Secondary metabolites responses of plants exposed to ozone: an update.

Tropospheric ozone (O3) is a secondary pollutant that causes oxidative stress in plants due to the generation of excess reactive oxygen species (ROS). Phenylpropanoid metabolism is induced as a usual response to stress in plants, and induction of key enzyme activities and accumulation of secondary metabolites occur, upon O3 exposure to provide resistance or tolerance. The phenylpropanoid, isoprenoid, and alkaloid pathways are the major secondary metabolic pathways from which plant defense metabolites emerge. Chronic exposure to O3 significantly accelerates the direction of carbon flows toward secondary metabolic pathways, resulting in a resource shift in favor of the synthesis of secondary products. Furthermore, since different cellular compartments have different levels of ROS sensitivity and metabolite sets, intracellular compartmentation of secondary antioxidative metabolites may play a role in O3-induced ROS detoxification. Plants' responses to resource partitioning often result in a trade-off between growth and defense under O3 stress. These metabolic adjustments help the plants to cope with the stress as well as for achieving new homeostasis. In this review, we discuss secondary metabolic pathways in response to O3 in plant species including crops, trees, and medicinal plants; and how the presence of this stressor affects their role as ROS scavengers and structural defense. Furthermore, we discussed how O3 affects key physiological traits in plants, foliar chemistry, and volatile emission, which affects plant-plant competition (allelopathy), and plant-insect interactions, along with an emphasis on soil dynamics, which affect the composition of soil communities via changing root exudation, litter decomposition, and other related processes.

Ultraviolet-B and Heavy Metal-Induced Regulation of Secondary Metabolites in Medicinal Plants: A Review.

Despite a rich history and economic importance, the potential of medicinal plants has not been fully explored under different abiotic stress conditions. Penetration of UV-B radiation and contamination of heavy metals are two important environmental stress for plants with remarkable influence on the defense-related and pharmaceutically important secondary metabolites of medicinal plants. UV-B and heavy metal contamination may become a critical issue that either positively or negatively affects the quality and quantity of secondary metabolites. Such effects may result from changes in the expression level of genes that encode the corresponding enzymes or the inactivation and/or stimulation of specific enzymes involved in the different biosynthetic pathways of the secondary metabolites. Therefore, a comprehensive study of the impact of UV-B and heavy metals individually and in combination on the biosynthesis and accumulation of secondary metabolites in medicinal plants is discussed in the present review.

Effect of carbon quantum dots derived from extracts of UV-B-exposed Eclipta alba on alcohol-induced liver cirrhosis in Golden Hamster.

The Eclipta alba plant is considered hepatoprotective, owing to its phytoconstituents wedelolactone. In the current study, effect of elevated ultraviolet-B (eUV-B) radiation was investigated on biochemical, phytochemical, and antioxidative enzymatic activities of E. alba (Bhringraj) plant. The UV-B exposure resulted in an increase in oxidative stress, which has caused an imbalance in phytochemical, biochemical constituents, and induced antioxidative enzymatic activities. It was observed that the UV-B exposure promoted wedelolactone yield by 23.64%. Further, the leaf extract of UV-B-exposed plants was used for the synthesis of carbon quantum dots (CQDs) using low cost, one-step hydrothermal technique and its biocompatibility was studied using in vitro MTT (3-[4,5-dimethylthiazol-2-yl]-2,5 diphenyl tetrazolium bromide) assay on HepG2 liver cell line. It revealed no toxicity in any treatment groups in comparison to the control. Both CQDs and leaf extract were orally administered to the golden hamster suffering from alcohol-induced liver cirrhosis. In the morphometric study, it was clearly observed that a combination of UV-B-exposed leaf extract and synthesized CQDs delivered the best result with maximum recovery of liver tissues. The present study reveals the positive impact of UV-B exposure on the medicinally important plant, increased yield of wedelolactone, and its enhanced hepatoprotective efficacy for the treatment of damaged liver tissues.

Photosynthetic, Biochemical and Secondary Metabolite Changes in a Medicinal Plant Chlorophytum borivillianum (Safed musli) against Low and High Doses of UV-B Radiation.

