed by 5-dihydroergosterol (6.3%) and 22-dihydrospinasterol (4.6%); cycloartenol (0.8%) and lanosterol (0.6%) were also detected.
- Leafminer-resistant genotypes did not show higher 20-hydroxyecdysone levels, contrary to the hypothesis that higher phytoecdysteroids account for greater leafminer resistance.
Methods
- Fifteen *Spinacia oleracea* accessions from USDA ARS GRIN were grown in vitro on Murashige and Skoog media; shoots from 5–9 plants per accession were harvested after the sixth true leaf (21–36 d after germination), and seeds were sampled in three replicates per accession.
- Phytoecdysteroids were extracted in methanol, partitioned with hexane, and quantified by HPLC as 20-hydroxyecdysone equivalents resulting from coelution of 20E and polypodine B; ANOVA with Fisher’s LSD and Pearson correlations were used.
- Phytosterols were extracted from shoots, saponified, purified, derivatized, and analyzed by GC-MS, with lathosterol as an internal standard and concentrations reported as lathosterol equivalents per gram dry mass.
Limitations
- The paper does not state a formal limitations section. Evident limitations include the use of in vitro-grown plants, which may not reflect field conditions, and a limited set of 15 accessions selected based on insect resistance, seed size, and flowering/bolting traits.
- The seed-to-shoot correlation was moderate and driven by an outlier; after its removal, correlations were no longer significant, so the conclusion that seed content cannot reliably predict shoot content is based on this dataset.
- Phytosterol profiling was performed only on accession NSL 92513, and lathosterol was not detected, limiting broader pathway inferences; the leafminer-resistance hypothesis was also not supported, with possible explanations including insect adaptation or other defensive compounds.
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- Findingown data
Phytoecdysteroid levels in shoots of in vitro grown spinach ranged from 0.7 to 1.2 mg/mg dry mass shoot.
“and from 0.7 to 1.2 mgÁmg -1 dry mass shoot”
- Also stated in: § Abstract · plain
“0.7 to 1.2 mgÁmg -1 dry mass shoot”
Quote in the paperNot a substantial claim - Also stated in: § Abstract · plain
- Findingown data
Phytoecdysteroid levels in shoots of in vitro grown spinach ranged from 19.9 to 44.1 mg per shoot.
“Levels of phytoecdysteroids from spinach grown in vitro ranged from 19.9 to 44.1 mg per shoot”
- Also stated in: § Abstract · plain
“Levels of phytoecdysteroids in spinach ranged from 19.9 to 44.1 mg per shoot”
Quote in the paperNot a substantial claim - Also stated in: § Abstract · plain
- Findingown data
Phytoecdysteroid content in seeds ranged from 0.5 to 1.1 mg/mg seed.
“and 0.5 to 1.1 mgÁmg -1 per seed”
- Also stated in: § Abstract · plain
“0.5 to 1.1 mgÁmg -1 seed”
Quote in the paperNot a substantial claim - Also stated in: § Abstract · plain
- Findingown data
Phytoecdysteroid content in seeds ranged from 3.2 to 9.6 mg per seed.
“In seeds, the phytoecdysteroid content ranged from 3.2 to 9.6 mg per seed”
- Also stated in: § Abstract · plain
“3.2 to 9.6 mg per seed”
Quote in the paperNot a substantial claim - Also stated in: § Abstract · plain
- Findingown data
Phytoecdysteroid accumulation varied significantly among the 15 spinach accessions on a per shoot, per dry mass shoot, per seed, and per seed mass basis (P < 0.05).
“Accumulation of phytoecdysteroids, measured as 20E equivalents in seeds and shoots of S. oleracea, varied significantly among accessions on a per shoot, per dry mass shoot, per seed, and per seed mass basis (Table 2).”
- Also stated in: § Abstract · plain
“Significant variations in phytoecdysteroid levels between accessions were observed (P < 0.05), suggesting the potential for genetic manipulation through traditional breeding or genetic engineering to increase phytoecdysteroid levels in spinach.”
