Bacopa monnieri (L.) wettst. Extract protects against glutamate toxicity and increases the longevity of Caenorhabditis elegans

Brimson, James Michael; Prasanth, Mani Iyer; Plaingam, Waluga; Tencomnao, Tewin

doi:10.1016/j.jtcme.2019.10.001

Cited by 51Medicinal Plants and NeuroprotectionAlzheimer's disease research and treatmentsPhytochemicals and Medicinal Plants

Bacopa monnieri (L.) wettst. Extract protects against glutamate toxicity and increases the longevity of Caenorhabditis elegans

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Brimson, James Michael, Prasanth, Mani Iyer, Plaingam, Waluga, Tencomnao, Tewin·2020·Journal of Traditional and Complementary Medicine

DOI 10.1016/j.jtcme.2019.10.001PMC PMC7484969

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Abstract

Neurodegenerative diseases, such as Alzheimer’s disease, cause a great deal of suffering for both patients and carers. Bacopa monnieri (L.) wettst. Is known for its memory-enhancing properties, and is of great interest in treating neurodegenerative disease. This study aimed to evaluate B.monnieri against glutamate toxicity, and identify whether B.monnieri reduces mitochondrial and ER stress, as well as to measure B.monnieri’s effect on the life span and aging of Caenorhabditis elegans. We hypothesized that B.monnieri would prevent cellular oxidative stress, prevent mitochondrial/ER stress, and increase the life span while reducing signs of aging in C.elegans. Glutamate toxicity was measured using viable cell staining assays and the MTT assay. ROS and mitochondrial stress were assessed by H2DCFDA and Rodamine123 staining, with fluorescence/confocal microscopy. C.elegans’ median and maximum life span were measured, in response to B.monnieri treatment, along with lipofuscin imaging to measure the health of the C.elegans population. B.monnieri hexane extract (but not ethanol extract) prevented the toxicity of 5 mM glutamate in HT-22 cells. We found that the mechanism involves the reduction of ROS production and the prevention of mitochondrial and ER stress. Furthermore, we showed that B.monnieri could increase the median and maximal lifespan of wild type C.elegans, maintain a younger appearing phenotype in the aged C.elegans. In conclusion, B.monnieri prevents mitochondrial, and oxidative stress in the cultured cells. Furthermore, it can prolong the healthy lifespan of C.elegans, indicating that B.monnieri the potential for therapeutic and preventative use in neurodegenerative disease.

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Written from the paper’s full text.

TL;DR

Hexane and dichloromethane extracts of *Bacopa monnieri* protected HT-22 neuronal cells against 5 mM glutamate toxicity by reducing ROS, mitochondrial stress, and ER stress, while the ethanol extract did not. The hexane extract also extended median and maximum lifespan and reduced lipofuscin autofluorescence in wild-type *Caenorhabditis elegans*.

Key findings

In HT-22 cells, the hexane extract protected against 5 mM glutamate with an EC50 of 1.2 ± 1.3 μg/ml (n = 4), and the DCM extract with an EC50 of 2.1 ± 1.37 μg/ml (n = 3); the ethanol extract had no effect within tested concentrations (EC50 > 10 μg/ml, n = 3).
At 10 μg/ml, hexane and DCM extracts restored viable cell numbers after glutamate to control levels by trypan blue exclusion (P < 0.0001, n = 4); ethanol showed no cytoprotective effect.
Glutamate increased ROS fluorescence (mean cellular fluorescence: control 10.93 ± 1.85; 5 mM glutamate 61.66 ± 17.60; glutamate + 10 μg/ml hexane extract 7.55 ± 1.20; n = 3), and hexane extract reduced this significantly versus glutamate alone (P < 0.0001, n = 3).
Glutamate caused mitochondrial stress/leakage by rhodamine 123 staining, which hexane extract reduced (P < 0.05, n = 4); glutamate doubled ERP57 and increased CHOP by approximately 50%, with co-treatment returning these ER-stress proteins to control levels.
In wild-type *C. elegans*, hexane extract at 7 μg/ml significantly increased survival (Log-rank p < 0.0001; Gehan-Breslow-Wilcoxon p = 0.0011, n = 3) and at 10 μg/ml (Log-rank p = 0.0085, n = 3); 3 μg/ml showed no significant difference (p = 0.09) and 80 μg/ml was not significant (p = 0.26).
Lipofuscin autofluorescence was lower in worms treated with 10 μg/ml hexane extract than controls (results text: 15.4 ± 1.7 vs 5.6 ± 1.1 fluorescence units; unpaired t-test P = 0.0006, n = 3).

Methods

HT-22 cells were treated with 5 mM glutamate with or without *B. monnieri* extracts (10 μg/ml unless otherwise stated). Assays included MTT, trypan blue exclusion, propidium iodide/Hoechst staining, rhodamine 123 for mitochondrial membrane potential, carboxy-H2DCFDA for ROS, and Western blot for CHOP and ERP57.
*B. monnieri* leaves were extracted by Soxhlet with hexane, dichloromethane, or 95% ethanol; extracts were standardized by reversed-phase HPLC for Bacoside A components, with Bacopaside II reported as most abundant at 0.38 ± 0.00001 mg/g.
Wild-type *C. elegans* (Bristol N2) were treated with hexane extract at 1–100 μg/ml; lifespan was counted every 24 h, and intestinal autofluorescence (lipofuscin) was imaged after five days. Statistical analysis used ANOVA with Dunnett’s post hoc test, log-rank (Mantel-Cox), Gehan-Breslow-Wilcoxon, and unpaired t-tests as appropriate.

Limitations

Not stated in the paper: the paper has no dedicated limitations section.
Evident from the text: the study is limited to HT-22 cultured cells and *C. elegans*, so findings are not directly demonstrated in humans or mammalian models.
Evident from the text: the active protective compounds were not identified; the authors note that future studies may use mass spectrometry to identify compounds differing between hexane and ethanol extracts, and that Bacoside A components were absent or very low in the protective hexane and DCM extracts.

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Crossref · 10.1016/j.jtcme.2019.10.001

Cites

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Claims the paper makes

0 of 35 cited sources checked · 0 supported · 0 flagged

Findingown data
Bacopa monnieri can increase the median lifespan of wild type Caenorhabditis elegans.
“B.monnieri could increase the median and maximal lifespan of wild type C.elegans”
Quote in the paperSubstantial claim
Findingown data
Bacopa monnieri can increase the maximal lifespan of wild type Caenorhabditis elegans.
“B.monnieri could increase the median and maximal lifespan of wild type C.elegans”
Quote in the paperSubstantial claim
Findingown data
Bacopa monnieri can maintain a younger appearing phenotype in aged Caenorhabditis elegans.
“maintain a younger appearing phenotype in the aged C.elegans.”
Quote in the paperSubstantial claim
Findingown data
5 mM glutamate treated HT22 cells showed significantly higher Rhodamine 123 fluorescence compared to control cells (ANOVA followed by Dunnett's post hoc test P < 0.05, n=4).
“Cells treated with 5 mM glutamate showed a statistically significant higher fluorescence compared to control cells (ANOVA followed by Dunnett’s post hoc test P < 0.05 (n = 4))”
Quote in the paperSubstantial claim
Findingown data
Glutamate caused an approximate 50% increase in CHOP expression in HT22 cells.
“an approximate 50% increase in CHOP expression”
Quote in the paperSubstantial claim

