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Industrialization-driven surface ozone (O 3 ) pollution significantly impairs plant growth. This study evaluates the effectiveness of exogenous protectants [3 mg L⁻ 1 abscisic acid (ABA), 400 mg L⁻ 1 ethylenediurea (EDU), and 80 mg L⁻ 1 spermidine (Spd)] on Trifolium repens subjected to O 3 stress in open-top chambers, focusing on plant growth and dynamics of culturable endophytic fungal communities. Results indicate that O 3 exposure adversely affects photosynthesis, reducing root biomass and altering root structure, which further impacts the ability of plant to absorb essential nutrients such as potassium (K), magnesium (Mg), and zinc (Zn). Conversely, the application of ABA, EDU, and Spd significantly enhanced total biomass and chlorophyll content in T. repens. Specifically, ABA and Spd significantly improved root length, root surface area, and root volume, while EDU effectively reduced leaves' malondialdehyde levels, indicating decreased oxidative stress. Moreover, ABA and Spd treatments significantly increased leaf endophytic fungal diversity, while root fungal abundance declined. The relative abundance of Alternaria in leaves was substantially reduced by these treatments, which correlated with enhanced chlorophyll content and photosynthesis. Concurrently, EDU and Spd treatments increased the abundance of Plectosphaerella, enhance the absorption of K, Ca, and Mg. In roots, ABA treatment increased the abundance of Paecilomyces, while Spd treatment enhanced the presence of Stemphylium, linked to improved nitrogen (N), phosphorus (P), and K uptake. These findings suggest that specific symbiotic fungi mitigate O 3 -induced stress by enhancing nutrient absorption, promoting growth. This study highlights the potential of exogenous protectants to enhance plant resilience against O 3 pollution through modulating interactions with endophytic fungal communities., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Masson SAS. All rights reserved.)

Three clover (Trifolium alexandrium L.) cultivars (Bundel, Wardan and JHB-146) were assessed for their responses to ambient ozone (O 3 ) with respect to growth, physiological and biochemical parameters at two rural sites (R1 and R2) using ethylenediurea (EDU). EDU solution (300ppm) was applied as soil drench, 10 days after germination (DAG) at an interval of 10 days up to 80 DAG. The average O 3 concentrations were 52.76 and 60.86 ppb at R1 and R2 sites, respectively during the experimental period. Ambient O 3 induced visible symptoms in all the cultivars at both the sites, with more at R2 site having high ambient O 3 levels. Visible injury was observed first in non-EDU treated plants of Wardan at R2 site. Wardan also showed maximum reduction in leaf injury under EDU treatment at both the sites with more at R2. Under EDU treatment, better adaptation to ambient O 3 at initial age of observation and higher acquisition of resources at later ages of observation at both the sites led to better physiological and biochemical adaptations in Wardan. Bundel retained more biomass in shoot as is reflected with higher shoot/root ratio and thus focused more on repair and defense. Shoot/root ratio of JHB-146 did not respond to EDU treatment and thus showed insignificant variations except at initial age of observation at R1 site. This study clearly suggests that Wardan and Bundel are sensitive to ambient O 3 and can be used as bioindicator species in areas having higher O 3 levels using EDU as a research tool., (Copyright © 2017 Elsevier Inc. All rights reserved.)

Ozone (O 3 ) critical levels have been established under the Long-Range Transboundary Air Pollution Convention to assess the risk of O 3 effects in European vegetation. A recent review study has led to the development of O 3 critical levels for annual Mediterranean pasture species using plants growing in well-watered pots at a coastal site and under low levels of competition. However, uncertainties remain in the extrapolation of the O 3 sensitivity of these species under natural conditions. The response of two O 3 -sensitive annual Mediterranean pasture Trifolium species at the coastal site was compared with the response of the same species growing at a continental site, in natural soil and subject to water-stress and inter-specific competition, representing more closely their natural habitat. The slopes of exposure- and dose-response relationships derived for the two sites showed differences in the response to O 3 between sites attributed to differences in environmental growing conditions, growing medium and the level of inter-specific competition, but the effect of the individual factors could not be assessed separately. Dose-based O 3 indices partially explained differences due to environmental growing conditions between sites. The slopes showed that plants were more sensitive to O 3 at the continental site, but homogeneity of slopes tests revealed that results from both experimental sites may be combined. Although more experimental data considering complex inter-specific competition situations and the effect of important interactive factors such as nitrogen would be needed, these results confirm the validity of applying the current flux-based O 3 critical level under close to natural growing conditions. The AOT40-based O 3 critical level derived at the coastal site was also considered a suitable risk indicator in close to natural growing conditions in the absence of soil moisture limitations on plant growth.

