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Photosynthesis provides carbon sources and energy for crop growth and development, and the widespread presence of microplastics and plastic plasticisers in agricultural soils affects crop photosynthesis, but the mechanism of the effect is not clear. This study aims to investigate the effects of different microplastics and plasticizers on cucumber photosynthesis. Using polyvinyl chloride (PVC), polyethylene (PE), polystyrene (PS), and di-n-octyl phthalate (DOP) as representative microplastics and plasticizers, we assessed their impact on cucumber photosynthesis. Our results reveal significant alterations in key parameters: intercellular CO 2 concentration (Ci) and transpiration rate (Tr) increased across all treatments, whereas stomatal limit value (Ls) and water use efficiency (WUE) decreased. Notably, PS + DOP treatment led to a significant reduction in the maximum efficiency of photosystem II (F v /F m ) and ATP accumulation. Furthermore, PE and PS + DOP treatments decreased lycopene and ɛ-carotene synthesis rates, as well as abscisic acid (ABA) accumulation. All treatments inhibited the conversion of β-carotene into strigolactone (SL) and decreased chlorophyll synthesis rates, with PS + DOP exhibiting the most severe impact. Regarding chlorophyll degradation pathways, PVC and PE treatments reduced chlorophyll decomposition rates, whereas DOP with PS promoted degradation. PE and PS treatments also impaired light energy capture, electron transport, and the structural stability of photosystems I and II, as well as photosynthetic capacity and NADPH and ATP synthesis rates. Our findings underscore the differential impacts of microplastics and plasticizers on cucumber photosynthesis, with PS + DOP having the most detrimental effect. These results shed light on the complex interactions between microplastics and plant physiology, highlighting the urgent need for mitigation strategies in agricultural practices to safeguard crop productivity and environmental sustainability., 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 B.V. All rights reserved.)

This study investigated the effects of corn cob biochar (CCB) and rice husk biochar (RHB) additions (at 0%, 5%, and 10% w/w) on nitrogen and carbon dynamics during co-composting with poultry litter, rice straw, and domestic bio-waste. The study further assessed the temperature, moisture, pH, and nutrient contents of the mature biochar co-composts, and their potential phytotoxicity effects on amaranth, cucumber, cowpea, and tomato. Biochar additions decreased NH 4 + -N and NO 3 - contents, but bacteria and fungi populations increased during the composting process. The mature biochar co-composts showed higher pH (9.0-9.7), and increased total carbon (24.7-37.6%), nitrogen (1.8-2.4%), phosphorus (6.5-8.1 g kg -1 ), potassium (26.8-42.5 g kg -1 ), calcium (25.1-49.5 g kg -1 ), and magnesium (4.8-7.2 g kg -1 ) contents compared to the compost without biochar. Germination indices (GI) recorded in all the plants tested with the different composts were greater than 60%. Regardless of the biochar additions, all composts treatments showed no or very minimal phytotoxic effects on cucumber, amaranth and cowpea seeds. We conclude that rice husk and corn cob biochar co-composts are nutrient-rich and safe soil amendment for crop production., (© 2024. The Author(s).)