Quote in the paperSubstantial claim - Also stated in: § Abstract · plain
- Findingown data
Lathosterol was not detected in this study.
“lathosterol was not detected in this study.”
Quote in the paperNot a substantial claim - Findingown data
Stigmasterol comprised 2.4% of total phytosterols in spinach shoots.
“Other common phytosterols accumulated lower levels, including 5-dihydroergosterol (6.3%), 22dihydrospinasterol (4.6%), stigmasterol (2.4%), and cholesterol (2.2%).”
Quote in the paperNot a substantial claim - Findingown data
A moderate but significant correlation was found between phytoecdysteroid accumulation per seed and per dry mass of shoots (r = 0.58, P = 0.02, n = 15) among the 15 spinach accessions.
“and between phytoecdysteroid accumulation per seed and phytoecdysteroid accumulation per dry mass of shoots (r = 0.58, P = 0.02, n = 15) among the 15 spinach accessions.”
Quote in the paperSubstantial claim - Findingown data
Lanosterol was detected in spinach at 0.6% of total phytosterols.
“Cycloartenol (0.8% of total phytosterols) and lanosterol (0.6% of total phytosterols) were detected in spinach.”
Quote in the paperNot a substantial claim - Findingown data
5-dihydroergosterol comprised 6.3% of total phytosterols in spinach shoots.
“Other common phytosterols accumulated lower levels, including 5-dihydroergosterol (6.3%), 22dihydrospinasterol (4.6%), stigmasterol (2.4%), and cholesterol (2.2%).”
Quote in the paperNot a substantial claim - Findingown data
Spinasterol was the predominant phytosterol in spinach shoots, making up an average of 79.8% of total phytosterols identified.
“Spinasterol was the predominant phytosterol, making up an average of 79.8% of total phytosterols identified”
- Also stated in: § Abstract · plain
“predominantly spinasterol, 5-dihydroergosterol, and 22-dihydrospinasterol, which comprised 79.8%, 6.3%, and 4.6% of the total phytosterol content, respectively.”
Quote in the paperNot a substantial claim - Also stated in: § Abstract · plain
- Findingown data
Phytoecdysteroids were detected in all seeds and shoots of S. oleracea accessions selected for this study.
“In this study, phytoecdysteroids were detected in all seeds and shoots of S. oleracea accessions selected for this study.”
Quote in the paperSubstantial claim - Findingown data
A moderate but significant correlation was found between phytoecdysteroid accumulation per mass of seeds and per dry mass of shoots (r = 0.52, P = 0.04, n = 15) among the 15 spinach accessions.
“A moderate but significant correlation was found in phytoecdysteroid accumulation per mass of seeds and per dry mass of shoots (r = 0.52, P = 0.04, n = 15)”
Quote in the paperSubstantial claim - Findingown data
The level of 20E in genotypes PI 274065, PI 174385, PI 169673, PI 175312, and PI 433208 did not reflect the hypothesis that high levels of phytoecdysteroids may account for greater leafminer resistance.
“However, the level of 20E in these genotypes, as measured in our screening (Tables 2), did not reflect the hypothesis that high levels of phytoecdysteroids may account for greater leafminer resistance.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Mou, 2008
Not checked in the free run.
- Rests on an assumption: The leafminer resistance rankings from Mou (2008) are comparable to the accessions' phytoecdysteroid levels measured in this study.
- Findingown data
Cycloartenol was detected in spinach at 0.8% of total phytosterols.
“Cycloartenol (0.8% of total phytosterols) and lanosterol (0.6% of total phytosterols) were detected in spinach.”
Quote in the paperNot a substantial claim - Findingown data
22-dihydrospinasterol comprised 4.6% of total phytosterols in spinach shoots.
“Other common phytosterols accumulated lower levels, including 5-dihydroergosterol (6.3%), 22dihydrospinasterol (4.6%), stigmasterol (2.4%), and cholesterol (2.2%).”