Findingown data
Lipofuscin imaging showed a significant reduction in Lipofuscin fluorescence in C. elegans treated with 10 μg/ml B. monnieri hexane extract compared to control.
“Lipofuscin Imaging (Fig. 8a–c), showed that there was a significant reduction in Lipofuscin fluorescence in the worms treated with 10 μg/ml B.monnieri hexane extract (Fig. 8b) when compared to the control worms (Fig. 8a).”
Quote in the paperSubstantial claim
Findingown data
Glutamate has an apparent effect on HT-22 cell morphology causing cell shrinkage and death.
“Glutamate has an apparent effect on cell morphology causing cell shrinkage, and death”
Quote in the paperSubstantial claim
Findingown data
None of the Bacopa monnieri extracts at 10 μg/ml affected viable HT-22 cell numbers when added alone.
“Fig. 2d also shows that none of the extracts (at 10 μg/ml) affected viable HT-22 cell numbers when added alone.”
Quote in the paperSubstantial claim
Findingown data
The hexane extract of Bacopa monnieri protects mitochondria from 5 mM glutamate-induced stress in HT-22 cells, as indicated by distinct and less diffuse rhodamine123 staining.
“In contrast to glutamate alone, the B.monnieri hexane extract appears to protect the mitochondria from 5 mM glutamate-induced stress as the staining shows distinct mitochondria, and less diffuse.”
Quote in the paperSubstantial claim
Findingown data
The hexane extract of Bacopa monnieri has a dose-dependent protective effect against toxicity caused by 5 mM glutamate in HT-22 cells, with an EC50 of 1.2 ± 1.3 μg/ml (n = 4).
“Fig. 2a shows the dose-dependent protective effect of the hexane extract of B.monnieri against the toxicity caused by 5 mM glutamate compared to control cells (treated with PBS); the EC50 value for the hexane extract was 1.2 ± 1.3 μg/ml (n = 4).”
Quote in the paperSubstantial claim
Findingown data
At 10 μg/ml, B. monnieri hexane extract caused a significant increase in C. elegans survival compared to control (Log-rank Mantel-Cox p = 0.0085, n=3).
“for 10 μg/ml Log-rank (Mantel-Cox) test (p = 0.0085 n = 3)”
Quote in the paperSubstantial claim
Findingown data
The ethanol extract of Bacopa monnieri did not affect the toxicity of 5 mM glutamate in HT-22 cells within the concentrations tested (n = 3).
“The ethanol extract of B.monnieri (Fig. 2c) did not affect (within the concentrations tested) the toxicity of 5 mM glutamate in HT-22 cells (n = 3).”
- Also stated in: § The effect of B.monnieri extracts, on glutamate toxicity in HT-22 cells · plain
  “As with the MTT assay, the ethanol extract had no cytoprotective effect in HT-22 cells treated with glutamate.”
Quote in the paperNot a substantial claim
Findingown data
At 7 μg/ml, B. monnieri hexane extract caused a significant increase in C. elegans survival compared to control (Log-rank Mantel-Cox p < 0.0001, Gehan-Breslow-Wilcoxon p = 0.0011, n=3).
“at doses between 7 and 10 μg/ml (Fig. 7b–c respectively) there was a significant increase in survival compared to control using the Log-rank (Mantel-Cox) test (p < 0.0001, n = 3) & Gehan-Breslow-Wilcoxon test (p = 0.0011, n = 3) for 7 μg/ml”
Quote in the paperSubstantial claim
Findingown data
The dichloromethane extract of Bacopa monnieri has a dose-dependent protective effect against toxicity caused by 5 mM glutamate in HT-22 cells, with an EC50 of 2.1 ± 1.37 μg/ml (n = 3).
“Fig. 2b shows the dose-dependent protective effect of the DCM extract of B.monnieri; with an EC50 value of 2.1 ± 1.37 μg/ml (n = 3).”
Quote in the paperSubstantial claim
Findingown data
All four constituents of bacoside A (bacoside A3, bacopaside II, bacopaside X, and bacopasaponin C) are present in the ethanol extract of Bacopa monnieri.
“It can be seen from Fig. 1, that all four constituents of bacoside A are present in the ethanol extract of B.monnieri.”
Quote in the paperNot a substantial claim
Findingown data
In the dichloromethane and hexane extracts of Bacopa monnieri, the bacoside A compounds appear to be missing or at very low concentrations.
“However, in the dichloromethane and hexane extracts these compounds appear to be missing or at very low concentrations (Fig. 1 c-d).”
Quote in the paperNot a substantial claim
Findingown data
Both the hexane and dichloromethane extracts of Bacopa monnieri returned the number of viable HT-22 cells up to that of the control after glutamate treatment (P < 0.0001, n = 4).
“Both the hexane and DCM extracts of B.monnieri returned the number of viable cells up to that of the control (statistically significant P= < 0.0001 (n = 4) using ANOVA followed by Dunnett’s multiple comparison post hoc test).”
Quote in the paperSubstantial claim
Findingown data
The present study did not see any indication of a protective antioxidant effect with the dose used with the ethanol extract.
“We did not see any indication of a protective antioxidant effect with the dose we used with the Ethanol extract.”
Quote in the paperSubstantial claim
Findingown data
Co-treatment with B. monnieri reduces expression of ERP57 and CHOP back to control levels in HT22 cells.
“co-treatment with B.monnieri reduces the expression of these proteins back to control levels”
Quote in the paperSubstantial claim
Findingown data
One-way ANOVA showed a significant difference between the mean cellular fluorescence of control, 5 mM glutamate, and 5 mM glutamate plus 10 μg/ml hexane extract of B. monnieri.
“One-way ANOVA analysis showed that there was a significant difference between the mean cellular fluorescence of the control, 5 mM glutamate, and 5 mM glutamate plus 10 μg/ml Hexane extract of B.monnieri.”
Quote in the paperSubstantial claim
Findingown data
At 3 μg/ml, B. monnieri hexane extract showed no significant difference in C. elegans survival compared to control (Log-rank Mantel-Cox test p = 0.09, n=3).
“At lower doses (3 μg/ml shown in Fig. 7a) B.monnieri Hexane extract showed no significant difference between the control and B.monnieri treated using Log-rank (Mantel-Cox) test (p = 0.09, n = 3).”
Quote in the paperNot a substantial claim
Findingown data
Glutamate caused a doubling in expression of ERP57 in HT22 cells.
“glutamate caused a doubling in the expression of ERP57”
Quote in the paperSubstantial claim
Findingown data
Dunnett's post hoc test showed a significant difference between 5 mM glutamate and 5 mM glutamate plus 10 μg/ml hexane extract of B. monnieri (P < 0.0001, n=3).
“Using Dunnet’s post hoc test for the comparison between glutamate 5 mM and glutamate plus 10 μg/ml hexane extract of B.monnieri P < 0.0001 (n = 3)”
Quote in the paperSubstantial claim
Findingown data
There was no significant difference between control and 5 mM glutamate plus 10 μg/ml hexane extract of B. monnieri in mean cellular fluorescence.
“while there was no significant difference between the control and glutamate plus 10 μg/ml hexane extract of B.monnieri.”
Quote in the paperSubstantial claim
Findingown data
The difference in average fluorescence per worm between control and B. monnieri treated C. elegans was statistically significant (unpaired t-test P = 0.0006, n=3).
“This difference was statistically significant (unpaired t-test P = 0.0006 n = 3).”
Quote in the paperSubstantial claim
Findingown data
Median survival time of C. elegans treated with B. monnieri hexane extract was significantly higher than control for some concentrations (ANOVA with Dunnett's post hoc test p < 0.05).
“* Median survival time in days significantly higher than control using ANOVA with Dunnett’s post hoc test p < 0.05.”
Quote in the paperSubstantial claim
Findingown data
Maximum survival time of C. elegans treated with B. monnieri hexane extract was significantly higher than control for some concentrations (ANOVA with Dunnett's post hoc test p < 0.05).
“* Maximum survival time in days significantly higher than control using ANOVA with Dunnett’s post hoc test p < 0.05.”
Quote in the paperSubstantial claim
Findingown data
Mean fluorescence per worm was 14.4 ± 1.7 for control and 5.6 ± 1.1 for treatment with 10 μg/ml B. monnieri.
“Mean fluorescence/worm ± SEM (AU) was 14.4 ± 1.7 for the control and 5.6 ± 1.1 for treatment with 10 μg/mll Bacopa monnieri.”
Quote in the paperSubstantial claim
Findingown data
Median survival time of C. elegans treated with B. monnieri hexane extract was significantly lower than control for some concentrations (ANOVA with Dunnett's post hoc test p < 0.05).
“§ Median survival time in days significantly lower than control using ANOVA with Dunnett’s post hoc test p < 0.05.”
Quote in the paperNot a substantial claim
Findingown data
B. monnieri extract could extend the median and maximum lifespan of C. elegans.
“In this study we demonstrated that B.monnieri extract could extend the median and maximum lifespan of the C. elegans”
Quote in the paperSubstantial claim
Findingown data
In this study, mitochondrial damage/leakage and build-up of ROS in cultured neurons can be prevented by treatment with hexane extract of B. monnieri.
“In this study we have shown that mitochondrial damage/leakage and the build-up of ROS in cultured neurons can be prevented by treatment with the hexane extract of B.monnieri.”
Quote in the paperSubstantial claim
Interpretationown data
The mechanism by which Bacopa monnieri protects against glutamate toxicity involves reduction of ROS production and prevention of mitochondrial and ER stress.
“We found that the mechanism involves the reduction of ROS production and the prevention of mitochondrial and ER stress.”
Quote in the paperSubstantial claim
Interpretationown data
Bacopa monnieri can prolong the healthy lifespan of Caenorhabditis elegans.
“Furthermore, it can prolong the healthy lifespan of C.elegans”
Quote in the paperSubstantial claim
Premisecitation
C. elegans intestinal autofluorescence is an indicator of health in terms of aging and oxidative stress in many species, including C. elegans.
“C.elegans intestinal autofluorescence is an indicator of health in terms of aging and oxidative stress in many species, including C.elegans.”
Quote in the paperSubstantial claim0 of 1 sources checked
- cites [25] Wang, 2016, Peptides from sesame cake extend healthspan of Caenorhabditis elegans via upregulation of skn-1 and inhibition of intracellular ROS levels, Exp Gerontol source
  Not checked in the free run.
Premisecitation
Cucurbitacin E and I are less likely to be protective compounds since they appear to induce cell death in various cancers.
“Curbitacini E & I, are less likely to be protective compounds since they apear to induce cell death in various cancers29,30”
Quote in the paperNot a substantial claim0 of 2 sources checked
- cites [29] Sörensen, 2012, The natural product cucurbitacin E inhibits depolymerization of actin filaments, ACS Chem Biol source
  Not checked in the free run.
- cites [30] Duncan, 1996, Cucurbitacin E-induced disruption of the actin and vimentin cytoskeleton in prostate carcinoma cells, Biochem Pharmacol source
  Not checked in the free run.
Premisecitation
There is no study related to glutamate toxicity in HT-22 cells.
“in particular, there is no study related to glutamate toxicity in HT-22 cells, a common in vitro model for measuring glutamate-induced ROS production in neuronal cells.22”
Quote in the paperSubstantial claim0 of 1 sources checked
- cites [22] Schubert, 1992, Growth factors and vitamin E modify neuronal glutamate toxicity, Proc Natl Acad Sci source
  Not checked in the free run.
Premisecitation
There have been very few in vitro studies regarding Bacopa monnieri and its effects on cultured neuronal cells.
“There have been very few in vitro studies regarding B.monnieri and its effects on cultured neuronal cells”
Quote in the paperSubstantial claim
Interpretationreasoning
The difference between ethanol, DCM and hexane extracts is likely due to compounds other than bacoside A constituents, due to lack of bacoside A compounds in dichloromethane and hexane extracts.
“This difference between the ethanol, DCM and hexane extracts is likely due to compounds other than the bacoside A constituents, due to the lack of bacoside A compounds in the dichloromethane and hexane extracts.”
Quote in the paperSubstantial claim
Interpretationreasoning
Bacopa monnieri has the potential for therapeutic and preventative use in neurodegenerative disease.
“indicating that B.monnieri the potential for therapeutic and preventative use in neurodegenerative disease.”
Quote in the paperSubstantial claim
- Rests on an assumption: Lifespan and aging effects of Bacopa monnieri in Caenorhabditis elegans are indicative of therapeutic and preventative potential in human neurodegenerative disease.
Interpretationreasoning
Less polar compounds such as monnieraside III and Plantainoside B are possible candidates for the protection seen in this study.
“Less polar compounds shown in suplamentary Fig. 1 such as monnieraside III and Plantainoside B are possible candidates for the protection seen in this study.”
Quote in the paperNot a substantial claim
Premisecitation
Limpeanchob et al. (2008) used at least 10 times higher concentrations of B. monnieri than the present study.
“Limpeanchob et al., (2008) used at least 10 times higher concentrations of B.monnieri”
Quote in the paperNot a substantial claim0 of 1 sources checked
- cites [27] Limpeanchob, 2008, Neuroprotective effect of Bacopa monnieri on beta-amyloid-induced cell death in primary cortical culture, J Ethnopharmacol source
  Not checked in the free run.
Premisecitation
Bacopa monnieri supplementation improves Alzheimer's like characteristics in mouse models, such as colchicine treatment, olfactory bulbectomy, and APP(SWE) expressing mice.
“and improves the Alzheimer’s like characteristics in mouse models, such as colchicine treatment,13 olfactory bulbectomy,14 and APP(SWE) expressing mice.14”
Quote in the paperNot a substantial claim0 of 2 sources checked
- cites [13] Saini, 2012, Neuroprotective effects of Bacopa monnieri in experimental model of dementia, Neurochem Res source
  Not checked in the free run.
- cites [14] Le, 2013, Bacopa monnieri ameliorates memory deficits in olfactory bulbectomized mice: possible involvement of glutamatergic and cholinergic systems, Neurochem Res source
  Not checked in the free run.
Interpretationcitation
B. monnieri's ability to protect mitochondria from ROS insult may explain why studies have shown positive behavioral changes in Alzheimer's experimental models but no change in size of plaques, even when amyloid β load is reduced.
“B.monnieri’s ability to protect the mitochondria from ROS insult may explain why studies have shown positive behavioral changes in Alzheimer’s experimental models but no change in the size of plaques, even when amyloid β load is reduced.13”
Quote in the paperNot a substantial claim0 of 1 sources checked
- cites [13] Saini, 2012, Neuroprotective effects of Bacopa monnieri in experimental model of dementia, Neurochem Res source
  Not checked in the free run.
Premisecitation
Reduction in glucose metabolism in the brain is common in Alzheimer's patients.
“it is well documented that a reduction in glucose metabolism in the brain is common in Alzheimer’s patients.36, 37, 38”
Quote in the paperSubstantial claim0 of 3 sources checked
- cites [36] Ding, 2013, Longitudinal progression of cognitive decline correlates with changes in the spatial pattern of brain 18F-FDG PET, J Nucl Med source
  Not checked in the free run.
- cites [37] Yamane, 2014, Visual-statistical interpretation of 18F-FDG-PET images for characteristic alzheimer patterns in a multicenter study: inter-rater concordance and relationship to automated quantitative evaluation, Am J Neuroradiol source
  Not checked in the free run.
- cites [38] Reiman, 2009, Associations between cognitive, functional, and FDG-PET measures of decline in AD and MCI, Neurobiol Aging
  Not checked in the free run.
Interpretationreasoning
This study provides more evidence that B. monnieri can potentially be of use as a therapeutic agent in aged patients.
“This study provides more evidence that B.monnieri, can potentially be of use as a therapeutic agent in aged patients.”
Quote in the paperSubstantial claim