Context: This study addresses the development of sustainable pyridinium ionic liquids (ILs) because of their potential applications in agriculture and pharmaceuticals. Pyridinium-based ILs are known for their low melting points, high thermal stability, and moderate solvation properties. We synthesized three novel pyridinium-based ILs: 1-(2-(isopentyloxy)-2-oxoethyl)pyridin-1-ium chloride, 1-(2-(hexyloxy)-2-oxoethyl)pyridin-1-ium chloride, and 1-(2-(benzyloxy)-2-oxoethyl)pyridin-1-ium chloride. The biological activities of these compounds were evaluated through plant growth promotion, herbicidal, and insecticidal assays. Our results show that the benzyloxy derivative significantly enhances wheat and cucumber growth, whereas the isopentyloxy compound has potent herbicidal effects. Computational methods, including DFT calculations and molecular docking, were applied to understand the structure‒activity relationships (SARs) and mechanisms of action., Methods: The computational techniques involved dispersion-corrected density functional theory (DFT) with the B3LYP functional and the 6-311G** basis set. Grimme's D3 corrections were included to account for dispersion interactions. The calculations were performed via GAMESS-US software. Quantum descriptors of reactivity, such as ionization potential, electron affinity, chemical potential, and electrophilicity index, were derived from the HOMO and LUMO energies. Molecular docking studies were conducted via the CB-Dock server via AutoDock Vina software to predict binding affinities to cancer-related proteins. Petra/Osiris/Molinspiration (POM) analysis was used to predict the drug likeness and other pharmaceutical properties of the synthesized ILs., (© 2024. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Low temperature (LT) is an important environmental factor affecting the growth and yield of plants. Melatonin (MT) can effectively enhance the LT tolerance of cucumber. This study found that LT stress induced the expression of Comt1 (caffeic acid O-methyltransferase 1), with the highest expression being about 2-times that of the control. Meanwhile, the content of MT was found to be roughly 63.16% of that in the control samples. Compared with LT treatment alone, exogenous MT pretreatment upregulated the expression levels of TOR (Target of rapamycin), PIN1 (Pin-formed 1), and YUC4 (YUCCA 4), with maximum upregulations reaching approximately 66.67%, 79.32%, and 42.86%, respectively. These results suggest that MT may modulate the tolerance of cucumber seedlings to LT stress by regulating the expression of TOR, PIN1, and YUC4. In addition, co-treatment with AZD-8055 (a TOR inhibitor) or NPA (N-1-naphthylphthalamic acid, an auxin polar transport inhibitor) and MT attenuated MT-induced resistance to LT stress, leading to higher levels of reactive oxygen species (ROS), reduced antioxidant defense capacity, and increased damage to the membrane system in cucumber seedlings. Concurrently, the content of osmoregulatory substances and the photosynthesis decreased. These results demonstrate that both TOR and auxin were required for MT to alleviate LT-induced damage in cucumber. In summary, the present study demonstrates that TOR and auxin signaling synergistically contribute to alleviating LT damage in cucumber seedlings by exogenous MT. These findings help us understand the function of MT and provide insights into the regulatory network of MT that regulates the LT tolerance of plants., 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.)

Copper (Cu) toxicity in crops is a result of excessive release of Cu into environment. Little is known about mitigation of Cu toxicity through the application of carbon-based nanomaterials including water-soluble fullerene C 60 derivatives. Two derivatives of fullerene were examined: polyhydroxylated C 60 (fullerenol) and arginine C 60 derivative. In order to study the response of Cu-stressed plants (Cucumis sativus L.) to these nanomaterials, metabolomics analysis by gas chromatography-mass spectrometry (GC-MS) was performed. Excess Cu (15 μM) caused substantial increase in xylem sap Cu, retarded dry biomass and leaf chlorosis of hydroponically grown cucumber. In Cu-stressed leaves, metabolomes was disturbed towards suppression metabolism of nitrogen (N) compounds and activation metabolism of hexoses. Also, upregulation of some metabolites involving in antioxidant defense system, such as ascorbic acid, tocopherol and ferulic acid, was occurred in Cu-stressed leaves. Hydroponically added fullerene adducts decreased the xylem sap Cu and alleviated Cu toxicity with effectiveness has been most pronounced for arginine C 60 derivative. Metabolic responses of plants subjected to high Cu with fullerene derivatives were opposite to that observed under Cu alone. Fatty acids up-regulation (linolenic acid) and antioxidant molecules (tocopherol) down-regulation might indicate that arginine C 60 adduct can alleviate Cu induced oxidative stress. Although fullerenol slightly improved cucumber growth, its effect on metabolic state of Cu-stressed plants was not statistically significant. We suggest that tested fullerene C 60 adducts have a potential to prevent Cu toxicity in plants through a mechanism associated with their capability to restrict xylem transport of Cu from roots to shoot, and to maintain antioxidative properties of plants., 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.)