Plants are inevitably grown in presence of sunlight, therefore bound to be exposed to natural UV-B radiation. Several studies have already been conducted with UV-B and medicinal plants and only few studies showed dose dependent variation. The present study aims to find out the variations and adaptation in Chlorophytum borivillianum under two different doses of UV-B radiation; ambient + low (3.2 kJm-2 d-1 ) and high (7.2 kJm-2 d-1 ) UV-B dose, denoted as LD and HD, respectively. Reduction in photosynthetic rate was higher at HD, while plants receiving LD displayed nonsignificant variation. During vegetative and reproductive stage, significant reduction (P ≤ 0.001) in stomatal conductance was obtained when exposed to HD-eUV-B. Fv /Fm showed more reductions in HD-eUV-B (12.6%) followed by LD-eUV-B (7.9%). Low and high doses of UV-B enhanced the anthocyanin content but the increase was significant in HD, indicates epidermal protection strategy by the plants. Under LD-eUV-B, the content of saponin, a major phytochemical constituent was enhanced by 26%. Phytochemical analysis of roots revealed reduction mostly in fatty acid components whereas the steroidal components (stigmasterol and sarsasapogenin) showed enhancement in response to LD. The study suggests the importance of LD-eUV-B in the stimulation of medicinal compounds in C. borivillianum.

Individual and combined effects of chromium and ultraviolet-B radiation on defense system, ultrastructural changes, and production of secondary metabolite psoralen in a medicinal plant Psoralea corylifolia L.

The present study focuses on the effects of individual and combined stress of chromium (Cr) and ultraviolet-B (UV-B) radiation on Psoralea corylifolia L. The experiment comprised four sets: (i) control, (ii) eUV-B (elevated UV-B i.e., ambient + 7.2 kJ m-2 day-1 UV-B), (iii) Cr (chromium; 30 mg kg-1 soil), and (iv) Cr + eUV-B (chromium and elevated UV-B; Cr 30 mg kg-1 and ambient + 7.2 kJ m-2 day-1 UV-B). The eUV-B and Cr individually and in combination showed the variable responses on ultrastructure, physiology and biomass however, the impact was more prominent under individual Cr treatment followed by Cr + eUV-B and eUV-B. Higher bioconcentration factor and the lowered translocation factor consequently led to a higher reduction in the below ground biomass and the lesser reduction in above ground biomass under Cr + eUV-B treatment as compared to individual Cr treatment. In addition, higher induction in the enzymatic (glutathione reductase, ascorbate peroxidase, superoxide dismutase, and glutathione-S-transferase) and non-enzymatic antioxidants (glutathione reduced) were found to be responsible for efficient scavenging of hydrogen peroxide and superoxide radical leading to lowered MDA content under combined treatment as compared to Cr treatment. Deposition of Cr as electron dense granules in the cytoplasm, vacuoles, and cell wall under Cr and Cr + eUV-B is contemplated as one of the cellular mechanisms of P. corylifolia against the toxicity of Cr. Psoralen increased under all treatments with a maximum increase under Cr + eUV-B treatment. Taken together our results accentuated that P. corylifolia can be grown in an area contaminated with Cr and has a higher influx of UV-B for the attainment of psoralen considering its pharmaceutical perspectives.

An assessment of dose-dependent UV-B sensitivity in Eclipta alba: Biochemical traits, antioxidative properties, and wedelolactone yield.

The UV-B-induced signals play a crucial role in improving the analeptic values of medicinally important plants. Eclipta alba L. (Hassak), commonly known as False Daisy, holds supreme stature with its pharmaceutical association in treating various ailments, particularly in Ayurvedic medicine. The present study aimed to evaluate the response of E. alba plants exposed to ambient (AT) and two different supplemental UV-B doses (eUV-B, ambient ±7.2 kJ m-2 day-1), i.e., intermittent (IT) and continuous (CT) UV-B treatment for 130 and 240 h respectively. Antioxidative activities and medicinally important compounds (wedelolactone) were measured in different plants' parts at three growth stages. Under both the eUV-B treatments, the photosynthetic pigments were adversely affected (along with reductions in protein content) with a concomitant increase in secondary metabolites. Substantial variations in enzymatic antioxidants and non-enzymatic compounds showed the adaptive resilience strategies of plants against eUV-B. The wedelolactone content increased in leaves but compromised in stem and roots under IT. The results concluded that IT UV-B exposure led to the improvement of plant growth and the yield of wedelolactone compared to CT, suggesting its ameliorative role in improving the test plant's medicinal value.