May combine two claims. 31
Classified as probably not a substantial claim. 22
The quote only partly states this claim. 1

May combine two claims.
Findingown data
Bacopa monnieri hexane extract, but not ethanol extract, prevented the toxicity of 5 mM glutamate in HT-22 cells.
“B.monnieri hexane extract (but not ethanol extract) prevented the toxicity of 5 mM glutamate in HT-22 cells.”
- Also stated in: § The effect of B.monnieri extracts, on glutamate toxicity in HT-22 cells · plain
  “The B.monnieri hexane extract protected the HT-22 cells from the toxicity of 5 mM glutamate.”
Quote in the paperSubstantial claim
May combine two claims.
Findingown data
Mean cellular fluorescence of HT22 cells stained with H2DCFDA was 10.93 ± 1.85 for control, 61.66 ± 17.60 for 5 mM glutamate treated, and 7.55 ± 1.20 for 5 mM glutamate plus 10 μg/ml hexane extract of B. monnieri (n=3).
“The mean cellular fluorescences were 10.93 ± 1.85, 61.66 ± 17.60 and 7.55 ± 1.20 for the control, 5 mM glutamate treated, and 5 mM glutamate plus 10 μg/ml Hexane extract of B.monnieri respectively (n = 3).”
Quote in the paperSubstantial claim
May combine two claims.
Findingown data
5 mM glutamate causes mitochondrial stress and leakage in HT-22 cells, as indicated by diffuse rhodamine123 staining.
“Fig. 4b shows a representative confocal micrograph of HT-22 cells treated with 5 mM glutamate, and stained with rhodamine123; here, the staining is more diffuse, suggesting that the mitochondria are stressed and leaking.”
Quote in the paperSubstantial claim
May combine two claims.
Findingown data
HT22 cells treated with 10 μg/ml B. monnieri hexane extract and 5 mM glutamate showed fluorescence more in line with control cells and significantly reduced fluorescence compared to 5 mM glutamate alone (ANOVA followed Dunnett's post hoc test P < 0.05, n=4).
“cells treated with 10 μg/ml B.monnieri Hexane extract and 5 mM glutamate, showed fluorescence more in line with control cells, and with statistically significant reduction in fluorescence compared to 5 mM glutamate alone (ANOVA followed Dunnett’s post hoc test P < 0.05 (n = 4))”
Quote in the paperSubstantial claim
May combine two claims.
Findingown data
The average fluorescence per worm for control C. elegans was 15.4 ± 1.7 (n=3) and for B. monnieri treated worms was 5.6 ± 1.1 (n=3).
“After three independent experiments, the average fluorescence per worm for the control was 15.4 ± 1.7 (n = 3) fluorescence units, and for the B.monnieri treated worms 5.6 ± 1.1 (n = 3).”
Quote in the paperSubstantial claim
May combine two claims.
Findingown data
At higher doses, there was an apparent reduction in C. elegans survival, but it was not statistically significant (Log-rank Mantel-Cox P = 0.26, n=3).
“At higher doses, there was an apparent reduction in C. elegans survival, although analysis showed it not to be statistically significant, Log-rank (Mantel-Cox) test (P = 0.26 n = 3).”
Quote in the paperNot a substantial claim
Probably not a substantial claim: may not pass S3 (load-bearing).
Findingown data
Bacopaside II was the most abundant compound from those measured in Bacopa monnieri (0.38 ± 0.00001 mg/g).
“Bacopaside II was found to be the most abundant compound from those measured (0.38 ± 0.00001 mg/g).”
Quote in the paperNot a substantial claim
May combine two claims.
Findingown data
The present study saw significant protection against glutamate toxicity with DCM and hexane extracts at much lower concentrations than in the previous study.
“However, we did see significant protection against glutamate-toxicity with the DCM and hexane extracts and furthermore, at much lower concentrations than in the previous study.”
Quote in the paperSubstantial claim
The quote only partly states this claim.
Findingown data
Maximum survival time of C. elegans treated with B. monnieri hexane extract was significantly lower than control for some concentrations (ANOVA with Dunnett's post hoc test p < 0.05).
“§ Maximum survival time in days significantly lower than control using ANOVA with Dunnett’s post hoc test p < 0.05.”
Quote in the paperNot a substantial claim
May combine two claims.
Interpretationown data
Bacopa monnieri prevents mitochondrial and oxidative stress in cultured cells.
“In conclusion, B.monnieri prevents mitochondrial, and oxidative stress in the cultured cells.”
Quote in the paperSubstantial claim
Probably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation
A meta-analysis of 9 randomized controlled trials concluded that Bacopa monnieri has the potential to improve cognitive function, particularly in the speed of attention.
“A meta-analysis of 9 randomized controlled trials concluded that B.monnieri has the potential to improve cognitive function, particularly in the speed of attention; however, the authors based their conclusions on nine small scale studies and suggest that a large scale study is required.18”
Quote in the paperNot a substantial claim0 of 1 sources checked
- cites [18] Norman Scholfield, 2013, Meta-analysis of randomized controlled trials on cognitive effects of Bacopa monnieri extract, J Ethnopharmacol
  Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductnone given
In each independent lipofuscin imaging experiment, at least ten worms were randomly selected and photographed for analysis.
“In each independent experiment, at least ten worms were randomly selected and photographed for analysis.”
Quote in the paperNot a substantial claim
May combine two claims.
Premisecitation
Increased ROS can have detrimental effects in neuronal cells, resulting in endoplasmic reticulum stress as proteins are miss-folded and mitochondrial stress and leakage, which can cause apoptosis and neurodegeneration.
“Increased ROS can have many detrimental effects in neuronal cells, resulting in endoplasmic reticulum stress, as proteins are miss-folded4 and mitochondrial stress and leakage, which can cause apoptosis and neurodegeneration.5”
Quote in the paperSubstantial claim0 of 2 sources checked
- cites [4] Kudo, 2008, A molecular chaperone inducer protects neurons from ER stress, Cell Death Differ source
  Not checked in the free run.
- cites [5] Cadenas, 2000, Mitochonrial free radical generation,oxidative stress,and aging, Free Radic Biol Med source
  Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Study design and conductown data
Each treatment in the trypan blue exclusion assay was carried out in triplicate, and data shown are from at least three independent experiments.
“Each treatment was carried out in triplicate; data shown are from at least three independent experiments, and represented as mean plus and minus the SEM.”
Quote in the paperNot a substantial claim
Classified as probably not a substantial claim. May combine two claims.
Premisecitation
Lipofuscin consists of oxidized lipid peroxide and protein residues that increase within cells in a manner dependent on age.
“Lipofuscin consists of oxidized lipid peroxide and protein residues that increase within cells in a manner dependent on age26⁠.”
Quote in the paperSubstantial claim0 of 1 sources checked
- cites [26] Jyotsna, 2015, Free Radic Res
  Not checked in the free run.
Probably not a substantial claim: may not pass S4 (based).
Premisenone given
There has been no study on the effect of Bacopa monnieri on the life span and aging of wild type Caenorhabditis elegans.
“Furthermore, to our knowledge, there has been no study on the effect of B.monnieri on the life span and aging of wild type Caenorhabditis elegans.”
Quote in the paperNot a substantial claim
May combine two claims.
Interpretationown data
Hexane and DCM extracts of B. monnieri protect against glutamate toxicity in HT-22 cells, whereas the ethanol extract does not.
“In this study, we have shown that the Hexane and DCM extracts of B.monnieri protect against glutamate toxicity in HT-22 cells, whereas the ethanol extract does not.”
Quote in the paperSubstantial claim
May combine two claims.
Premisecitation
High concentrations of glutamate can block cystine uptake into cells via the cysteine/glutamate anti-porter, resulting in depletion of intracellular cysteine and glutathione, and subsequently increasing ROS production.
“High concentrations of glutamate can block cystine uptake into the cells via the cysteine/glutamate anti-porter, which results in depletion of intracellular cysteine and glutathione (a cellular anti-oxidant), and subsequently, increasing ROS production.3”
Quote in the paperSubstantial claim0 of 1 sources checked
- cites [3] Suh, 2007, Curcumin attenuates glutamate-induced HT22 cell death by suppressing MAP kinase signaling, Mol Cell Biochem source
  Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation
Animal studies found that Bacopa monnieri improves memory in the form of pain avoidance in rats.
“and improve memory in the form of pain avoidance in rats.11”
Quote in the paperNot a substantial claim0 of 1 sources checked
- cites [11] Singh, 1982, Effect of Bacopa monniera Linn. (Brāhmi) extract on avoidance responses in rat, J Ethnopharmacol source
  Not checked in the free run.
Probably not a substantial claim: may not pass S4 (based). May combine two claims.
Interpretationnone given
Non-polar compounds more likely to be extracted by hexane will have a greater ability to be absorbed by the gut and cross the blood brain barrier, making them more useful in treatment of neurodegenerative diseases.
“since the non-polar copounds more likely to be extracted by hexane, will have a greater ability to be absorbed by the gut, and cross the blood brain barrier, making them more usefull in the treatment of neurodegernative diseases.”
Quote in the paperNot a substantial claim
May combine two claims.
Premisecitation
Glutamate toxicity is a factor in the pathology of neurodegeneration in aging patients, with the toxicity mediated through multiple pathways, one being a reactive oxygen species (ROS) mechanism.
“Glutamate toxicity is thought to be a factor in the pathology of neurodegeneration in aging patients, with the toxicity mediated through multiple pathways, one being a reactive oxygen species (ROS) mechanism.”
Quote in the paperSubstantial claim0 of 1 sources checked
- cites [3] Suh, 2007, Curcumin attenuates glutamate-induced HT22 cell death by suppressing MAP kinase signaling, Mol Cell Biochem source
  Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation
Bacopa monnieri supplementation improves symptoms of paraquat treatment of mice in a Parkinson's disease model.
“Furthermore, B.monnieri supplementation improves symptoms of paraquat treatment of mice in a Parkinson’s disease model12”
Quote in the paperNot a substantial claim0 of 1 sources checked
- cites [12] Krishna, 2019, Bacopa monnieri supplements offset paraquat-induced behavioral phenotype and brain oxidative pathways in mice, Cent Nerv Syst Agents Med Chem source
  Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Study design and conductnone given
Bacopa monnieri was identified by an expert botanist and assigned voucher number 016299 (BUC).
“B.monnieri was identified by, expert botanist, Nirun Vipunngeun, College of Pharmacy, Rangsit University, Thailand, voucher number 016299 (BUC).”
