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Aucubin and catalpol concentrations in leaves and roots of Plantago lanceolata.
Plants were grown at different light and nutrient availability (N-: low nutrient, N+: high nutrient, L-: low light, L+: high light). Values are means across all plants per treatment (+ 1 SE). Results of Tukey`s test applied to test for significant differences between different light x nutrient treatments are indicated with letters.

Aucubin and catalpol concentrations in leaves and roots of Plantago lanceolata. Plants were grown at different light and nutrient availability (N-: low nutrient, N+: high nutrient, L-: low light, L+: high light). Values are means across all plants per treatment (+ 1 SE). Results of Tukey`s test applied to test for significant differences between different light x nutrient treatments are indicated with letters.

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Article
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A few studies in the past have shown that plant diversity in terms of species richness and functional composition can modify plant defense chemistry. However, it is not yet clear to what extent genetic differentiation of plant chemotypes or phenotypic plasticity in response to diversity-induced variation in growth conditions or a combination of bot...

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Context 1
... iridoid glycosides, aucubin and catalpol concentrations in leaves did not depend on species number and presence of legumes in the plots of origin (Table 1). High light conditions had positive effects on the concentrations of aucubin and catalpol in both above- and belowground plant tissue (Fig 1, Table 1). While leaf aucubin concentrations were nearly four times higher under high light than under low light conditions, leaf catalpol concentrations had twice the concentrations under high light compared to low light conditions. ...
Context 2
... of nutrient addition on foliar aucubin concentration depended on light availability (significant interaction nutrient x light). Aucubin concentrations were on average 12% higher in the high nutrient treatment than in the low nutrient treatment when plants were grown under high light conditions, while aucubin concentrations were on average 8% lower in the high nutrient treatment than in the low nutrient treatment when plants were grown under low light conditions although the group means between low and high light conditions did not differ significantly (Fig 1). Aucubin concentrations in the roots were not affected by nutrient availability. ...
Context 3
... we found consistent positive effects of light on P. lanceolata defense compounds, the effects of nutrient availability on verbascoside and iridoid glycoside concentrations differed: Verbascoside concentrations decreased under high nutrient conditions in our experiment ( Fig 2) and also in other studies [1], but iridoid glycoside concentrations were only marginally affected by nutrient availability (Fig 1). Phenolics such as verbascoside are derived from the shikimic acid pathway, whereas terpenoids, including the monoterpene-derived iridoid glyco- sides, are synthesized via the methylerythritol-phosphate (MEP) pathway [62,63]. ...

Citations

... The wide differences in the results obtained for aucubin content in the leaves between Tamura and Nishibe 2002 19 and the experiment conducted at Frick (Switzerland) are most likely due to environmental conditions. Further research conducted by Miehe-Steier et al. in a greenhouse concluded that phenotypic plasticity responding to environmental changes could be responsible for the variation in the aucubin content of the aboveground parts of ribwort plantain 22 . The authors obtained aucubin content results ranging from 1.6% DM to just over 1.8% DM under high light conditions (14 hours of light per day). ...
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This study investigated the potential of Plantago lanceolata as an alternative plant to mitigate the environmental impacts of agriculture in the face of climate change. Eleven genotypes of ribwort were compared under field conditions in northern Switzerland, focusing on the characteristics of the root system and the aucubin content in the leaves and roots. Aucubin, known to inhibit nitrification process in the soil, varied according to genotype and season. The content of secondary metabolite in leaves was higher than in roots, which may indicate the greater importance of leaves in reducing soil nitrification. Dry matter yield and for the first time root characteristics, such as length density and specific root length, were evaluated at two different depths, and at different positions in the experimental plots. The results showed that ribwort is well adapted in terms of rooting in topsoil. In addition, the plant's ability to root deeply after one year of cultivation under field conditions has been demonstrated, reaching depths of up to 1.88 metres. This allows the plant to effectively absorb water and nutrients from the subsoil layers.
... Tannins are highly variable in their chemical structures, with several hundred unique molecules detectable across various plant species (Salminen & Karonen, 2011). Genetic-based variation among host plant species (e.g., Bernhardsson et al., 2013;Bowers & Stamp, 1992) and variation in environmental factors such as nutrients, water, temperature, CO 2 , light (Miehe-Steier et al., 2015) and herbivory (Kessler & Kalske, 2018) determine the concentrations of plant defense compounds. As invasive alien plants often experience less herbivory than native plants in the exotic range (Geppert et al., 2021), it is plausible that invasive species should express lower concentrations of flavonoids and tannins than co-occurring native plant species. ...