Ultraviolet-B mediated biochemical and metabolic responses of a medicinal plant Adhatoda vasica Nees. at different growth stages.

In the present study, the effects of elevated UV-B (eUVB; ambient ± 7.2 kJ m-2 day-1) were evaluated on the biochemical and metabolic profile of Adhatoda vasica Nees. (an indigenous medicinal plant) at different growth stages. The results showed reduction in superoxide radical production rate, whereas increase in the content of hydrogen peroxide which was also substantiated by the histochemical localization. Malondialdehyde content, which is a measure of oxidative stress, did not show significant changes at any of the growth stages however photosynthetic rate and chlorophyll content showed reduction at all growth stages under eUV-B exposure. Increased activities of the enzymatic and non-enzymatic antioxidants were noticed except ascorbic acid, which was reduced under eUV-B exposure. The metabolic profile of A. vasica revealed 43 major compounds (assigned under different classes) at different growth stages. Triterpenes, phytosterols, unsaturated fatty acids, diterpenes, tocopherols, and alkaloids showed increment, whereas reduction in saturated fatty acids and sesquiterpenes were observed under eUV-B treatment. Vasicinone and vasicoline, the two important alkaloids of A. vasica, showed significant induction under eUV-B exposure as compared to control. Treatment of eUV-B leads to the synthesis of some new compounds, such as oridonin oxide (diterpene) and α-Bisabolol oxide-B (sesquiterpene), which possess potent anti-inflammatory and anticancerous activities. The study displayed that differential crosstalk between antioxidants and secondary metabolites at different growth stages, were responsible for providing protection to A. vasica against eUV-B induced oxidative stress and enhancing its medicinal properties.

An assessment of growth, floral morphology, and metabolites of a medicinal plant Sida cordifolia L. under the influence of elevated ozone.

Tropospheric ozone (O3) is a major secondary air pollutant and greenhouse gas, and its impact on growth, yield, and its quality is well established in the case of crop plants. However, the effects of tropospheric O3 have not been comprehensively studied on medicinal plants. Therefore, a field study was planned on a medicinally important Sida cordifolia L. plant (commonly known as country mallow or Bala) to assess the expected changes on the morphology, growth, and leaf injury under elevated O3 (ambient + 20 ppb) by using open-top chambers (OTCs) at 30, 60, and 90 days after treatment (DAT), while leaf and root metabolites were observed at 60 DAT. At all the growth stages, significant leaf damage was recorded as foliar injury symptoms. Most of the growth parameters also showed significant reductions at all the growth stages. Plants under elevated O3 showed a significant negative impact on most of the reproductive parts of the plant. Leaf weight ratio (LWR) showed significant increment at early stages while reduced at 90 DAT; however, root shoot ratio (RSR) showed a significant reduction at 60 DAT. The majority of the steroid metabolites showed an increase in root and leaves under elevated O3, while terpenes showed variable response. Due to O3 stress, most of the major metabolites showed an increase possibly due to their role in defense and other metabolic activities. Based on the outcomes, it is concluded that the future increase in the levels of tropospheric O3 will impact a significant effect on important metabolites of medicinal plants growing in tropical countries like India.

Effect on essential oil components and wedelolactone content of a medicinal plant Eclipta alba due to modifications in the growth and morphology under different exposures of ultraviolet-B.