Quote in the paperNot a substantial claim
Probably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductnone given
All lifespan experiments were carried out in triplicate.
“All the experiments were carried out in triplicate.”
Quote in the paperNot a substantial claim
Probably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductown data
The identity of the Bacopa monnieri extract was confirmed by reversed-phase HPLC.
“B.monnieri extract was analyzed by reversed-phase HPLC to confirm the identity of the herb.”
Quote in the paperNot a substantial claim
May combine two claims.
Interpretationown data
B. monnieri hexane extract reduces mitochondrial stress and increases mitochondrial membrane integrity in glutamate-treated HT22 cells.
“which would suggest that B.monnieri is reducing mitochondrial stress, and increasing the mitochondrial membrane integrity.”
Quote in the paperSubstantial claim
Probably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation
Limpeanchob et al. (2008) showed that ethanol extract of B. monnieri at 100 μg/ml could reduce ROS content of control cells (not stressed).
“Limpeanchob et al. (2008) did show that the ethanol extract of B.monnieri at 100 μg/ml could reduce the ROS content of the control cells (i.e., not stressed).”
Quote in the paperNot a substantial claim0 of 1 sources checked
- cites [27] Limpeanchob, 2008, Neuroprotective effect of Bacopa monnieri on beta-amyloid-induced cell death in primary cortical culture, J Ethnopharmacol source
  Not checked in the free run.
Classified as probably not a substantial claim. May combine two claims.
Study design and conductnone given
All results come from 3 or more independent experiments and are represented as means ± SEM.
“All results come from 3 or more independent experiments and are represented as means ± SEM.”
Quote in the paperSubstantial claim
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Premisecitation
Bacopa monnieri is a medicinal herb found throughout Asia and is known for its memory-enhancing effects.
“B.monnieri is a medicinal herb found throughout Asia and is known for its memory-enhancing effects.6”
Quote in the paperNot a substantial claim0 of 1 sources checked
- cites [6] Kean, 2017, Systematic overview of bacopa monnieri (L.) wettst. Dominant poly-herbal formulas in children and adolescents, Medicine
  Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation
Animal studies found that Bacopa monnieri can reverse the amnesic effects of scopolamine in mice.
“Animal studies have found that the herb can reverse the amnesic effects of scopolamine9,10 in mice”
Quote in the paperNot a substantial claim0 of 2 sources checked
- cites [9] Saraf, 2010, Bacopa monniera attenuates scopolamine-induced impairment of spatial memory in mice, Evidence-Based Complement Altern Med
  Not checked in the free run.
- cites [10] Saraf, 2010, Scopolamine induced amnesia is reversed by bacopa monniera through participation of kinase-CREB pathway, Neurochem Res source
  Not checked in the free run.
May combine two claims.
Premisecitation
Previous work by Limpeanchob et al. (2008) found that ethanol extract of B. monnieri did not prevent glutamate toxicity but did protect against effects of amyloid β in primary cortical cells.
“Previous work by Limpeanchob et al., (2008)27 investigated the ethanol extract concerning glutamate-toxicity in primary cortical cells, found that the ethanol extract did not prevent glutamate toxicity, but did protect against the effects of amyloid β.”
Quote in the paperSubstantial claim0 of 1 sources checked
- cites Limpeanchob et al., (2008)27 source
  Not checked in the free run.
May combine two claims.
Interpretationown data
Glutamate treatment (5 mM) causes significant increase in mitochondrial stress and loss of mitochondrial membrane integrity in HT22 cells.
“indicating a significant increase in mitochondrial stress, and a loss of mitochondrial membrane integrity, in glutamate treated cells”
Quote in the paperSubstantial claim
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Premisecitation
Apigenin is found in B. monnieri and hexane extracts of other herbs, and has potential neuroprotective effects with relation to glutamate release.
“One other possible compound found in B.monnieri8 and hexane extracts of other herbs is apigenin,31 and has potential neuroprotective effects with realtion to glutamate release.32”
Quote in the paperNot a substantial claim0 of 3 sources checked
- cites [8] Deepak, 2005, Quantitative determination of the major saponin mixture bacoside A in Bacopa monnieri by HPLC, Phytochem Anal An Int J Plant Chem Biochem Tech
  Not checked in the free run.
- cites [31] Mallhi, 2018, Determination of phytoconstituents of n-hexane extract of leaves of Morus nigra and evaluation of their effects on biochemical and histopathological parameters in paracetamol intoxicated mice liver, Brazilian J Pharm Sci
  Not checked in the free run.
- cites [32] Brimson, 2019, Clerodendrum petasites S. Moore: the therapeutic potential of phytochemicals, hispidulin, vanillic acid, verbascoside, and apigenin, Biomed Pharmacother source
  Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Premisecitation
The % w/w of bacoside A compounds in the ethanol extract were relatively lower than other studies, but the ratio of compounds was comparable to previous studies.
“While the % w/w of bacoside A compounds in the ethanol exract were relatively lower than other studies the ratio of compounds was comparable to previous studies.27,28”
Quote in the paperNot a substantial claim0 of 2 sources checked
- cites [27] Limpeanchob, 2008, Neuroprotective effect of Bacopa monnieri on beta-amyloid-induced cell death in primary cortical culture, J Ethnopharmacol source
  Not checked in the free run.
- cites [28] Ganzera, 2004, Separation of the major triterpenoid saponins in Bacopa monnieri by high-performance liquid chromatography, Anal Chim Acta source
  Not checked in the free run.
May combine two claims.
Interpretationown data
The hexane extract (but not the ethanol extract) of B. monnieri protects cultured neuronal cells against ROS production induced by glutamate treatment and prevents mitochondrial stress.
“In summary, the hexane extract (but not the ethanol extract) of B.monnieri protects cultured neuronal cells against the ROS production induced by glutamate treatment and prevents mitochondrial stress.”
Quote in the paperSubstantial claim
May combine two claims.
Interpretationreasoning
The protection of mitochondria in surviving neurons and ability to maintain glucose metabolism at higher levels may explain B. monnieri's effectiveness in improving memory in Alzheimer's patients and healthy patients alike.
“The protection of mitochondria in the surviving neurons, and the ability to maintain glucose metabolism at higher levels may explain B.monnieri’s effectiveness in improving memory in Alzheimer’s patients and healthy patients alike.”
Quote in the paperSubstantial claim
May combine two claims.
Interpretationown data
From our results, B. monnieri extract can extend the lifespan and reduce the effects of aging in C. elegans, resulting in younger appearing worms compared to control.
“From our results it would appear that the B.monnieri extract can extend the lifespan and reduce the effects of aging in C. elegans, resulting in younger appearing worms when compared to the control.”
Quote in the paperSubstantial claim
May combine two claims.
Premisecitation
C. elegans intestinal autofluorescence is an indicator of the individual's apparent age and thus linked to the "health-span" of the population.
“C. elegans intestinal autofluorescence is an indicator of the individual’s apparent age and thus linked to the “health-span” of the population.39,40”
Quote in the paperSubstantial claim0 of 2 sources checked
- cites [39] Pincus, 2016, Autofluorescence as a measure of senescence in C. elegans: look to red, not blue or green, Aging (Albany NY) source
  Not checked in the free run.
- cites [40] Forge, 1989, Nematode autofluorescence and its sse as an indicator of viability, J Nematol
  Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Premisecitation
Neuronal cells in the brain have a particularly high metabolism, using 20% and 25% of total consumed oxygen and glucose, making neurons and their mitochondria particularly susceptible to ROS damage.
“Neuronal cells in the brain have a particularly high metabolism, using 20% and 25% of the total consumed oxygen and glucose,34 this makes neurons and their mitochondria particularly susceptible to ROS damage.”
Quote in the paperNot a substantial claim0 of 1 sources checked
- cites [34] Bélanger, 2011, Brain energy metabolism: focus on Astrocyte-neuron metabolic cooperation, Cell Metabol source
  Not checked in the free run.
Classified as probably not a substantial claim. May combine two claims.
Premisecitation
Mitochondria are a source of free radicals and ROS, and are susceptible to ROS damage.
“The mitochondria, are a source of free radicals and ROS, as well as being susceptible to ROS damage.33”
Quote in the paperSubstantial claim0 of 1 sources checked
- cites [33] Hirai, 2001, Mitochondrial abnormalities in Alzheimer’s disease, J Neurosci source
  Not checked in the free run.
Probably not a substantial claim: may not pass S2 (checkable).
Interpretationreasoning
The fact that ethanol extract and hexane extract had differing efficacies could lead to identification of new lead compounds for pharmaceutical study.
“Furthermore, the fact that the ethanol extract and the hexane extract had differing efficacies could lead to the identification of new lead compounds for pharmaceutical study.”
Quote in the paperNot a substantial claim
May combine two claims.
Interpretationreasoning
Bacoside A compounds may be a defining feature of B. monnieri, but they are not necessarily the only bioactive compounds found in the herb.
“This suggests that while the bacoside A compounds may be a defining feature of B.monnieri, they are not necessarily the only bioactive compounds found in the herb.”
Quote in the paperNot a substantial claim
May combine two claims.
Interpretationnone given
The extract can extend the lifespan of C. elegans while maintaining a younger like phenotype in aged worms.
“Furthermore, the extract can extend the lifespan of the model organism, C.elegans, while maintaining a younger like phenotype in the aged worms.”
Quote in the paperSubstantial claim
Probably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation
Mitochondrial damage in neurons is considered a risk factor for neurodegenerative diseases such as Alzheimer's and Parkinson's disease.
“Mitochondrial damage in neurons is considered to be a risk factor for neurodegenerative diseases such as Alzheimer’s and Parkinson’s disease.33,35”
Quote in the paperNot a substantial claim0 of 2 sources checked
- cites [33] Hirai, 2001, Mitochondrial abnormalities in Alzheimer’s disease, J Neurosci source
  Not checked in the free run.
- cites [35] Wang, 2014, Oxidative stress and mitochondrial dysfunction in Alzheimer’s disease, Biochim Biophys Acta (BBA) - Mol Basis Dis source
  Not checked in the free run.