Melatonin (Mel) is recognized as a prominent plant growth regulator. This study investigated the alleviating effect of Mel pretreatment on growth inhibition caused by low-temperature (LT) stress (10 °C/6 °C) in cucumber seedlings and explored the role of the Ca 2+ /Calcium-dependent protein kinases (CPKs) signaling pathway in Mel-regulated LT tolerance. The main results are as follows: compared to LT treatment alone, 100 μM Mel increased both the content of Ca 2+ (highest about 42.01%) and the expression levels of Ca 2+ transporter and cyclic nucleotide-gated channel (CNGC) genes under LT. Similarly, Mel enhanced the content of CPKs (highest about 27.49%) and the expression levels of CPKs family genes in cucumber leaves under LT. Additionally, pretreatment with 100 μM Mel for three days strengthened the antioxidant defense and photosynthesis of seedlings under LT. Genes in the ICE-CBF-COR pathway and the MAPK cascade were upregulated by Mel, with maximum upregulations reaching approximately 2.5-fold and 1.9-fold, respectively, thus conferring LT tolerance to cucumber seedlings. However, the above beneficial effects of Mel were weakened by co-treatment with calcium signaling blockers (LaCl 3 or EGTA) or CPKs inhibitors (TFP or W-7), suggesting that the Ca 2+ /CPKs pathway is involved in the Mel-mediated regulation of LT tolerance. In conclusion, this study revealed that Mel can alleviate growth inhibition in cucumber seedlings under LT stress and demonstrated that the Ca 2+ /CPKs signaling pathway is crucial for the Mel-mediated enhancement of LT tolerance. The findings hold promise for providing theoretical insights into the application of Mel in agricultural production and for investigating its underlying mechanisms of action., 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. Published by Elsevier Masson SAS.)

In this paper, we discussed the physiological mechanism of enhanced chilling tolerance with combined treatment of nitric oxide (NO) and reduced glutathione (GSH) in cucumber seedlings. With prolonged low temperature (10 °C/6 °C), oxidative stress improved, which was manifested as an increase the hydrogen peroxide (H 2 O 2 ) and malondialdehyde (MDA), causing cell membrane damage, particularly after 48 h of chilling stress. Exogenous sodium nitroprusside (SNP, NO donor) enhanced the activity of nitric oxide synthase NOS-like, the contents of GSH and polyamines (PAs), and the cellular redox state, thus regulating the activities of mitochondrial oxidative phosphorylation components (CI, CII, CIV, CV). However, buthionine sulfoximine (BSO, a GSH synthase inhibitor) treatment drastically reversed or attenuated the effects of NO. Importantly, the combination of SNP and GSH treatment had the best effect in alleviating chilling-induced oxidative stress by upregulating the activities of antioxidant enzyme, including superoxidase dismutase (SOD), catalase (CAT), ascorbate peroxidase (APX) and peroxidase (POD) and improved the PAs content, thereby increased activities of CI, CII, CIII, CIV, and CV. This potentially contributes to the maintenance of oxidative phosphorylation originating from mitochondria. In addition, the high activity of S-nitrosoglutathione reductase (GSNOR) in the combined treatment of SNP and GSH possibly mediates the conversion of NO and GSH to S-nitrosoglutathione. Our study revealed that the combined treatment with NO and GSH to synergistically improve the cold tolerance of cucumber seedlings under prolonged low-temperature stress., 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.)

Nickel phytotoxicity has been attributed, among others, to oxidative stress. However, little is known about Ni-induced phospholipid modifications, including the oxidative ones. Accumulation of reactive oxygen species (ROS), antioxidative enzyme activities, malondialdehyde and the early lipid oxidation products contents, membrane permeability, phospholipid profile as well as phospholipid unsaturation degree were studied in the 1st and the 2nd leaves of hydroponically grown cucumber seedlings subjected to Ni stress. Compared to the 2nd leaf the 1st one showed stronger visual Ni toxicity symptoms, higher Ni, O 2 .- and H 2 O 2 accumulation as well as greater enhancement in membrane permeability. Enzyme activities were differently influenced by Ni stress, however most pronounced changes were generally found in the 1st leaf. Ni treatment resulted in oxidation of leaf lipids, which was evidenced by appearance of increased contents of MDA and the early produced oxylipins. Among the latter 9-hydroxyoctadecatrienoic acid (9-HOTrE) and 13-hydroxyoctadecatrienoic acid (13-HOTrE) contents showed the most pronounced increase in response to Ni treatment. Exposure to the metal led to the changes in the leaf phospholipid profile and increased degree of phospholipid unsaturation. The obtained results have been discussed in relation to the difference in Ni stress severity between the 1st and the 2nd leaves., 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 The Authors. Published by Elsevier B.V. All rights reserved.)