In the present study sensitivity of a medicinal plant Eclipta alba L. (Hassk) (False daisy) was assessed under intermittent (IT) and continuous (CT) doses of elevated ultraviolet-B (eUV-B). Eclipta alba is rich in medicinally important phytochemical constituents, used against several diseases. The hypothesis of this study is that alterations in UV-B dose may modify the quantity and quality of medicinally valuable components with changes in the morphological and physiological parameters of test plant. To fulfill our hypothesis IT and CT of eUV-B (ambient ± 7.2 kJ m-2 day-2) was given for 130 and 240 h respectively to assess the impact of UV-B stress. Growth and physiological parameters were adversely affected under both the treatments with varying magnitude. The observation of leaf surfaces showed increase in stomatal and trichome densities suggesting the adaptive resilience of the plants against UV-B. Besides, biosynthesis of wedelolactone, a major medicinal compound of E. alba was observed to be stimulated under UV-B exposure. The essential oil content was reduced under IT while increased under CT. A total of 114 compounds were identified from oil extract of E. alba. n-Pentadecane (25.79%), n-Octadecane (12.98%), ß-Farnesene (9.43%), α-Humulene (4.95%) (E)-Caryophyllene (4.87%), Phytol (4.25%), α-Copaene (2.26%), Humulene epoxide (1.46%), ß-Pinene (1.07) and ß-Caryophyllene oxide (1.06%) were identified as major components of oil. CT induced the synthesis of some medicinally important compounds such as α-terpineol, δ-cadinene, linolenic acid, methyl linoleate and myristic acid amide. Hence, the study revealed that continuous UV-B exposure of low intensity could be helpful for commercial exploitation of essential oil in E. alba.

Differential response of radish plants to supplemental ultraviolet-B radiation under varying NPK levels: chlorophyll fluorescence, gas exchange and antioxidants.

Current and projected increases in ultraviolet-B (UV-B; 280-315 nm) radiation may alter crop growth and yield by modifying the physiological and biochemical functions. This study was conducted to assess the possibility of alleviating the negative effects of supplemental UV-B (sUV-B; 7.2 kJ m⁻² day⁻¹; 280-315 nm) on radish (Raphanus sativus var Pusa Himani) by modifying soil nitrogen (N), phosphorus (P) and potassium (K) levels. The N, P and K treatments were recommended dose of N, P and K, 1.5 times recommended dose of N, P and K, 1.5 times recommended dose of N and 1.5 times recommended dose of K. Plants showed variations in their response to UV-B radiation under varying soil NPK levels. The minimum damaging effects of sUV-B on photosynthesis rate and stomatal conductance coupled with minimum reduction in chlorophyll content were recorded for plants grown at recommended dose of NPK. Flavonoids increased under sUV-B except in plants grown at 1.5 times recommended dose of N. Lipid peroxidation (LPO) also increased in response to sUV-B at all NPK levels with maximum at 1.5 times recommended dose of K and minimum at recommended dose of NPK. This study revealed that sUV-B radiation negatively affected the radish plants by reducing the photosynthetic efficiency and increasing LPO. The plants grown at 1.5 times recommended dose of NPK/N/K could not enhance antioxidative potential to the extent as recorded at recommended dose of NPK and hence showed more sensitivity to sUV-B.

Effects of fly ash incorporation on heavy metal accumulation, growth and yield responses of Beta vulgaris plants.

Use of fly ash (FA) to agriculture is not always beneficial, however, utilization of lower concentration of FA as soil amendment is suitable for better management of few crops. The present study was conducted to study the effects of various concentrations of FA (0%, 5%, 10%, 15% and 20%) on heavy metal accumulation, growth, and yield responses of palak (Beta vulgaris L. var All Green H1). The results showed that application of FA caused significant reductions in growth, biomass and yield responses of B. vulgaris plants at different ages of observations. The concentrations of all the heavy metals increased significantly with increasing concentrations of FA. Metal pollution index (MPI) of both roots and shoots showed significant and negative relationships with the yield of B. vulgaris plants. The study concludes that B. vulgaris plant is sensitive to FA concentrations used in this study. It is further recommended that leafy vegetable like B. vulgaris is not a suitable crop to be grown in a region where FA is used for amendment of agricultural soils.