44 of 44

The paper relies on this without support.
Lifespan and aging effects of Bacopa monnieri in Caenorhabditis elegans are indicative of therapeutic and preventative potential in human neurodegenerative disease.
“indicating that B.monnieri the potential for therapeutic and preventative use in neurodegenerative disease.”
The paper relies on this without support.
This is the first study to investigate the effects of glutamate-induced toxicity with different solvents used to extract B. monnieri.
“This is the first study to investigate the effects of glutamate-induced toxicity with different solvents used to extract the contents of the herb.”
The paper relies on this without support.
Glutamate-induced cell death in cortical neurons is likely caused by excitotoxicity, which is a different mechanism from that in HT-22 cells.
“the glutamate-induced cell death in cortical neurons is likely to be caused by excitotoxicity, which is different mechanism to that in HT-22 cells”
The paper relies on this without support.
In HT-22 cells, glutamate-induced cell death is caused by a block on the production of glutathione, resulting in ROS build up.
“which is a block on the production of glutathione, resulting in ROS build up.”
The paper relies on this without support.
B. monnieri has previously been shown to have positive outcomes in the improvement of memory in both normal and aged/Alzheimer’s subjects.
“B.monnieri has previously been shown to have positive outcomes in the improvement of memory in both normal and aged/Alzheimer’s subjects.”
The paper relies on this without support.
Intact and functioning mitochondria are essential for proper neuronal function.
“Intact and functioning mitochondria are essential for proper neuronal function”

6 of 6

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?