Quote in the paperNot a substantial claim - Interpretationown data
Detection of cycloartenol and lanosterol in spinach suggests that spinach may also have dual biosynthetic pathways to phytosterols that contribute to the production of phytoecdysteroids.
“Detection of the phytosterols cycloartenol and lanosterol in spinach suggests that spinach may also have dual biosynthetic pathways to phytosterols that contribute to the production of phytoecdysteroids.”
Quote in the paperSubstantial claim - Interpretationown data
Estimation of phytoecdysteroid levels in shoots may not be achieved by measuring levels in the seeds.
“These results suggest that estimation of phytoecdysteroid levels in shoots may not be achieved by measuring levels in the seeds.”
- Also stated in: § Abstract · hedged
“However, results suggest that estimation of phytoecdysteroid levels in shoots may not be achieved by measuring levels in the seeds.”
Quote in the paperSubstantial claim- Rests on an assumption: The outlier data point should be excluded from the correlation analysis.
- Also stated in: § Abstract · hedged
- Interpretationown data
Significant variation in phytoecdysteroid content between spinach accessions suggests potential for molecular biotechnology or conventional breeding to enhance levels of phytoecdysteroid accumulation.
“Significant variation in phytoecdysteroid content between spinach accessions suggests potential for molecular biotechnology or conventional breeding to enhance levels of phytoecdysteroid accumulation.”
- Also stated in: § Abstract · hedged
“suggesting the potential for genetic manipulation through traditional breeding or genetic engineering to increase phytoecdysteroid levels in spinach.”
Quote in the paperSubstantial claim- Rests on an assumption: Variation in phytoecdysteroid levels among accessions is due to genetic differences rather than environmental or developmental variation.
- Also stated in: § Abstract · hedged
- Premisecitation
Phytoecdysteroids play a role in plant defense.
“Taken together, these and many other studies make a firm case that phytoecdysteroids play a role in plant defense.”
Quote in the paperSubstantial claim0 of 3 sources checked- cites Adler and Grebenok, 1999
Not checked in the free run.
- cites Jones and Firn, 1978
Not checked in the free run.
- cites Schmelz et al., 1999
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- Interpretationcitation
Inhibition of carbon flux into lathosterol suggests that phytoecdysteroid regulation may also occur before the final hydroxylation steps, impacting carbon allocation early in the phytoecdysteroid pathway such as during phytosterol biosynthesis.
“Inhibition of carbon flux into lathosterol suggests that phytoecdysteroid regulation may also occur before the final hydroxylation steps, impacting carbon allocation early in the phytoecdysteroid pathway such as during phytosterol biosynthesis.”
Quote in the paperSubstantial claim
- May combine two claims. 7
- The quote only partly states this claim. 1
- Classified as probably not a substantial claim. 17
May combine two claims. The quote only partly states this claim.
Findingown dataAccessions NSL 92513 ('Bloomsdale Long-Standing') and PI 606707 ('America') had the highest and lowest phytoecdysteroid content per dry mass of shoots, respectively.
“('America') had the highest and lowest phytoecdysteroid content per dry mass of shoots, respectively.”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataAccession NSL 92513 also had the highest phytoecdysteroid content per gram seed and per seed.
“Accession NSL 92513 also had the highest phytoecdysteroid content per gram seed and per seed.”
Quote in the paperNot a substantial claimMay combine two claims.
Findingown dataAccessions PI 531456 ('Popey') and PI 606707 ('America') had the highest and lowest phytoecdysteroid content per shoot, respectively.
“Accessions PI 531456 ('Popey') and PI 606707 ('America') had the highest and lowest phytoecdysteroid content per shoot, respectively”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Findingown dataCholesterol comprised 2.2% of total phytosterols in spinach shoots.