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In support of the prediction of the enemy release hypothesis regarding a growth–defense trade‐off, invasive alien plants often exhibit greater growth and lower anti‐herbivory defenses than native plants. However, it remains unclear how nutrient enrichment of invaded habitats may influence competitive interactions between invasive alien and co‐occurring native plants, as well as production of anti‐herbivore defense compounds, growth‐promoting hormones, and defense‐regulating hormones by the two groups of plants. Here, we tested whether: (i) nutrient enrichment causes invasive alien plants to produce greater biomass and lower concentrations of the defense compounds flavonoids and tannins than native plants; and (ii) invasive alien plants produce lower concentrations of a defense‐regulating hormone jasmonic acid (JA) and higher concentrations of a growth‐promoting hormone gibberellic acid (GA3). In a greenhouse experiment, we grew five congeneric pairs of invasive alien and native plant species under two levels each of nutrient enrichment (low vs. high), simulated herbivory (leaf clipping vs. no‐clipping), and competition (alone vs. competition) in 2.5‐L pots. In the absence of competition, high‐nutrient treatment induced a greater increase in total biomass of invasive alien species than that of native species, whereas the reverse was true under competition as native species benefitted more from nutrient enrichment than invasive alien species. Moreover, high‐nutrient treatment caused a greater increase in total biomass of invasive alien species than that of native species in the presence of simulated herbivory. Competition induced higher production of flavonoids and tannins. Simulated herbivory induced higher flavonoid expression in invasive alien plants under low‐nutrient than high‐nutrient treatments. However, flavonoid concentrations of native plants did not change under nutrient enrichment and simulated herbivory treatments. Invasive alien plants produced higher concentrations of GA3 than native plants. Taken together, these results suggest that impact of nutrient enrichment on growth of invasive alien and co‐occurring native plants may depend on the level of competition that they experience. Moreover, invasive alien plants might adjust their flavonoid‐based defense more efficiently than native plants in response to variation in soil nutrient availability and herbivory pressure. Our findings suggest that large‐scale efforts to reduce nutrient enrichment of invaded habitats may help to control future invasiveness of target alien plant species.
... Importantly, these plants differ in their secondary chemistry, which may influence herbivore interactions with enemies. Plantago contains iridoid glycosides (IGs), monoterpenoid metabolites that can be toxic and/or deterrent to non-adapted herbivores (Abate et al., 2022;Bowers & Puttick, 1988;Miehe-Steier et al., 2015). Anartia jatrophae larvae are capable of sequestering and retaining small amounts of IGs into the pupal and adult stages (Knerl & Bowers, 2013), which provide chemical defence in other sequestering species (Bowers, 1993). ...
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Incorporation of exotic plants into the diets of native herbivores is a common phenomenon, influencing interactions with natural enemies and providing insight into the tritrophic costs and benefits of dietary expansion. We evaluated how use of an exotic plant, Plantago lanceolata, impacted immune performance, development and susceptibility to pathogen infection in the neotropical herbivore Anartia jatrophae (Lepidoptera: Nymphalidae). Caterpillars were reared on P. lanceolata or a native plant, Bacopa monnieri, and experimentally infected with a pathogenic virus, Junonia coenia densovirus. We found that virus‐challenged herbivores exhibited higher survival rates and lower viral burdens when reared on P. lanceolata compared to B. monnieri, though immune performance and development time were largely similar on the two plants. These findings reveal that use of an exotic plant can impact the vulnerability of a native herbivore to pathogen infection, suggesting diet‐mediated protection against disease as a potential mechanism facilitating the incorporation of novel resources. In this study, we evaluated how use of an exotic host plant, Plantago lanceolata, impacted immune performance, development and susceptibility to pathogen infection in the neotropical herbivore Anartia jatrophae (Lepidoptera: Nymphalidae). We found that virus‐challenged herbivores exhibited higher survival rates and lower viral burdens when reared on P. lanceolata compared to a native host plant, though immune performance and development time were largely similar on the two plants. These findings reveal that use of an exotic plant can impact the vulnerability of a native herbivore to pathogen infection, suggesting diet‐mediated protection against disease as a potential mechanism facilitating the incorporation of novel resources.