Bacopa monnieri reduces oxidative and mitochondrial stress, enhances lifespan, and reduces aging symptoms in Caenorhabditis elegans.
“In this context, in recent research, BM shows significant results by reducing oxidative and mitochondrial stress and enhancing life span while reducing aging symptoms in Caenorhabditis elegans.”
From Unlocking the Therapeutic Potential of Medicinal Plants for Alzheimer’s Disease: Preclinical to Clinical Trial Insights (2023) · cites this paper as [128]
Mechanism of Bacopa monnieri hexane extract includes lowering ROS generation and preventing ER and mitochondrial stress.
“The mechanism behind this includes lowering ROS generation and preventing ER and mitochondrial stress [128].”
From Unlocking the Therapeutic Potential of Medicinal Plants for Alzheimer’s Disease: Preclinical to Clinical Trial Insights (2023) · cites this paper as [128]
Hexane extract of Bacopa monnieri (but not ethanol extract) reduces toxicity of 5 mM glutamate in HT-22 cells.
“The results demonstrate that in HT-22 cells, the hexane extract of BM (but not an ethanol extract) reduced the toxicity of 5 mM glutamate.”
From Unlocking the Therapeutic Potential of Medicinal Plants for Alzheimer’s Disease: Preclinical to Clinical Trial Insights (2023) · cites this paper as [128]

Self-check

Percentages, sums and reported differences re-computed from the paper’s own numbers.

Each contradiction between two passages listed here was confirmed by an independent check.

Conflicting control-worm fluorescence values
Minor
The results report control-worm fluorescence as 15.4 ± 1.7, while the Fig. 8 caption reports 14.4 ± 1.7 for the same measurement.
Conflicting sample counts for trypan blue results
Minor
The text gives n = 4 for the trypan blue comparison, but the Fig. 2 caption gives n = 3.
ROS staining images assigned to two figures
Minor
The ROS results first identify the carboxy-H2DCFDA staining images as Fig. 7a-c, then identify them as Fig. 5a-c. The Fig. 5 caption describes those images, while Fig. 7 concerns worm survival.
Conflicting sample counts for rhodamine fluorescence
Minor
The Fig. 4 caption gives n = 3 for the control-versus-glutamate fluorescence comparison, whereas the results text gives n = 4.