Salinity stress poses a significant treat to crop yields and product quality worldwide. Application of a humic acid bio stimulant and grafting onto tolerant rootstocks can both be considered sustainable agronomic practices that can effectively ameliorate the negative effects of salinity stress. This study aimed to assess the above mentioned ameliorative effects of both practices on cucumber plants subjected to saline environments. To attain this goal a factorial experiment was carried out in the form of a completely randomized design with three replications. The three factors considered were (a) three different salinity levels (0, 5, and 10 dS m -1 of NaCl), (b) foliar application of humic acid at three levels (0, 100, and 200 mg L -1 ), and (c) both grafted and ungrafted plants. Vegetative traits including plant height, fresh and dry weight and number of leaf exhibited a significant decrease under increasing salinity stress. However, the application of humic acid at both levels mitigated these effects compared to control plants. The reduction in relative water content (RWC) of the leaf caused by salinity, was compensated by the application of humic acid and grafting. Thus, the highest RWC (86.65%) was observed in grafting plants with 0 dS m -1 of NaCl and 20 mg L -1 of humic acid. Electrolyte leakage (EL) increased under salinity stress, but the application of humic acid and grafting improved this trait and the lowest amount of EL (26.95%) was in grafting plants with 0 dS m -1 of NaCl and 20 mg L -1 of humic acid. The highest amount of catalase (0.53 mmol H 2 O 2 g -1  fw min -1 ) and peroxidase (12.290 mmol H 2 O 2 g -1  fw min -1 ) enzymes were observed in the treatment of 10 dS m -1 of NaCl and 200 mg L -1 humic acid. The highest amount of total phenol (1.99 mg g -1 FW), total flavonoid (0.486 mg g -1 FW), total soluble carbohydrate (30.80 mg g -1 FW), soluble protein (34.56 mg g -1 FW), proline (3.86 µg g -1 FW) was in grafting plants with 0 dS m -1 of NaCl and 200 mg L -1 of humic acid. Phenolic acids and phenylalanine ammonia lyase (PAL) and polyphenol oxidase (PPO) enzymes increased with increasing salinity and humic acid levels. Contrary to humic acid, salt stress increased the sodium (Na + ) and chlorine (Cl - ) and decreased the amount of potassium (K + ) and calcium (Ca 2+ ) in the root and leaf of ungrafted cucumber. However, the application 200 mg L -1 humic acid appeared to mitigate these effects, thereby suggesting a potential role in moderating physiological processes and improving growth of cucumber plants subjected to salinity stress. According to the obtained results, spraying of humic acid (200 mg L -1 ) and the use of salt resistant rootstocks are recommended to increase tolerance to salt stress in cucumber. These results, for the first time, clearly demonstrated that fig leaf gourd a new highly salt-tolerant rootstock, enhances salt tolerance and improves yield and quality of grafted cucumber plants by reducing sodium transport to the shoot and increasing the amount of compatible osmolytes., (© 2024. The Author(s).)

Avobenzone (AVO) is a sunscreen with high global production and is constantly released into the environment. Incorporating sewage biosolids for fertilization purposes, the leaching from cultivated soils, and the use of wastewater for irrigation explain its presence in the soil. There is a lack of information about the impact of this sunscreen on plants. In the present study, the ecotoxicity of AVO was tested at concentrations 1, 10, 100, and 1,000 ng/L. All concentrations caused a reduction in root growth of Allium cepa, Cucumis sativus, and Lycopersicum esculentum seeds, as well as a mitodepressive effect, changes in the mitotic spindle and a reduction in root growth of A. cepa bulbs. The cell cycle was disturbed because AVO disarmed the enzymatic defense system of root meristems, leading to an accumulation of hydroxyl radicals and superoxides, besides lipid peroxidation in cells. Therefore, AVO shows a high potential to cause damage to plants and can negatively affect agricultural production and the growth of non-cultivated plants., (© 2024. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