“Other common phytosterols accumulated lower levels, including 5-dihydroergosterol (6.3%), 22dihydrospinasterol (4.6%), stigmasterol (2.4%), and cholesterol (2.2%).”
Quote in the paperNot a substantial claimMay combine two claims.
Findingown dataThe significant correlations between seed and shoot phytoecdysteroid accumulation were driven by an outlier; when the outlier was removed, the correlations were no longer significant.
“However, the significant correlation was driven by an outlier and when this data point was removed, the Pearson's correlation coefficients were no longer significant.”
Quote in the paperSubstantial claimMay combine two claims.
InterpretationcitationLathosterol was not detected likely because it did not accumulate and was subsequently metabolized, as reported by Grebenok and Adler (1993).
“However, Grebenok and Adler (1993) also reported that lathosterol did not accumulate and instead was subsequently metabolized, which likely explains why lathosterol was not identified in this analysis.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Grebenok and Adler, 1993
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationPhytoecdysteroids are biosynthesized from phytosterols through the mevalonic acid pathway.
“Phytoecdysteroids are polyhydroxylated triterpenoids biosynthesized from phytosterols through the mevalonic acid pathway (Adler and Grebenok, 1999).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Adler and Grebenok, 1999
Not checked in the free run.
Classified as probably not a substantial claim.
PremisecitationWithin a plant species, the phytoecdysteroid content in seeds may provide a gauge of the foliar content for the plant.
“Within a plant species, the phytoecdysteroid content in seeds may provide a gauge of the foliar content for the plant (Dinan et al., 2001b).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Dinan et al., 2001b
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationIn spinach, phytoecdysteroids are actively biosynthesized in older leaves and transported to newly developing apical parts of the plant, including flowers, seeds, and young leaves.
“In spinach, phytoecdysteroids are actively biosynthesized in older leaves and transported to newly developing apical parts of the plant, including flowers, seeds, and young leaves (Bakrim et al., 2008).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Bakrim et al., 2008
Not checked in the free run.
Probably not a substantial claim: may not pass S2 (checkable), S3 (load-bearing), S4 (based).
Interpretationnone givenManipulation of phytosterol biosynthesis may help to elucidate regulation of phytoecdysteroid biosynthesis.
“Additionally, manipulation of phytosterol biosynthesis may help to elucidate regulation of phytoecdysteroid biosynthesis.”
Quote in the paperNot a substantial claimClassified as probably not a substantial claim.
Study design and conductreasoningAll data were normalized to the internal standard lathosterol to allow comparison between samples.
“To allow comparison between samples, all data were normalized to the internal standard, lathosterol (Sigma), which was not detected in preliminary extracts of this accession.”
Quote in the paperSubstantial claimClassified as probably not a substantial claim. May combine two claims.
PremisecitationIn an evaluation of 180 randomly selected plant species, phytoecdysteroids were more readily detected in leaves than in seeds; however, the highest levels of phytoecdysteroids were detected only for species that were also positive for phytoecdysteroids in the seeds.
“In an evaluation of 180 randomly selected plant species, phytoecdysteroids were more readily detected in leaves than in seeds; however, the highest levels of phytoecdysteroids were detected only for species that were also positive for phytoecdysteroids in the seeds (Dinan et al., 2001b).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Dinan et al., 2001b
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationSimilar relationships between secondary compound accumulation in seed and vegetative tissues have been demonstrated for aliphatic glucosinolates, which were correlated in 35 different Arabidopsis ecotypes.
“Similar relationships between secondary compound accumulation in seed and vegetative tissues have been demonstrated for aliphatic glucosinolates, which were correlated in 35 different Arabidopsis ecotypes (Kliebenstein et al., 2001).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Kliebenstein et al., 2001
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductown dataA 100% recovery rate was achieved for the lathosterol internal standard.
“Lathosterol was added immediately before the derivatization procedure at 1 mgÁmL -1 in each chromatogram and a 100% recovery rate was achieved.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
PremisecitationLanosterol synthase genes have been induced by methyl jasmonate, suggesting that secondary metabolites produced through the lanosterol pathway may contribute to plant defense.