... At this point, the increase and consequent decrease in the relative percentage of eupatoriochromene stands out. Some chromenes can act in the defense of vegetative organs against natural predators or phytopathogens [8,[109][110][111][112]. In addition, eupatoriochromene may be related to the process of maturation and development of inflorescences. ...
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The aim of this study was to monitor the volatile chemical composition from leaves and reproductive organs of Piper mollicomum Kunth (PM), in its reproduction period, as well as register inflorescence visitors, microclimate and phenological information. The essential oils (EOs) obtained from the different fresh organs by hydrodistillation were identified and quantified by Gas Chromatography/Mass Spectrometry (GC/MS) and by GC coupled to a Flame Ionization Detector (GC/FID), respectively. The cercentage content of some volatiles present in reproductive organs, such as limonene, 1,8-cineole, linalool and eupatoriochromene, increased during the maturation period of the inflorescences, and decreased during the fruiting period, suggesting a defense/attraction activities. Furtermore, a biosynthetic dichotomy between 1,8-cineole (leaves) and linalool (reproductive organs) was recorded. A high frequency of bee visits was registered weekly, and some correlations showed a positive relationship between this variable and terpenes. Microclimate has an impact on this species’ phenological cycles and insect visiting behavior. All correlations between volatiles, insects, phenology and microclimate allowed us to present important data about the complex information network in PM. These results are extremely relevant for the understanding of the mechanisms of chemical–ecological plant–insect interactions in Piperaceae, a basal angiosperm.
... The accumulation of compounds 5-9 can be related with the effect of sunlight and nutrients availability, etc. In harmony, the derivates of plant iridoids and phenolic metabolites have been studied, and found that the production of such compounds have significant correlation with the light and nutrients availability 38 . The accumulation of these metabolites can also suggest the result of the soil that contains, for example an increased level of aluminum 39 . ...
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This research characterizes key metabolites in the leaf from Citronella gongonha Martius (Mart.) Howard (Cardiopteridaceae). All metabolites were assessed in intact leaf tissue by proton (1H) high-resolution magic angle spinning (HR-MAS) nuclear magnetic resonance (NMR) spectroscopy integrated with the principal component analysis (PCA) to depict molecular association with the seasonal change. The major ‘known unknown’ metabolites detected in 1H HR-MAS NMR were derivatives of flavonoid, polyphenolic and monoterpenoid compounds such as kaempferol-3-O-dihexoside, caffeoyl glucoside (2), 3-O-caffeoylquinic acid (3), 5-O-caffeoylquinic acid (4), kingiside (5), 8-epi-kingisidic acid (6), (7α)-7-O-methylmorroniside (7), (7β)-7-O-methylmorroniside (8) and alpigenoside (9) together with the universally occurring sucrose (10), α-glucoses (11, 12), alanine (13), and fatty (linolenic) acid (14). Several of the major metabolites (1, 2–9) were additionally confirmed by liquid chromatography tandem mass spectrometry (LC–MS/MS). In regard with the PCA results, metabolites 1, 2–9 and 14 were influenced by seasonal variation and/or from further (a) biotic environmental conditions. The findings in this work indicate that C. gongonha Mart. is an effective medicinal plant by preserving particularly compounds 2, 3–9 in abundant amounts. Because of close susceptibility with seasonal shift and ecological trends, further longitudinal studies are needed to realize the physiology and mechanism involved in the production of these and new metabolites in this plant under controlled conditions. Also, future studies are recommended to classify different epimers, especially of the phenolics and monoterpenoids in the given plant.
... Another iridoid glycoside aucubin (tR = 11.04, with m/z 346.1258, Table 1), along with catalpol, are pharmacologically active natural components of numerous plants (Sertic et al., 2015;Miehe-Steier et al., 2015). Ganolucidic acid B (tR = 17.73, with m/z 502.3296, ...