This study explores the impact of juglone on cucumber (Cucumis sativus cv. Beith Alpha), scrutinizing its effects on seed germination, growth, and the polyphenol oxidase (PPO) enzyme's activity and gene expression. Employing concentrations ranging from 0.01 to 0.5 mM, we found juglone's effects to be concentration-dependent. At lower concentrations (0.01 and 0.1 mM), juglone promoted root and shoot growth along with germination, whereas higher concentrations (0.25 and 0.5 mM) exerted inhibitory effects, delineating a threshold for its allelopathic influence. Notably, PPO activity surged, especially at 0.5 mM in roots, hinting at oxidative stress involvement. Real-time PCR unveiled that juglone modulates PPO gene expression in cotyledons, peaking at 0.1 mM and diminishing at elevated levels. Correlation analyses elucidated a positive link between juglone-induced root growth and cotyledon PPO gene expression but a negative correlation with heightened root enzyme activity. Additionally, germination percentage inversely correlated with root PPO activity, while PPO activities positively associated with dopa and catechol substrates in both roots and cotyledons. Molecular docking studies revealed juglone's selective interactions with PPO's B chain, suggesting regulatory impacts. Protein interaction assessments highlighted juglone's influence on amino acid metabolism, and molecular dynamics indicated juglone's stronger, more stable binding to PPO, inferring potential alterations in enzyme function and stability. Conclusively, our findings elucidate juglone's dose-dependent physiological and biochemical shifts in cucumber plants, offering insights into its role in plant growth, stress response, and metabolic modulation., (© 2024 The Author(s). Physiologia Plantarum published by John Wiley & Sons Ltd on behalf of Scandinavian Plant Physiology Society.)

Background: Environmental pollution by cadmium (Cd) is currently a common problem in many countries, especially in highly industrialised areas. Cd present in the soil can be absorbed by plants through the root system., Aim: The aim of the present study was to investigate the effects of cadmium on the metabolic activity of cucumber plants (Cucumis sativus L.) and the accumulation and distribution of Cd in the organs of the plants., Methods: Cucumber seeds (3 g) were exposed to 0.76, 1.58 or 4.17 mg Cd/L (applied as CdCl 2 solutions). The activity of selected antioxidant enzymes - glutathione peroxidase (GSH-Px), superoxide dismutase (SOD) and catalase (CAT), lipid peroxidation and the content of photosynthetic pigments were determined in 6-week-old cucumber plants. In addition, intake of Cd has been determined by flame atomic absorption spectrometry (F-AAS)., Results: The results show that the applied cadmium concentrations affected the activity of antioxidant enzymes. An increase in CAT activity and a decrease in SOD activity were observed in all cucumber organs analysed. GSH-Px activity increased in the roots and stems. Surprisingly, GSH-Px activity decreased in the leaves. The level of lipid peroxidation was usually unchanged (the only one statistically significant change was a decrease in the concentration of malondialdehyde in the leaves which was observed after exposure to the highest Cd concentration). The applied Cd concentrations had no effect on the content of photosynthetic pigments. The highest cadmium content was found in the roots of cucumber plants. Cd tends to accumulate in the roots and a small amount was translocated to the stems and leaves, which was confirmed with the translocation factor (TF)., Conclusions: The results indicate that the range of cadmium concentrations used, corresponding to the level of environmental pollution recorded in Europe, effectively activates the antioxidant enzyme system, without intensifying lipid peroxidation or reducing the content of photosynthetic pigments., 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 The Authors. Published by Elsevier GmbH.. All rights reserved.)