4 of 4

Ageing populations: the challenges ahead
Christensen, K., Doblhammer, G., Rau, R., & Vaupel, J. W. (2009) · The Lancet · doi:10.1016/s0140-6736(09)61460-4 · Cited by 3,764 · see citing works
Mitochondrial free radical generation, oxidative stress, and aging
Cadenas, E., & Davies, K. J. A. (2000) · Free Radical Biology and Medicine · doi:10.1016/s0891-5849(00)00317-8 · Cited by 3,010 · see citing works
Brain energy metabolism: focus on Astrocyte-neuron metabolic cooperation
Bélanger, M., Allaman, I., & Magistretti, P. J. (2011) · Cell Metabolism · doi:10.1016/j.cmet.2011.08.016 · Cited by 2,313 · see citing works
Mitochondrial abnormalities in Alzheimer's disease
Hirai, K., Aliev, G., Nunomura, A., Fujioka, H., Russell, R. L., Atwood, C. S., Johnson, A. B., Kress, Y., Vinters, H. V., Tabaton, M., et al. (2001) · The Journal of Neuroscience · doi:10.1523/jneurosci.21-09-03017.2001 · Cited by 1,367 · see citing works
Oxidative stress and mitochondrial dysfunction in Alzheimer’s disease
Wang, X., Wang, W., Li, L., Perry, G., Lee, H.-G., & Zhu, X. (2014) · Biochimica et Biophysica Acta (BBA) - Molecular Basis of Disease · doi:10.1016/j.bbadis.2013.10.015 · Cited by 1,357 · see citing works
Associations between cognitive, functional, and FDG-PET measures of decline in AD and MCI
Landau, S. M., Harvey, D., Madison, C. M., Koeppe, R. A., Reiman, E. M., Foster, N. L., Weiner, M. W., & Jagust, W. J. (2011) · Neurobiology of Aging · doi:10.1016/j.neurobiolaging.2009.07.002 · Cited by 750 · see citing works
Chronic Effects of Brahmi (Bacopa monnieri) on Human Memory
Roodenrys, S. (2002) · Neuropsychopharmacology · doi:10.1016/s0893-133x(01)00419-5 · Cited by 310 · see citing works
A molecular chaperone inducer protects neurons from ER stress
Kudo, T., Kanemoto, S., Hara, H., Morimoto, N., Morihara, T., Kimura, R., Tabira, T., Imaizumi, K., & Takeda, M. (2008) · Cell Death & Differentiation · doi:10.1038/sj.cdd.4402276 · Cited by 250 · see citing works
World Alzheimer Report 2013: Journey of Caring: An analysis of long-term care for dementia
Guerchet, M., Prince, M. J., & Prina, M. A. (2013) · Cited by 210 · see citing works
Neuroprotective effect of Bacopa monnieri on beta-amyloid-induced cell death in primary cortical culture
Limpeanchob, N., Jaipan, S., Rattanakaruna, S., Phrompittayarat, W., & Ingkaninan, K. (2008) · Journal of Ethnopharmacology · doi:10.1016/j.jep.2008.07.039 · Cited by 206 · see citing works
Growth factors and vitamin E modify neuronal glutamate toxicity
Schubert, D., Kimura, H., & Maher, P. (1992) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.89.17.8264 · Cited by 172 · see citing works
Meta-analysis of randomized controlled trials on cognitive effects of Bacopa monnieri extract
Kongkeaw, C., Dilokthornsakul, P., Thanarangsarit, P., Limpeanchob, N., & Norman Scholfield, C. (2014) · Journal of Ethnopharmacology · doi:10.1016/j.jep.2013.11.008 · Cited by 166 · see citing works
Quantitative determination of the major saponin mixture bacoside A in <i>Bacopa monnieri</i> by HPLC
Deepak, M., Sangli, G. K., Arun, P. C., & Amit, A. (2005) · Phytochemical Analysis · doi:10.1002/pca.805 · Cited by 164 · see citing works
Effect of Bacopa monniera Linn. (Brāhmi) extract on avoidance responses in rat
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Autofluorescence as a measure of senescence in C. elegans: look to red, not blue or green
Pincus, Z., Mazer, T. C., & Slack, F. J. (2016) · Aging · doi:10.18632/aging.100936 · Cited by 153 · see citing works
Cucurbitacin E-induced disruption of the actin and vimentin cytoskeleton in prostate carcinoma cells
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Effects of 12-Week Bacopa monnieri Consumption on Attention, Cognitive Processing, Working Memory, and Functions of Both Cholinergic and Monoaminergic Systems in Healthy Elderly Volunteers
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Neuroprotective effects of Bacopa monnieri in experimental model of dementia
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Randomized controlled trial of standardized Bacopa monniera extract in age-associated memory impairment
Singh, H., Raghav, S., Dalal, P., Srivastava, J., & Asthana, O. (2006) · Indian Journal of Psychiatry · doi:10.4103/0019-5545.31555 · Cited by 94 · see citing works
Efficacy of Standardized Extract of Bacopa monnieri (Bacognize®) on Cognitive Functions of Medical Students: A Six‐Week, Randomized Placebo‐Controlled Trial
Kumar, N., Abichandani, L. G., Thawani, V., Gharpure, K. J., Naidu, M. U. R., & Ramana, G. V. (2016) · Evidence-based Complementary and Alternative Medicine · doi:10.1155/2016/4103423 · Cited by 94 · see citing works
Bacopa monniera Attenuates Scopolamine‐Induced Impairment of Spatial Memory in Mice
Saraf, M. K., Prabhakar, S. K., KHANDUJA, K. L., & Anand, A. (2010) · Evidence-based Complementary and Alternative Medicine · doi:10.1093/ecam/neq038 · Cited by 89 · see citing works
Neuropharmacological and cognitive effects of Bacopa monnieri (L.) Wettst – A review on its mechanistic aspects
Sukumaran, N. P., Amalraj, A., & Gopi, S. (2019) · Complementary Therapies in Medicine · doi:10.1016/j.ctim.2019.03.016 · Cited by 88 · see citing works
Bacopa monnieri ameliorates memory deficits in olfactory bulbectomized mice: Possible involvement of glutamatergic and cholinergic systems
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Curcumin attenuates glutamate-induced HT22 cell death by suppressing MAP kinase signaling
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The natural product cucurbitacin E inhibits depolymerization of actin filaments
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Effect of Bacopa monnieri on Cognitive functions in Alzheimer???s disease patients
Goswami, S., An, Saoji, V., Kumar, N., Thawani, V., Tiwari, M., & Thawani, M. (2011) · International journal of collaborative research on internal medicine & public health · Cited by 55 · see citing works
Efficacy and tolerability of bacomind® on memory improvement in elderly participants—A double blind placebo controlled study
Barbhaiya, H. C., Desai, R. P., Saxena, V. S., Pravina, K., Wasim, P., Geetharani, P., Allan, J. J., Venkateshw, K., & Amit, A. (2008) · Journal of Pharmacology and Toxicology · doi:10.3923/jpt.2008.425.434 · Cited by 54 · see citing works
Longitudinal Progression of Cognitive Decline Correlates with Changes in the Spatial Pattern of Brain <sup>18</sup> F-FDG PET
Shokouhi, S., Claassen, D., Kang, H., Ding, Z., Rogers, B., Mishra, A., Riddle, W. R., & Initiative, F. T. A. D. N. (2013) · Journal of Nuclear Medicine · doi:10.2967/jnumed.112.116137 · Cited by 54 · see citing works
Ultraviolet-A triggers photoaging in model nematode Caenorhabditis elegans in a DAF-16 dependent pathway
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Self-citation
Scopolamine induced amnesia is reversed by bacopa monniera through participation of kinase-CREB pathway
Saraf, M. K., Anand, A., & Prabhakar, S. (2010) · Neurochemical Research · doi:10.1007/s11064-009-0051-4 · Cited by 48 · see citing works
Visual-statistical interpretation of 18F-FDG-PET images for characteristic alzheimer patterns in a multicenter study: inter-rater concordance and relationship to automated quantitative evaluation
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Separation of the major triterpenoid saponins in Bacopa monnieri by high-performance liquid chromatography
Ganzera, M., Gampenrieder, J., Pawar, R. S., Khan, I. A., & Stuppner, H. (2004) · Analytica Chimica Acta · doi:10.1016/j.aca.2004.04.002 · Cited by 45 · see citing works
Clerodendrum petasites S. Moore: the therapeutic potential of phytochemicals, hispidulin, vanillic acid, verbascoside, and apigenin
Brimson, J. M., Onlamoon, N., Tencomnao, T., & Thitilertdecha, P. (2019) · Biomedicine & Pharmacotherapy · doi:10.1016/j.biopha.2019.109319 · Cited by 44 · see citing works
Self-citation
Ethanolic extract of Streblus asper leaves protects against glutamate-induced toxicity in HT22 hippocampal neuronal cells and extends lifespan of Caenorhabditis elegans
Prasansuklab, A., Meemon, K., Sobhon, P., & Tencomnao, T. (2017) · BMC Complementary and Alternative Medicine · doi:10.1186/s12906-017-2050-3 · Cited by 42 · see citing works
Self-citation
Peptides from sesame cake extend healthspan of Caenorhabditis elegans via upregulation of skn-1 and inhibition of intracellular ROS levels
Wang, Z., Ma, X., Li, J., & Cui, X. (2016) · Experimental Gerontology · doi:10.1016/j.exger.2016.07.001 · Cited by 39 · see citing works
Acacetin promotes healthy aging by altering stress response inCaenorhabditis elegans
Asthana, J., Mishra, B. N., & Pandey, R. (2016) · Free Radical Research · doi:10.1080/10715762.2016.1187268 · Cited by 35 · see citing works
Systematic Overview of Bacopa monnieri (L.) Wettst. Dominant Poly-Herbal Formulas in Children and Adolescents
Kean, J., Downey, L., & Stough, C. (2017) · Medicines · doi:10.3390/medicines4040086 · Cited by 29 · see citing works
Nematode autofluorescence and its use as an indicator of viability.
Forge, T. A., & MacGuidwin, A. E. (1989) · PubMed · Cited by 26 · see citing works
Bacopa monnieri supplements offset paraquat-induced behavioral phenotype and brain oxidative pathways in mice
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Investigating the Neuroprotective and Cognitive-Enhancing Effects of Bacopa monnieri: A Systematic Review Focused on Inflammation, Oxidative Stress, Mitochondrial Dysfunction, and Apoptosis
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Bacopa monnieri reduces Tau aggregation and Tau-mediated toxicity in cells
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Role of Medicinal Plants against Neurodegenerative Diseases
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Rhinacanthus nasutus “Tea” Infusions and the Medicinal Benefits of the Constituent Phytochemicals
Brimson, J. M., Prasanth, M. I., Malar, D. S., Brimson, S. J., & Tencomnao, T. (2020) · Nutrients · doi:10.3390/nu12123776 · Cited by 33 · see citing works
“Herbal remedies and traditional medicines for the treatment of cognitive decline have been used for centuries, and many have been shown to have positive effects on neurons in a laboratory setting, such as Curcuma longa [45], Mucuna pruriens [46], and Bacopa monnieri [47].”
BackgroundSelf-citation
Use of Bacopa monnieri in the Treatment of Dementia Due to Alzheimer Disease: Systematic Review of Randomized Controlled Trials
Basheer, A., Agarwal, A., Mishra, B., Gupta, A., Srivastava, M. V. P., Kirubakaran, R., & Vishnu, V. Y. (2022) · Interactive Journal of Medical Research · doi:10.2196/38542 · Cited by 33 · see citing works
Potential of Medicinal Plants as Neuroprotective and Therapeutic Properties Against Amyloid-β-Related Toxicity, and Glutamate-Induced Excitotoxicity in Human Neural Cells
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RETRACTED: Prevention of cardiovascular disease for healthy aging and longevity: A new scoring system and related “mechanisms-hallmarks-biomarkers”
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Retracted
RETRACTED ARTICLE: Biotechnology for propagation and secondary metabolite production in Bacopa monnieri
Sanyal, R., Nandi, S., Pandey, S., et al. (2022) · Applied Microbiology and Biotechnology · doi:10.1007/s00253-022-11820-6 · Cited by 25 · see citing works
“Additionally, the plant possesses neuroprotective, anti-neuro-inflammatory, pro-cognitive, neuropsychiatric, antinociceptive, analgesic, anticancer, antioxidant, antipyretic, and anticonvulsant properties (Nemetchek et al. 2017; Ranjan et al. 2018; Abdul Manap et al. 2019; Brimson et al. 2020; Castelli et al. 2020; Je…”
