Abiotic stressors such as extreme temperatures, drought, flood, light, salt, and heavy metals alter biological diversity and crop production worldwide. Therefore, it is important to know the mechanisms by which plants cope with stress conditions. Polyphenols, which are the largest group of plant-specialized metabolites, are generally recognized as molecules involved in stress protection in plants. This diverse group of metabolites contains various structures, from simple forms consisting of one aromatic ring to more complex ones consisting of large number of polymerized molecules. Consequently, all these molecules, depending on their structure, may show different roles in plant growth, development, and stress protection. In the present review, we aimed to summarize data on how different polyphenol structures influence their biological activity and their roles in abiotic stress responses. We focused our review on phenolic acids, flavonoids, stilbenoids, and lignans.
Salinity is a major abiotic stress negatively affecting plant growth and consequently crop production. The effects of short-term salt stress were evaluated on seedlings of three globally important Brassica crops—Chinese cabbage (Brassica rapa ssp. pekinensis), white cabbage (Brassica oleracea var. capitata), and kale (Brassica oleracea var. acephala)—with particular focus on phenolic acids. The physiological and biochemical stress parameters in the seedlings and the levels of three main groups of metabolites (total glucosinolates, carotenoids, and phenolics) and individual phenolic acids were determined. The salt treatments caused a dose-dependent reduction in root growth and biomass and an increase in stress parameters (Na+/K+ ratio, reactive oxygen species (ROS) and glutathione (GSH)) in all seedlings but most prominently in Chinese cabbage. Based on PCA, specific metabolites grouped close to the more tolerant species, white cabbage and kale. The highest levels of phenolic acids, particularly hydroxycinnamic acids, were determined in the more tolerant kale and white cabbage. A reduction in caffeic, salicylic, and 4-coumaric acid was found in Chinese cabbage and kale, and an increase in ferulic acid levels was found in kale upon salinity treatments. Phenolic acids are species-specific among Brassicaceae, and some may participate in stress tolerance. Salt-tolerant varieties have higher levels of some phenolic acids and suffer less from metabolic stress disorders under salinity stress.
Drought is one of the major abiotic stresses affecting the productivity of Brassica crops. To understand the role of phytohormones in drought tolerance, we subjected Chinese cabbage (B. rapa ssp. pekinensis), white cabbage (B. oleracea var. capitata), and kale (B. oleracea var. acephala) to drought and examined the stress response on the physiological, biochemical and hormonal levels. The phytohormones abscisic acid (ABA), auxin indole-3-acetic acid (IAA), brassinosteroids (BRs), cytokinins (CKs), jasmonates (JAs), and salicylic acid (SA) were analyzed by ultra-high-performance liquid chromatography–tandem mass spectrometry (UHPLC-MS/MS). Based on the physiological and biochemical markers the Chinese cabbage exhibited the lowest tolerance, followed by the white cabbage, while the kale appeared to be the most tolerant to drought. The drought tolerance of the kale correlated with increased levels of SA, ABA, IAA, CKs iP(R) and cZ(R), and typhasterol (TY), a precursor of active BRs. In contrast, the drought sensitivity of the Chinese cabbage correlated with a significant increase in ABA, JAs and the active BRs castasterol (CS) and brassinolide (BL). The moderately tolerant white cabbage, positioned between the kale and Chinese cabbage, showed more similarity in terms of the phytohormone patterns with the kale. We concluded that the drought tolerance in Brassicaceae is mostly determined by the increased endogenous levels of IAA, CKs, ABA and SA and the decreased levels of active BRs.
The potential role of NaCl (50–200 mM) as an eustressor for the accumulation of health promoting phytochemicals and maintaining the homeostasis of macro- and micro-elements in three, hydroponically grown Brassica leafy vegetables (Chinese cabbage, white cabbage, and kale) was investigated. Considering K+/Na+ ratio and proline contents as reliable stress markers, we confirmed more prominent stress status in Chinese cabbage followed by white cabbage and kale. Low to moderate salinity treatments (50 and 100 mM NaCl) caused an increase in most of the phenolic compounds in the analyzed Brassica leafy vegetables. Total glucosinolates were elicited by NaCl in a dose dependent manner. Salt treatment caused an increase in total chlorophylls but did not significantly affect carotenoid content. Furthermore, low to moderate treatments did not significantly disturb homeostasis of macro- and micro-elements, particularly in white cabbage and kale where the K level did not decrease significantly and Ca was even increased in white cabbage. We may conclude that salinity may elicit phytochemical accumulation in selecting vegetables grown on saline soils without undesirable disturbance in macro- and micro-elements homeostasis depending on salt concentration and species/varieties. This information may be of great importance in the selection of crops grown on saline soils.