Background: Faced with the need to develop new herbicides with modes of action different to those observed for existing agrochemicals, one of the most promising strategies employed by synthetic chemists involves the structural modification of molecules found in natural products. Molecules containing amides, imides, and epoxides as functional groups are prevalent in nature and find extensive application in synthesizing more intricate compounds due to their biological properties. In this context, this paper delineates the synthesis of N-phenylnorbornenesuccinimide derivatives, conducts biological assays, and carries out in silico investigation of the protein target associated with the most potent compound in plant organisms. The phytotoxic effects of the synthesized compounds (2-29) were evaluated on Allium cepa, Bidens pilosa, Cucumis sativus, Sorghum bicolor, and Solanum lycopersicum., Results: Reaction of endo-bicyclo[2.2.1]hept-5-ene-3a,7a-dicarboxylic anhydride (1) with aromatic amines led to the N-phenylnorbornenesuccinic acids (2-11) with yields ranging from 75% to 90%. Cyclization of compounds (2-11) in the presence of acetic anhydride and sodium acetate afforded N-phenylnorbornenesuccinimides (12-20) with yields varying from 65% to 89%. Those imides were then subjected to epoxidation reaction to afford N-phenylepoxynorbornanesuccimides (21-29) with yields from 60% to 90%. All compounds inhibited the growth of seedlings of the plants evaluated. Substance 23 was the most active against the plants tested, inhibiting 100% the growth of all species in all concentrations. Cyclophilin was found to be the enzymatic target of compound 23., Conclusion: These findings suggest that derivatives of N-phenylnorbornenesuccinimide are promising compounds in the quest for more selective and stable agrochemicals. This perspective reinforces the significance of these derivatives as potential innovative herbicides and emphasizes the importance of further exploring their biological activity on weeds. © 2024 Society of Chemical Industry., (© 2024 Society of Chemical Industry.)

Propylparaben (PrP) and dichloropropylparaben (diClPrP) are found in soil worldwide, mainly due to the incorporation of urban sludge in crop soils and the use of non-raw wastewater for irrigation. Studies on the adverse effects of PrP on plants are incipient and not found for diClPrP. PrP and diClPrP were evaluated at concentrations 4, 40, and 400 µg/L for their phytotoxic potential to seeds of Allium cepa (onion), Cucumis sativus (cucumber), Lycopersicum sculentum (tomato), and Lactuca sativa (lettuce), and cytotoxic, genotoxic potential, and for generating oxygen-reactive substances in root meristems of A. cepa bulbs. PrP and diClPrP caused a significant reduction in seed root elongation in all four species. In A. cepa bulb roots, PrP and diClPrP resulted in a high prophase index; in addition, PrP at 400 µg/L and diClPrP at the three concentrations significantly decreased cell proliferation and caused alterations in a significant number of cells. Furthermore, diClPrP concentrations induced the development of hooked roots in onion bulbs. The two chemical compounds caused significant changes in the modulation of catalase, ascorbate peroxidase, and guaiacol peroxidase, disarming the root meristems against hydroxyl radicals and superoxides. Therefore, PrP and diClPrP were phytotoxic and cytogenotoxic to the species tested, proving dangerous to plants., (© 2024. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Silicon nanoparticles (Si NPs) have an eminent role in improving plant yield through improving yield. The present study was conducted to find the effect of Si NPs on plant yield, biochemical attributes, and minerals of different cucumber cultivars. The greenhouse experiment with foliar application of Si NPs (100, 200, and 300 mg L -1 ) was carried out on cucumber cultivars (Emilie, Mirsoltan, Mitio, and Viola). The application of Si NPs at 300 mg L -1 led to the highest fruit yield, with a 17% increase in fruit production compared to the control. Fruit firmness differed by 31% between Emilie and Si NPs at 100 mg L -1 and Mito at 300 mg L -1 . Plants experiencing Si NPs at 300 mg L -1 had the greatest chlorophyll (Chl) a+b. Compared to the other cultivars, Mito had a greater fruit yield and Chl content. The Si NPs increased TSS by 11% while lowering TA by 24% when compared to the control at 300 mg L -1 . Foliar application of Si NPs reduced the value of TSS/TA. The largest value of K was reached in the Mito cultivar with Si NPs at 200 mg L -1 , with a 22% increase in comparison to the control, indicating that Si NPs considerably boosted the K content. The Si NPs at 200 mg L -1 significantly increased leaf N and P in the Mito cultivar by 16 and 50%, respectively. By using agglomerative hierarchy clustering (AHC), Emilie and Mito were located in two separate clusters, whilst Viola and Mirsoltan were grouped in one cluster. In conclusion, Si NPs at 200-300 mg L -1 enhanced fruit yield, and Mito showed the highest yield when compared to other cultivars., (© 2024. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