Retracted
Piquiá Shells (Caryocar villosum): A Fruit by-Product with Antioxidant and Antiaging Properties in Caenorhabditis elegans
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“However, an isolated triterpenoid saponin from Panax ginseng [95], a steroidal saponin from Asparagus racemosus [96], and plant extracts particularly rich in triterpenoid saponins [97, 98] were previously reported to enhance stress resistance and to prolong lifespan in C. elegans .”
Herbal medicines and botanicals for managing insomnia, stress, anxiety, and depression: A critical review of the emerging evidence focusing on the Middle East and Africa
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“It protects against glutamate toxicity induced by 5 mM glutamate in Caenorhabditis elegans ( C. elegans ) and modulates glutamate receptor binding and N-methyl-D-aspartate (NMDA) R1 gene expression in hippocampus of temporal lobe epileptic rats [76,77].”
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Hibiscus sabdariffa extract protects HT-22 cells from glutamate-induced neurodegeneration by upregulating glutamate transporters and exerts lifespan extension in C. elegans via DAF-16 mediated pathway
Malar, D. S., Prasanth, M. I., Brimson, J. M., Verma, K., Prasansuklab, A., & Tencomnao, T. (2021) · Nutrition and Healthy Aging · doi:10.3233/nha-210131 · Cited by 18 · see citing works
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Antinociceptive activity of standardized extract of Bacopa monnieri in different pain models of zebrafish
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Cleistocalyx nervosum var. paniala seed extracts exhibit sigma-1 antagonist sensitive neuroprotective effects in PC12 cells and protects C. elegans from stress via the SKN-1/NRF-2 pathway
Brimson, J. M., Prasanth, M. I., Isidoro, C., Sukprasansap, M., & Tencomnao, T. (2021) · Nutrition and Healthy Aging · doi:10.3233/nha-200108 · Cited by 13 · see citing works
“It has been observed that many plants or plant products including green tea [41], Streblus asper [42], Bacopa monnieri [43], Paullinia cupanna [44], and mulberry [45] can extend the lifespan and health-span and protect against stress in C.”
BackgroundMethodSelf-citation
Bacopa Protects against Neurotoxicity Induced by MPP+ and Methamphetamine
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Bacopa monnieri for Disorders Affecting Brain: Current Perspectives
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Spectroscopic, Electrochemical, and Biological Assays of Copper-Binding Molecules for Screening of Different Drugs and Plant Extracts against Neurodegenerative Disorders
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“monnieri hexane extract (Brimson et al., 2020), sanguinarine (Liu et al.”
Background
The Role of Bacopa monnieri in Alzheimer’s Disease: Mechanisms and Potential Clinical Use—A Review
Wiciński, M., Fajkiel-Madajczyk, A., Wójcicki, J., Ozorowski, M., & Szambelan, M. (2025) · Nutrients · doi:10.3390/nu17223538 · Cited by 7 · see citing works
“Brimson et al. [48] In vitro HT-22 cell line and wild C. elegans Anti-glutamate toxicity action.”
Influential
RETRACTED ARTICLE: Bacopa monnieri attenuates glutamate-induced nociception and brain mitochondrial toxicity in Zebrafish
Sharma, M., Gupta, P. K., & Garabadu, D. (2021) · Metabolic Brain Disease · doi:10.1007/s11011-021-00874-6 · Cited by 7 · see citing works
Retracted
HydroZitLa inhibits calcium oxalate stone formation in nephrolithic rats and promotes longevity in nematode Caenorhabditis elegans
Lordumrongkiat, N., Chotechuang, N., Prasanth, M. I., Jindatip, D., Ma-on, C., Chuenwisad, K., Leelahavanichkul, A., Tencomnao, T., & Boonla, C. (2022) · Scientific Reports · doi:10.1038/s41598-022-08316-8 · Cited by 7 · see citing works
Self-citation
Rhinacanthin-C but Not -D Extracted from Rhinacanthus nasutus (L.) Kurz Offers Neuroprotection via ERK, CHOP, and LC3B Pathways
Rakkhittawattana, V., Panichayupakaranant, P., Prasanth, M. I., et al. (2022) · Pharmaceuticals · doi:10.3390/ph15050627 · Cited by 7 · see citing works
“Glutamate has been shown in previous studies to induce the production of reactive oxygen species (ROS) in HT-22 cells [16,20,21].”
BackgroundSelf-citation
Agarwood leaf ethanol extract provides neuroprotective properties and promotes cholinergic differentiation of HT22 hippocampal neurons
Kumaree, K. K., Brimson, J. M., Verma, K., Chuchawankul, S., Tencomnao, T., & Prasansuklab, A. (2025) · Scientific Reports · doi:10.1038/s41598-025-93462-y · Cited by 6 · see citing works
Self-citation
Alzheimer’s Disease: The Current and Emerging Treatment Approaches
Pang, R., Jia, Q., Ma, C., Li, T., Bi, W., Wang, H. P., Liu, R., Chen, P., Lee, E., & Jiang, H. B. (2025) · Behavioural Neurology · doi:10.1155/bn/9627699 · Cited by 6 · see citing works
Simple ammonium salt and sigma-1 receptor ligand dipentylammonium provides neuroprotective effects in cell culture and Caenorhabditis elegans models of Alzheimer's disease
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Self-citation
Streblus asper Lour. exerts MAPK and SKN-1 mediated anti-aging, anti-photoaging activities and imparts neuroprotection by ameliorating Aβ in Caenorhabditis elegans
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“Caenorhabditis elegans , the in vivo model system, has been widely used to study aging, development, reproduction, obesity, stress resistance, neuroprotection and many more diverse mechanisms [21–24].”
BackgroundSelf-citation
Identification and pathogenicity of Alternaria alternata causing leaf blight of Bacopa monnieri (L.) Wettst. and its biocontrol by Trichoderma species in agrifields--an ecofriendly approach
Ghosh, S. K., Pal, S., & Banerjee, S. (2022) · Journal of Applied Research on Medicinal and Aromatic Plants · doi:10.1016/j.jarmap.2022.100406 · Cited by 5 · see citing works
DAF-16 and SKN-1 mediate Anti-aging and Neuroprotective efficacies of “thai ginseng” Kaempferia parviflora Rhizome extract in Caenorhabditis elegans
Prasanth, M. I., Malar, D. S., Brimson, J. M., et al. (2022) · Nutrition and Healthy Aging · doi:10.3233/nha-210148 · Cited by 5 · see citing works
“aging, stress resistance and neuroprotective effects of 113 various traditional medicinal plants including, Cleis114 tocalyx nervosum [13, 14], Lignosus rhinoceros [15], 115 Bacopa monnieri [16], Caesalpinia mimosoides [17] 116 and Diplocyclos palmatus [18].”
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Physical, chemical, and biological evaluation of nanoparticles containing phenolic compounds from wine production residues
Dávila-Trujillo, R., Fernandes, S. S., Dora, C. L., et al. (2021) · Journal of Food Processing and Preservation · doi:10.1111/jfpp.15629 · Cited by 4 · see citing works
Dammarane triterpene saponins in Bacopa monnieri (Brahmi): Unraveling their pharmacological potential
Tanwar, S. S., & Sharma, S. (2026) · Pharmacological Research - Natural Products · doi:10.1016/j.prenap.2026.100726 · Cited by 3 · see citing works
Effect of Thai medicinal plants Acanthus ebracteatus Vahl. Carthamus tinctorius L. and Streblus asper Lour. on neurite outgrowth activity in Neuro-2A cells
Rakkhitawatthana, V., Sillapachaiyaporn, C., Nilkhet, S., Brimson, J. M., & Tencomnao, T. (2022) · Journal of Associated Medical Sciences · doi:10.12982/jams.2023.010 · Cited by 3 · see citing works
Self-citation
Hylocereus undatus extends lifespan and exerts neuroprotection in Caenorhabditis elegans via DAF-16 mediated pathway
Prasanth, M. I., Prasansuklab, A., Verma, K., et al. (2023) · Nutrition and Healthy Aging · doi:10.3233/nha-220196 · Cited by 3 · see citing works
“Previous reports of plant extracts including Cleistocalyx nervosum [22, 23], Bacopa monnieri [24], Streblus asper [25], Hibiscus sabdar-iffa [26], and Kaempferia parviﬂora [27, 28] in C. elegans showed extension of lifespan and reduction of lipofuscin accumulation indicating its anti-aging…”
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Shramahara Mahakasya, a traditional polyherbal formulation, induces anti-anxiety activity in hippocampal neurons by effectuating SOD2-mediated protection against oxidative stress
Saini, S., Kumar, V., Haldar, S., Chauhan, S., Demiwal, P., Ghosh, S., Gupta, S., Sircar, D., Mahajon, B., & Roy, P. (2024) · Phytomedicine Plus · doi:10.1016/j.phyplu.2024.100682 · Cited by 3 · see citing works
Bacopaside-I Alleviates the Detrimental Effects of Acute Paraquat Intoxication in the Adult Zebrafish Brain
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Ligand docking in the sigma-1 receptor compared to the sigma-1 receptor-BiP complex and the effects of agonists and antagonists on C. elegans lifespans
Verma, K., Prasanth, M. I., Tencomnao, T., & Brimson, J. M. (2024) · Biomedicine & Pharmacotherapy · doi:10.1016/j.biopha.2024.117783 · Cited by 2 · see citing works
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Review of ethnomedicinal uses and phytopharmacology of herbs for the treatment of Alzheimer's disease
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Nutri Ayur in Cognition and Memory Dysfunction
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“The intestinal autofluorescence analysis sug - gested decreased aging and oxidative stress in the group with 10 µg/mL of extracts (Brimson et al. 2020).”
The Role of Herbal Medicines in Cognitive Impairment: A Narrative Review of Neuroprotective Mechanisms and Therapeutic Perspectives
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Authors: Brimson, J. M., Prasanth, M. I., Plaingam, W., & Tencomnao, T.
Affiliations: Chulalongkorn University (TH)
Rangsit University (TH)
Corresponding author: Tencomnao, Tewin
ORCID: Brimson, James Michael
Prasanth, Mani Iyer
Plaingam, Waluga
Tencomnao, Tewin
Venue: Journal of Traditional and Complementary Medicine
Publisher: Elsevier BV
Published: 2020-09
Volume: 10
Issue: 5
Pages: 460-470
ISSN: 2225-4110 (ISSN-L)
DOI: 10.1016/j.jtcme.2019.10.001
PMC: PMC7484969
OpenAlex: W2978806321
Semantic Scholar: a20e924f522440b3f7eb2d2b4c450290dd7b4525
Cited by: 51
References: 43
Citations by year: 2026: 4 · 2025: 7 · 2024: 9 · 2023: 7 · 2022: 11 · 2021: 10 · 2020: 3
FWCI: 7.73
Type: journal article
Language: en
Open access: diamond
doi.org
Licence: cc-by-nc-nd
www.elsevier.com/tdm/userlicense/1.0/ (tdm)
www.elsevier.com/legal/tdmrep-license (tdm)
creativecommons.org/licenses/by-nc-nd/4.0/ (vor)
Locations: Journal of Traditional and Complementary Medicine (publishedVersion, cc-by-nc-nd)
PubMed Central (publishedVersion, cc-by-nc-nd)
PubMed (publishedVersion)
DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
Funders: Chulalongkorn University
Topics: Health Sciences › Medicine › Complementary and alternative medicine › Medicinal Plants and Neuroprotection
Health Sciences › Medicine › Physiology › Alzheimer's disease research and treatments
Health Sciences › Medicine › Complementary and alternative medicine › Phytochemicals and Medicinal Plants
Keywords: Bacopa monnieri, glutamate excitotoxicity, neuroprotection, Caenorhabditis elegans, lifespan extension, aging, mitochondrial stress, endoplasmic reticulum stress, oxidative stress, reactive oxygen species, lipofuscin, neurodegenerative diseases, Alzheimer's disease, HT22 cells, plant extracts

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Bacopa monnieri (L.) wettst. Extract protects against glutamate toxicity and increases the longevity of Caenorhabditis elegans

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