High sugar ryegrasses (HSG) have been developed to improve the uptake, digestion and nitrogen (N)-utilisation of grazing stock, with the potential to increase production yields and benefit climate by reducing methane (CH4) and nitrous oxide (N2O) emissions from livestock farming. In this study, the effects of tropospheric ozone pollution on the seasonal growth dynamics of HSG pasture mesocosms containing Lolium perenne cv. AberMagic and Trifolium repens cv. Crusader were investigated. Species-specific ozone (O3) dose-response relationships (seasonal means: 35, 41, 47, 51, 59 & 67ppb) based on the Phytotoxic Ozone Dose (PODy) were constructed for above and below ground biomass, injury, N-fixation and forage quality. The dynamics of effects of ozone exposure on HSG pasture changed over the course of a season, with the strongest responses occurring in the first 4-8weeks. Overall, strong negative responses to ozone flux were found for root biomass, root nodule mass and N-fixation rates, and ozone adversely impacted a range of forage quality parameters including total sugar content and relative and consumable food values. These results indicate that increasing ozone pollution could decrease the N-use efficiency and reduce the sugar content of managed pasture, and thereby partially detract from some of the suggested benefits of HSG., (Copyright © 2016 Elsevier B.V. All rights reserved.)

Forage species common to the southern USA Piedmont region, Lolium arundinacea, Paspalum dilatatum, Cynodon dactylon and Trifolium repens, were established in a model pasture system to test the future climate change scenario of increasing ozone exposure in combination with varying rainfall amounts on community structure and nutritive quality. Forages were exposed to two levels of ozone [ambient (non-filtered; NF) and twice ambient (2×) concentrations] with three levels of precipitation (average or ±20% of average) in modified open-top chambers (OTCs) from June to September 2009. Dry matter (DM) yield did not differ over the growing season between forage types, except in primary growth grasses where DM yield was higher in 2× than NF treatment. Primary growth clover decreased in nutritive quality in 2× ozone because of increased concentrations of neutral detergent fibre (NDF), acid detergent fibre (ADF) and acid detergent lignin (ADL). Re-growth clover exhibited the largest decrease in nutritive quality, whereas grasses were not adversely affected in 2× ozone. Re-growth grasses responded positively to 2× ozone exposure, as indicated in increased relative food value (RFV) and percentage crude protein (CP) than NF-exposed re-growth grasses. Effects of precipitation were not significant over the growing season for primary or re-growth forage, except in primary growth grasses where DM yield was higher in chambers with above average (+20%) precipitation. Total canopy cover was significantly higher over the growing season in chambers receiving above average precipitation, but no significant effects were observed with ozone. Results indicate shifts in plant community structure and functioning related to mammalian herbivore herbivory in future climate change scenarios., (© 2015 German Botanical Society and The Royal Botanical Society of the Netherlands.)