“Lanosterol synthase genes have been induced by methyl jasmonate, suggesting that secondary metabolites produced through the lanosterol pathway may contribute to plant defense (Ohyama et al., 2009;Suzuki et al., 2006).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Ohyama et al., 2009
Not checked in the free run.
- cites Suzuki et al., 2006
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Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationLevels of total phytosterols increased or decreased in coordination with phytoecdysteroid levels during the growth and development of spinach.
“Levels of total phytosterols increased or decreased in coordination with phytoecdysteroid levels during the growth and development of spinach (Grebenok et al., 1991).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Grebenok et al., 1991
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationFor Chenopodium species, if phytoecdysteroids were detected in the seed, they were consistently detected in the germinated plant foliage.
“For Chenopodium species, if phytoecdysteroids were detected in the seed, they were consistently detected in the germinated plant foliage (Dinan, 1992).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Dinan, 1992
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Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationIn higher plants, there are two biosynthetic pathways for phytosterol biosynthesis, through cycloartenol or through lanosterol.
“In higher plants, there are two biosynthetic pathways for phytosterol biosynthesis, through cycloartenol or through lanosterol (Benveniste, 2004).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Benveniste, 2004
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationLanosterol synthase genes were only recently identified in dicotyledonous plants.
“Lanosterol synthase genes were only recently identified in dicotyledonous plants (previously only found in animals and fungi).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Ohyama et al., 2009
Not checked in the free run.
- cites Suzuki et al., 2006
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Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationIn an excised leaf assay, [2-14C] mevalonic acid was incorporated into the phytosterol lathosterol before incorporation into 20E and other phytoecdysteroids.
“In an excised leaf assay, [2-14 C] mevalonic acid was incorporated into the phytosterol lathosterol before incorporation into 20E and other phytoecdysteroids (Grebenok and Adler, 1993).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Grebenok and Adler, 1993
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductown dataLathosterol was not detected in preliminary extracts of this accession.
“which was not detected in preliminary extracts of this accession.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductnone givenExtracted sterol samples were prepared and analyzed in triplicate.
“Extracted sterol samples were prepared and analyzed in triplicate by the Metabolomics Center in the Roy J. Carver Biotechnology Center at the University of Illinois at Urbana Champaign.”
Quote in the paperNot a substantial claim
The paper relies on this without support.
The 15 spinach accessions selected are representative of S. oleracea genetic diversity.
“Spinach seeds were obtained from the USDA germplasm repository and are representative varieties from a range of geographic locations.”
The paper relies on this without support.
Variation in phytoecdysteroid levels among accessions is due to genetic differences rather than environmental or developmental variation.
“The significant differences in phytoecdysteroid accumulation between varieties illustrate the potential for further manipulation of phytoecdysteroid levels.”
The paper relies on this without support.
The outlier data point should be excluded from the correlation analysis.
“However, the significant correlation was driven by an outlier and when this data point was removed, the Pearson's correlation coefficients were no longer significant.”
The paper relies on this without support.
The leafminer resistance rankings from Mou (2008) are comparable to the accessions' phytoecdysteroid levels measured in this study.
“In cage and field tests, S. oleracea accessions PI 274065, PI 174385, and PI 169673 had the fewest mines or lowest sting density produced by leafminers and PI 175312 and PI 433208 had high sting density in field trials (Mou, 2008).”
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
Fifteen accessions of spinach grown under controlled conditions displayed significant differences in ecdysteroid content.
“Fifteen accessions of spinach grown under controlled conditions displayed significant differences in ecdysteroid content [20].”
From Ecdysteroid Content and Therapeutic Activity in Elicited Spinach Accessions (2020) · cites this paper as [20]
Variation of PE types at growth stages is essential for conventional breeding and molecular biotechnology to promote PE levels, facilitating plant growth and productivity and improving plant immunity against stress.