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Honey represents a valuable food commodity, known since ancient times for its delicate taste and health benefits due to its specific compositional characteristics, mainly the phenolic compound content. “Anama” honey is a monofloral honey produced from the nectar of Erica manipuliflora plant, a heather bush of the Greek island of Ikaria, one of the Mediterranean’s longevity regions. “Anama” is characterized by a unique aroma and taste, with a growing demand for consumption and the potential to be included in the list of products with a protected designation of origin. The aim of this study was to determine the chemical and botanical profile of authentic Anama honey samples and find similarities and differences with honey samples of a different botanical origin from the same geographical area. Untargeted Ultrahigh-Performance Liquid Chromatography-Hybrid Quadrupole-Orbitrap High-Resolution Mass Spectrometry (UHPLC-HRMS) metabolomics study was conducted on authentic heather, pine, and thyme honey samples from Ikaria and neighboring islands. The Principal Component Analysis (PCA), Orthogonal Projections to Latent Structures Discriminant Analysis (OPLS-DA), and differential analysis were performed using the entire metabolic profile of the samples and allowed the identification of chemical markers for sample discrimination. Thirty-two characteristic secondary metabolites (cinnamic acids, phenolic acids, flavonoids, terpenes) and other bioactive phenolic compounds, some of them not previously reported in a heather honey (aucubin, catalpol, domesticoside, leonuriside A, picein among others), emerged as potential chemical indicators of Anama honey. Melissopalynological analysis was also carried out to decipher the botanical and geographical origin of Anama honey. The relative frequency of the pollen of dominant plants of the Ericaceae family and a multitude of nectariferous and nectarless plants contributing to the botanical profile of Anama was evaluated. The identification of the pollen sources enabled a potential correlation of differentially increased secondary metabolites and chemicals with their botanical origin. The physicochemical profile of Anama was also determined, including the parameters of pH, color, electrical conductivity, diastase, moisture, as well as sugars, supporting the high quality of this heather honey.
... Plantain leaf and stem material could be incorporated into the soil through animal trampling during grazing. Although the aucubin concentration in roots of plantain pastures was not determined, another pathway for aucubin to be released into the soil could be via root senescence as the aucubin concentration in the roots is similar to that in the leaves (De Deyn et al. 2009;Wurst et al. 2010;Miehe-Steier et al. 2015). Recently, de Klein et al. (2019) hypothesised that plantain could incorporate secondary metabolites into the soil through the release of aucubin in root exudates. ...
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Plantain (Plantago lanceolata L.) is a forage that produces secondary metabolites with one, aucubin, known to inhibit soil nitrification. This study aimed to quantify the exudation of aucubin and catalpol by plantain root systems in a hydroponic experiment; evaluate the effect of aucubin on nitrogen (N) leaching and nitrous oxide (N2O) emissions from urine applied to plantain and ryegrass-white clover swards; and compare N losses from a ryegrass-white clover sward after urine from cows fed plantain and cows fed ryegrass-white clover was applied. Nitrate (NO3−) leaching and N2O losses were measured in a lysimeter experiment. Catalpol, but not aucubin, was exudated by plantain roots. N2O emissions were decreased by plantain swards and by ryegrass-white clover swards to which aucubin was also applied. Aucubin had no effect on NO3− leaching. Urine from cows grazing plantain had no effect on N2O emissions, and N leaching when compared to the urine from cows grazing ryegrass-white clover with the same N content. We conclude the plantain sward and the aucubin applied to the ryegrass-white clover sward decreased N2O emissions via mechanisms in the soil separate from the decreased emissions resulting from the lower N concentration of urine derived from cows grazing plantain.
... Under exposure to Ni-and Al-oxides nanoparticles, iridoid glycosides, phenolics, and flavonoids contents were significantly increased with the accumulation of reactive oxygen species in Nigella arvensis, suggesting the possible roles of iridoid glycosides during metal stress [49]. In our study, the aucubin concentrations (summer, 132 mg/g DW; winter, 114 mg/ g DW; calculated using root water content) in A. japonica roots were considerably higher than those of general antioxidants in plants; for example, the concentrations of antioxidative total phenolics in tree leaves were approximately 20 to 40 mg/g DW [50], and aucubin concentration in Plantago species and Eucommia ulmoides (abundant in aucubin) were approximately 7 to 20 mg/kg DW [51,52]. This suggests that A. japonica constantly produces a high concentration of aucubin in roots, which can scavenge reactive oxygen species in plants. ...