Increasing tropospheric ozone (O3) and nitrogen soil availability (N) are two of the main drivers of global change. They both may affect gas exchange, including plant emission of volatiles such as terpenes. We conducted an experiment using open-top chambers to analyze these possible effects on two leguminous species of Mediterranean pastures that are known to have different O3 sensitivity, Ornithopus compressus and Trifolium striatum. O3 exposure and N fertilization did not affect the photosynthetic rates of O. compressus and T. striatum, although O3 tended to induce an increase in the stomatal conductance of both species, especially T. striatum, the most sensitive species. O3 and N soil availability reduced the emission of terpenes in O. compressus and T. striatum. If these responses are confirmed as a general pattern, O3 could affect the competitiveness of these species., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Intraspecific variation in six cultivars of clover Trifolium alexandrinum L., (Bundel, Wardan, JHB-146, Saidi, Fahli, and Mescavi) has been studied with ambient and elevated O3 (ambient + 10 ppb O3) in open top chambers. Significant effect of elevated O3 was detected on different morphological, physiological, and biochemical parameters depicting differential response among the test cultivars. Results showed that the magnitude of O3 induced foliar injury symptoms varied in all the cultivars. Ozone significantly depressed photosynthetic rate, stomatal conductance, and photosynthetic efficiency, although variations were cultivar specific. Ozone treatment diminished total biomass of all the cultivars; reduction was highest in Wardan with least O3 resistance followed by Bundel, JHB-146, Saidi, Mescavi, and Fahli. According to the cumulative sensitive index, variations in the sensitivity showed that two cultivars (Wardan and Bundel) were sensitive to elevated O3, while other three cultivars (Fahli, Saidi, and Mescavi) were resistant, and JHB-146 showed intermediate sensitivity. Therefore, the present study supported the selection of sensitive cultivar of clover as a bioindicator for O3 under Indian conditions for the areas experiencing higher concentrations of O3.

Three ethylene diurea (EDU) concentrations (0, 150 and 300 mg/L) were used to evaluate the negative impact of ozone (O3) on two cultivars of Trifolium repens L. cv. Vardan and Bundel grown under natural field conditions in a suburban area of Varanasi, India. Mean O3 concentrations varied from 30.3 to 46.6 microg/L during the experimental period. Higher photosynthetic pigments and ascorbic acid concentrations were noticed in both EDU-treated cultivars over non-EDU-treated ones, but a reverse trend was found for lipid peroxidation. Growth parameters and biomass also showed increments under EDU treatment of both cultivars. The ratio of variable fluorescence to maximum fluorescence increased significantly in Vardan but not in Bundel upon EDU treatment. Results revealed that EDU concentration of 300 mg/L was more effective to combat the oxidative stress as well as protecting plants from O3 injury symptoms. The test cultivar Vardan is relatively sensitive to O3, thereby can be used as a bioindicator of O3 pollution in areas having higher O3 concentrations. Results also indicated that Bundel has more efficient antioxidant defense system than Vardan and hence was more tolerant to O3 stress.

Plants in Nordic regions can be more ozone sensitive at a given ozone concentration than plants at lower latitudes. A recent study shows that the Nordic summer photoperiod, particularly the dim nighttime light, can increase visible foliar injury and alter leaf transpiration in subterranean clover. Effects of photoperiod on the ozone sensitivity of white and red clover cultivars adapted to Nordic conditions were investigated. Although ozone induced visible foliar injury and leaf transpirational changes in white clover, the effects were independent of photoperiod. In red clover, ozone combined with a long photoperiod with dim nights (8 nights) induced more severe visible injuries than with a short photoperiod. Furthermore, transpirational changes in red clover depended on photoperiod. Thus, a long photoperiod can increase ozone sensitivity differently in clover cultivars with different degrees of adaptation to northern conditions, suggesting that ozone indices used in risk analysis should take this effect into account.

The aim of the study was to assess ambient ozone phytotoxicity in Kyiv (Ukraine) using bioindicator clover plants (Trifolium subterraneum cv. Geraldton) and to test some natural and synthetic antioxidants as ozone protectants. The results obtained showed that ambient ozone concentrations were high enough to cause visible leaf injury in clover. All used substances showed partial ozone protective effect on clover. Water extracts from the leaves of plants, known to contain flavonoids-antioxidants showed weaker ozone protective effect and were less stable in the field conditions than synthetic antioxidants. Among the studied extracts, those from Ocimum basilicum and Tagetes patula were more effective as ozone protectants than the one from Salvia sclarea.