“Their variation at growth stages is essential for conventional breeding and molecular biotechnology to further promote PE levels in plants, facilitating plant growth and productivity and improving plant immunity against stress [180].”
From Phytoecdysteroids: Distribution, Structural Diversity, Biosynthesis, Activity, and Crosstalk with Phytohormones (2022) · cites this paper as [180]
In spinach, several PE types are detected in seeds and shoots.
“In spinach, several PE types are detected in seeds and shoots.”
From Phytoecdysteroids: Distribution, Structural Diversity, Biosynthesis, Activity, and Crosstalk with Phytohormones (2022) · cites this paper as [180]
In spinach, several PEs that protect the plant from insects and induce plant fitness and yield are found in different concentrations at different phases of the plants' life cycle.
“In spinach, several PEs that protect the plant from insects and induce plant fitness and yield are found in different concentrations at different phases of the plants’ life cycle [180].”
From Phytoecdysteroids: Distribution, Structural Diversity, Biosynthesis, Activity, and Crosstalk with Phytohormones (2022) · cites this paper as [180]
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Research works printed but not linked: 1
- Suzuki,, 2006, 47, 565
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Arif, Y., Singh, P., Bajguz, A., & Hayat, S. (2022) · International Journal of Molecular Sciences · doi:10.3390/ijms23158664 · Cited by 50 · see citing works
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Ecdysteroid Content and Therapeutic Activity in Elicited Spinach Accessions
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Urinary Excretion of Ecdysterone and Its Metabolites Following Spinach Consumption
Yuliandra, T., Touvleliou, K., de la Torre, X., Botrè, F., Loke, S., Isenmann, E., Valder, S., Diel, P., & Parr, M. K. (2023) · Molecular Nutrition & Food Research · doi:10.1002/mnfr.202200518 · Cited by 12 · see citing works
Phytoecdysteroid and Clerodane Content in Three Wild <i>Ajuga</i> Species in Israel
Taha-Salaime, L., Davidovich-Rikanati, R., Sadeh, A., Abu-Nassar, J., Marzouk-Kheredin, S., Yahyaa, Y., Ibdah, M., Ghanim, M., Lewinsohn, E., Inbar, M., et al. (2019) · ACS Omega · doi:10.1021/acsomega.8b03029 · Cited by 10 · see citing works
Dissemond, J., Havers, T., Held, S., Geisler, S., Kostov, T., Diel, P., Türschmann, S., Parr, M. K., & Isenmann, E. (2025) · Journal of the International Society of Sports Nutrition · doi:10.1080/15502783.2025.2540408 · Cited by 2 · see citing works
- Authors
- Cheng, D. M., Yousef, G. G., & Lila, M. A.
- Affiliations
- University of Illinois Urbana-Champaign (US)
- Venue
- HortScience
- Published
- 2010-11
- Volume
- 45
- Issue
- 11
- Pages
- 1634-1638
- ISSN
- 0018-5345 (ISSN-L)
- 2327-9834
- OpenAlex
- W2150189324
- Cited by
- 18
- References
- 38
- Citations by year
- 2026: 2 · 2025: 4 · 2024: 1 · 2023: 2 · 2022: 2 · 2020: 2 · 2019: 1 · 2017: 2 · 2015: 1 · 2014: 1
- FWCI
- 0.00
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Locations
- HortScience (publishedVersion)
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Plant biochemistry and biosynthesis
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Phytochemical Studies and Bioactivities
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Natural product bioactivities and synthesis
- Keywords
- phytoecdysteroids, spinach, phytosterols, seed content, spinasterol, cycloartenol, lanosterol, genetic variation, plant defense, health benefits
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Claims read by deepseek-v4.1-flash under definition version 3 (extract-v3) on 2026-10-10.
Classified by jev-1.13-20260917 (gold-guideline-v0.2) on 2026-10-10.
Statistics and arithmetic are recomputed, not judged.
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