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Aucuba japonica Thunb. is an evergreen understory shrub that grows naturally at a mine site. The mine soil contains high concentrations of heavy metals, and A . japonica appears to maintain detoxification mechanisms against heavy metals in the study site’s understory. This study aimed to investigate the heavy metal tolerance mechanisms in A . japonica , considering the possible roles of arbuscular mycorrhizal and root-endophytic fungi. We conducted fieldwork in summer (canopy-foliation season) and winter (canopy-defoliation season) to measure the heavy metal concentrations in leaves, branches, and roots and analyze possible detoxicants in the roots. The infection rates of arbuscular mycorrhizal and root-endophytic fungi were evaluated via microscopic observation, and heavy metal (Zn) localization in A . japonica roots was observed using confocal laser scanning microscopy. Field analysis showed that A . japonica accumulated excessive Zn and produced aucubin and citric acid in the roots in both summer and winter. Zn localization observations clarified that Zn was distributed in thickened epidermal and cortical cell walls, suggesting that the cell walls functioned as Zn deposition sites, reducing Zn toxicity. It was further clarified that Zn was contained within cortical cells, indicating that Zn might be detoxified by aucubin and citric acid. Arbuscular mycorrhizal and root-endophytic fungi within cortical cells adsorbed Zn on fungal cell walls, indicating that these fungi would reduce Zn content within root cells and might alleviate Zn toxicity. Our results indicated that A . japonica would maintain Zn tolerance in both summer and winter via Zn immobilization in the cell walls and production of aucubin and citric acid, and that arbuscular mycorrhizal and root-endophytic fungi might play important roles in the Zn tolerance of A . japonica .
... In addition, they can be an oviposition stimulant for specialized insects [35]. The amount is stimulated by environmental factors in accordance with Miehe-Steier et al. [40]; ...
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This article gives an overview of the widely distributed herb ribwort plantain (Plantago lanceolata). Currently, this plant is mostly grown in grasslands, rarely in arable land, and its secondary metabolites are used in medicine. Studies cited in the article indicate its very rapid growth and deep rooting in the soil, which results in high drought tolerance and uptake of valuable nutrients from deep soil layers. An intensive symbiosis with various mycorrhizal fungi is characteristic of plantain for a high capacity for nutrient and water appropriation. It is also characterized by different contents of iridoid glycosides like aucubin and catalpol in shoots and roots of different varieties. The use of P. lanceolata is discussed on permanent and non-permanent grasslands where agriculturally specific varieties have been developed for grazing animals showing positive health effects in them. Information is provided on the cultivation of ribwort plantain, including studies on sowing and fertilization, the yield and infestation of the plant with pathogens, and the occurrence of iridoid glycosides in the plant. In addition, information is included on pests that threaten the plant’s cultivation.
... Probably, fungal infestation depends to a large extent on weather conditions, the actual growth environment, and neighboring plants, and to a lower extent on differences in the plant history.In line with the leaf damage results, we found no effects of plant history on the expression of leaf traits (except higher leaf greenness of P. trivialis in home than in away-same soil in May 2019). This suggests that phenotypic differences in leaf trait expression in response to community diversity, which was shown in several studies(Bachmann et al., 2018;Gubsch et al., 2011), were not induced by eco-evolutionary feedbacks, but only by phenotypic plasticity in response to diversity and composition of the actual community, which is in line with a study byMiehe-Steier et al. (2015). Nevertheless, our findings contrast previous studies Moorsel et al., 2018; Zuppinger-Dingley et al., 2014) showing different phenotypic trait expression of plants selected at low and high diversity. ...
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Long‐term biodiversity experiments have shown increasing strengths of biodiversity effects on plant productivity over time. However, little is known about rapid evolutionary processes in response to plant community diversity, which could contribute to explaining the strengthening positive relationship. To address this issue, we performed a transplant experiment with offspring of seeds collected from four grass species in a 14‐year‐old biodiversity experiment (Jena Experiment). We used two‐ and six‐species communities and removed the vegetation of the study plots to exclude plant–plant interactions. In a reciprocal design, we transplanted five “home” phytometers (same origin and actual environment), five “away‐same” phytometers (same species richness of origin and actual environment, but different plant composition), and five “away‐different” phytometers (different species richness of origin and actual environment) of the same species in the study plots. In the establishment year, plants transplanted in home soil produced more shoots than plants in away soil indicating that plant populations at low and high diversity developed differently over time depending on their associated soil community and/or conditions. In the second year, offspring of individuals selected at high diversity generally had a higher performance (biomass production and fitness) than offspring of individuals selected at low diversity, regardless of the transplant environment. This suggests that plants at low and high diversity showed rapid evolutionary responses measurable in their phenotype. Our findings provide first empirical evidence that loss of productivity at low diversity is not only caused by changes in abiotic and biotic conditions but also that plants respond to this by a change in their micro‐evolution. Thus, we conclude that eco‐evolutionary feedbacks of plants at low and high diversity are critical to fully understand why the positive influence of diversity on plant productivity is strengthening through time.