Salinity stress is one of the most damaging abiotic stresses to plants, causing disturbances in physiological, biochemical, and metabolic processes. The exogenous application of natural metabolites is a useful strategy to reduce the adverse effects of stress on crops. We investigated the effect of foliar application of salicylic acid (SA) and ferulic acid (FA) (10–100 μM) on short-term salt-stressed (150 mM NaCl, 72 h) Chinese cabbage plants. Subsequently, proline level, photosynthetic performance, phenolic metabolites with special focus on selected phenolic acids (sinapic acid (SiA), FA, SA), flavonoids (quercetin (QUE), kaempferol (KAE)), and antioxidant activity were investigated in salt-stressed and phenolic acid-treated plants compared with the corresponding controls. Salt stress caused a significant increase in SA and proline contents, a decrease in phenolic compounds, antioxidant activity, and photosynthetic performance, especially due to the impairment of PSI function. SA and FA treatments, with a concentration of 10 μM, had attenuated effects on salt-stressed plants, causing a decrease in proline and SA level, and indicating that the plants suffered less metabolic disturbance. Polyphenolic compounds, especially FA, SiA, KAE, and QUE, were increased in FA and SA treatments in salt-stressed plants. Consequently, antioxidant activities were increased, and photosynthetic performances were improved. FA resulted in a better ameliorative effect on salt stress compared to SA.
Brassica oleracea var. acephala is known to have a strong tolerance to low temperatures, but the protective mechanisms enabling this tolerance are unknown. Simultaneously, this species is rich in health-promoting compounds such as polyphenols, carotenoids, and glucosinolates. We hypothesize that these metabolites play an important role in the ability to adapt to low temperature stress. To test this hypothesis, we exposed plants to chilling (8 °C) and additional freezing (−8 °C) temperatures under controlled laboratory conditions and determined the levels of proline, chlorophylls, carotenoids, polyphenols, and glucosinolates. Compared with that of the control (21 °C), the chilling and freezing temperatures increased the contents of proline, phenolic acids, and flavonoids. Detailed analysis of individual glucosinolates showed that chilling increased the total amount of aliphatic glucosinolates, while freezing increased the total amount of indolic glucosinolates, including the most abundant indolic glucosinolate glucobrassicin. Our data suggest that glucosinolates are involved in protection against low temperature stress. Individual glucosinolate species are likely to be involved in different protective mechanisms because they show different accumulation trends at chilling and freezing temperatures.
Salt and osmotic stress are the main abiotic stress factors affecting plant root growth and architecture. We investigated the effect of salt (100 mM NaCl) and osmotic (200 mM mannitol) stress on the auxin metabolome by UHPLC-MS/MS, auxin distribution by confocal microscopy, and transcript levels of selected genes by qRT-PCR in Arabidopsis thaliana ecotype Columbia-0 (Col-0) and DR5rev::GFP (DR5) line. During long-term stress (13 days), a stability of the auxin metabolome and a tendency to increase indole-3-acetic acid (IAA) were observed, especially during salt stress. Short-term stress (3 h) caused significant changes in the auxin metabolome, especially NaCl treatment resulted in a significant reduction of IAA. The data derived from auxin profiling were consistent with gene expressions showing the most striking changes in the transcripts of YUC, GH3, and UGT transcripts, suggesting disruption of auxin biosynthesis, but especially in the processes of amide and ester conjugation. These data were consistent with the auxin distribution observed in the DR5 line. Moreover, NaCl treatment caused a redistribution of auxin signals from the quiescent center and the inner layers of the root cap to the epidermal and cortical cells of the root elongation zone. The distribution of PIN proteins was also disrupted by salt stress; in particular, PIN2 was suppressed, even after 5 min of treatment. Based on our results, the DR5 line was more sensitive to the applied stresses than Col-0, although both lines showed similar trends in root morphology, as well as transcriptome and metabolome parameters under stress conditions.
Kale (Brassica oleracea var acephala) is known as a vegetable with good tolerance of environmental stress and numerous beneficial properties for human health, which are attributed to different phytochemicals. In the present study, investigation of how low temperatures affect proline, pigments and specialized metabolites content was performed using 8-weeks old kale plants subjected to chilling (at 8 °C, for 24 h) followed by short freezing (at −8 °C, for 1 h after previous acclimation at 8 °C, for 23 h). Plants growing at 21 °C served as a control. In both groups of plants (exposed to low temperatures and exposed to short freezing) a significant increase in proline content (14% and 49%, respectively) was recorded. Low temperatures (8 °C) induced an increase of pigments (total chlorophylls 7%) and phytochemicals (phenolic acids 3%; flavonoids 5%; carotenoids 15%; glucosinolates 21%) content, while exposure to freezing showed a different trend dependent upon observed parameter. After freezing, the content of chlorophylls, carotenoids, and total phenolic acids retained similar levels as in control plants and amounted to 14.65 ± 0.36 mg dw g−1, 2.58 ± 0.05 mg dw g−1 and 13.75 ± 0.07 mg dw CEA g−1, respectively. At the freezing temperature, total polyphenol content increased 13% and total flavonoids and glucosinolates content decreased 21% and 54%, respectively. Our results suggest that acclimatization (23 h at 8 °C) of kale plants can be beneficial for the accumulation of pigments and phytochemicals, while freezing temperatures affect differently specialized metabolite synthesis. The study suggests that growing temperature during kale cultivation must be considered as an important parameter for producers that are orientated towards production of crops with an increasing content of health-related compounds.
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