Intra-genus and intra-specific variation and the influence of nitrogen enrichment on net assimilation and stomatal conductance of some annual Trifolium species of Mediterranean dehesa grasslands were assessed under experimental conditions. Also gas exchange rates were compared between some Leguminosae and Poaceae species growing in the field in a dehesa ecosystem in central Spain. The results showed that the previously reported different O3 sensitivity of some Trifolium species growing in pots does not seem to be related to different maximum g(s) values. In addition, no clear differences on gas exchange rates could be attributed to Leguminosae and Poaceae families growing in the field, with intra-genus variation being more important than differences found between families. Further studies are needed to increase the database for developing a flux-based approach for setting O3 critical levels for semi-natural Mediterranean species.

Ozone-sensitive (NC-S clone) and resistant plants (NC-R clone) of Trifolium repens and Centaurea jacea were exposed to moderate ozone concentrations in ambient air. The aim of this study was the investigation of the relation between ozone-sensitivity and leaf concentrations of antioxidants (ascorbic acid, total phenolics and total antioxidant capacity). NC-R clone showed the highest concentrations of antioxidants with 50-70% more ascorbic acid than NC-S. NC-R had about 5 times more ascorbic acid in the young leaves and 9 times more in the old leaves than Centaurea. In a fumigation experiment with acute ozone stress (100 nl L(-1)) the antioxidant levels changed profoundly. The ozone-injured leaves of NC-S had 6-8 times more total phenolics than uninjured leaves. Generally older leaves had lower antioxidant concentrations and were more prone to ozone injury than younger leaves. Ascorbic acid concentrations were closer related to the appearance of visible ozone injury than the other antioxidative parameters.

The effect of acute ozone exposure (150 ppb for 3 h) on two clover species, white clover (Trifolium repens L.) and red clover (Trifolium pratense L.) was investigated through the analysis of 10 different physiological and biochemical parameters. Twenty-four hours after fumigation, visible symptoms of injury on leaves were observed only in red clover, but from the biochemical point of view, both species revealed significant ozone-induced modifications. A decrease in the photosynthetic efficiency as well as an increase in the de-epoxidation index and a decrease in the redox state of ascorbate were detected only in T. pratense leaves; no significant change in pigment content was found in either of the two species. On the other hand, both white and red clover showed, although to different extents, significant decreases in the activities of the antioxidant enzymes peroxidase and ascorbate peroxidase. Multivariate statistical analysis revealed not only that ozone affects both species, but also that they differentially respond to the pollutant, confirming the higher sensitivity of Trifolium pratense to ozone exposure.

The aim of this study was to test and select one or more highly sensitive, specific and environmentally successful Egyptian bioindicator plants for ozone (O3). For that purpose more than 30 Egyptian species and cultivars were subjected to extensive screening studies under controlled environmental and pollutant exposure conditions to mimic the Egyptian environmental conditions and O3 levels in urban and rural sites. Four plant species were found to be more sensitive to O3 than the universally used O3-bioindicator, tobacco Bel W3, under the Egyptian environmental conditions used. These plant species, jute (Corchorus olitorius c.v. local), clover (Trifolium alexandrinum L. c.v. Masry), garden rocket (Eruca sativa c.v. local) and alfalfa (Medicago sativa L. c.v. local), ranked in order of decreasing sensitivity, exhibited typical O3 injury symptoms faster and at lower 03 concentrations than Bel W3. Three variables were tested in search of a reliable tool for the diagnosis and prediction of O3 response prior to the appearance of visible foliar symptoms: pigment degradation, stomatal conductance (g(s)) and net photosynthetic CO2 assimilation (Pnet). Pigment degradation was found to be unreliable in predicting species sensitivity to O3. Evidence supporting stomatal conductance involvement in 03 tolerance was found only in tolerant species. A good correlation was found between g(s), restriction of O3 and CO2 influx into the mesophyll tissues, and Pnet. Changes in Pnet seemed to depend largely on fluctuations in g(s).