The development of cost-effective and eco-friendly fertilizers is crucial for enhancing iron (Fe) uptake in crops and can help alleviate dietary Fe deficiencies, especially in populations with limited access to meat. This study focused on the application of MgFe-layered double hydroxide nanoparticles (MgFe-LDHs) as a potential solution. We successfully synthesized and characterized MgFe-LDHs and observed that 1-10 mg/L MgFe-LDHs improved cucumber seed germination and water uptake. Notably, the application of 10 mg/L MgFe-LDHs to roots significantly increased the seedling emergence rate and growth under low-temperature stress. The application of 10 mg/L MgFe-LDHs during sowing increased the root length, lateral root number, root fresh weight, aboveground fresh weight, and hypocotyl length under low-temperature stress. A comprehensive analysis integrating plant physiology, nutrition, and transcriptomics suggested that MgFe-LDHs improve cold tolerance by upregulating SA to stimulate CsFAD3 expression, elevating GA 3 levels for enhanced nitrogen metabolism and protein synthesis, and reducing levels of ABA and JA to support seedling emergence rate and growth, along with increasing the expression and activity of peroxidase genes. SEM and FTIR further confirmed the adsorption of MgFe-LDHs onto the root hairs in the mature zone of the root apex. Remarkably, MgFe-LDHs application led to a 46% increase (p < 0.05) in the Fe content within cucumber seedlings, a phenomenon not observed with comparable iron salt solutions, suggesting that the nanocrystalline nature of MgFe-LDHs enhances their absorption efficiency in plants. Additionally, MgFe-LDHs significantly increased the nitrogen (N) content of the seedlings by 12% (p < 0.05), promoting nitrogen fixation in the cucumber seedlings. These results pave the way for the development and use of LDH-based Fe fertilizers., (© 2024. The Author(s).)

Salt stress is one of the major factors limiting plant growth and productivity. Many studies have shown that serine hydroxymethyltransferase (SHMT) gene play an important role in growth, development and stress response in plants. However, to date, there have been few studies on whether SHMT3 can enhance salt tolerance in plants. Therefore, the effects of overexpression or silencing of CsSHMT3 gene on cucumber seedling growth under salt stress were investigated in this study. The results showed that overexpression of CsSHMT3 gene in cucumber seedlings resulted in a significant increase in chlorophyll content, photosynthetic rate and proline (Pro) content, and antioxidant enzyme activity under salt stress condition; whereas the content of malondialdehyde (MDA), superoxide anion (H 2 O 2 ), hydrogen peroxide (O 2 ·- ) and relative conductivity were significantly decreased when CsSHMT3 gene was overexpressed. However, the content of chlorophyll and Pro, photosynthetic rate, and antioxidant enzyme activity of the silenced CsSHMT3 gene lines under salt stress were significantly reduced, while MDA, H 2 O 2 , O 2 ·- content and relative conductivity showed higher level in the silenced CsSHMT3 gene lines. It was further found that the expression of stress-related genes SOD, CAT, SOS1, SOS2, NHX, and HKT was significantly up-regulated by overexpressing CsSHMT3 gene in cucumber seedlings; while stress-related gene expression showed significant decrease in silenced CsSHMT3 gene seedlings under salt stress. This suggests that overexpression of CsSHMT3 gene increased the salt tolerance of cucumber seedlings, while silencing of CsSHMT3 gene decreased the salt tolerance. In conclusion, CsSHMT3 gene might positively regulate salt stress tolerance in cucumber and be involved in regulating antioxidant activity, osmotic adjustment, and photosynthesis under salt stress. KEY MESSAGE: CsSHMT3 gene may positively regulate the expression of osmotic system, photosynthesis, antioxidant system and stress-related genes in cucumber., (© 2024. The Author(s), under exclusive licence to Springer Nature B.V.)

Riboflavins are secreted under iron deficiency as a part of the iron acquisition Strategy I, mainly when the external pH is acidic. In plants growing under Fe-deficiency and alkaline conditions, riboflavins have been reported to accumulate inside the roots, with very low or negligible secretion. However, the fact that riboflavins may undergo hydrolysis under alkaline conditions has been so far disregarded. In this paper, we report the presence of riboflavin derivatives and products of their alkaline hydrolysis (lumichrome, lumiflavin and carboxymethylflavin) in nutrient solutions of Cucumis sativus plants grown under different iron regimes (soluble Fe-EDDHA in the nutrient solution, total absence of iron in the nutrient solution, or two different doses of FeSO 4 supplied as a foliar spray), either cultivated in slightly acidic (pH 6) or alkaline (pH 8.8, 10 mM bicarbonate) nutrient solutions. The results show that root synthesis and exudation of riboflavins is controlled by shoot iron status, and that exuded riboflavins undergo hydrolysis, especially at alkaline pH, with lumichrome being the main product of hydrolysis., 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 The Authors. Published by Elsevier Masson SAS.. All rights reserved.)

Strigolactone is a newly discovered type of plant hormone that has multiple roles in modulating plant responses to abiotic stress. Herein, we aimed to investigate the effects of exogenous GR24 (a synthetic analogue of strigolactone) on plant growth, photosynthetic characteristics, carbohydrate levels, endogenous strigolactone content and antioxidant metabolism in cucumber seedlings under low light stress. The results showed that the application of 10 μM GR24 can increase the photosynthetic efficiency and plant biomass of low light-stressed cucumber seedlings. GR24 increased the accumulation of carbohydrates and the synthesis of sucrose-related enzyme activities, enhanced antioxidant enzyme activities and antioxidant substance contents, and reduced the levels of H 2 O 2 and MDA in cucumber seedlings under low light stress. These results indicate that exogenous GR24 might alleviate low light stress-induced growth inhibition by regulating the assimilation of carbon and antioxidants and endogenous strigolactone contents, thereby enhancing the tolerance of cucumber seedlings to low light stress., (© 2022. The Author(s).)

Cucumber is an important vegetable but highly sensitive to salt stress. The present study was designed to investigate the comparative performance of cucumber genotypes under salt stress (50 mmol L -1 ) and stress alleviation through an optimized level of triacontanol @ 0.8 mg L -1 . Four cucumber genotypes were subjected to foliar application of triacontanol under stress. Different physiological, biochemical, water relations and ionic traits were observed to determine the role of triacontanol in salt stress alleviation. Triacontanol ameliorated the lethal impact of salt stress in all genotypes, but Green long and Marketmore were more responsive than Summer green and 20252 in almost all the attributes that define the genetic potential of genotypes. Triacontanol performs as a good scavenger of ROS by accelerating the activity of antioxidant enzymes (SOD, POD, CAT) and compatible solutes (proline, glycinebetaine, phenolic contents), which lead to improved gas exchange attributes and water relations and in that way enhance the calcium and potassium contents or decline the sodium and chloride contents in cucumber leaves. Furthermore, triacontanol feeding also shows the answer to yield traits of cucumber. It was concluded from the results that the salinity tolerance efficacy of triacontanol is valid in enhancing the productivity of cucumber plants under salt stress. Triacontanol was more pronounced in green long and marketer green than in summer green and 20252. Hence, the findings of this study pave the way towards the usage of triacontanol @ 0.8 mg L -1 , and green long and marketer genotypes may be recommended for saline soil., (© 2021. The Author(s).)

As an important gas signaling molecule, hydrogen sulfide (H 2 S) plays a crucial role in regulating cold tolerance. H 2 S cooperates with phytohormones such as abscisic acid, ethylene, and salicylic acid to regulate the plant stress response. However, the synergistic regulation of H 2 S and auxin in the plant response to cold stress has not been reported. This study showed that sodium hydrosulfide (NaHS, an H 2 S donor) treatment enhanced the cold stress tolerance of cucumber seedlings and increased the level of auxin. CsARF5 , a cucumber auxin response factor (ARF) gene, was isolated, and its role in regulating H 2 S-mediated cold stress tolerance was described. Transgenic cucumber leaves overexpressing CsARF5 were obtained. Physiological analysis indicated that overexpression of CsARF5 enhanced the cold stress tolerance of cucumber and the regulation of the cold stress response by CsARF5 depends on H 2 S. In addition, molecular assays showed that CsARF5 modulated cold stress response by directly activating the expression of the dehydration-responsive element-binding (DREB)/C-repeat binding factor (CBF) gene CsDREB3 , which was identified as a positive regulator of cold stress. Taken together, the above results suggest that CsARF5 plays an important role in H 2 S-mediated cold stress in cucumber. These results shed light on the molecular mechanism by which H 2 S regulates cold stress response by mediating auxin signaling; this will provide insights for further studies on the molecular mechanism by which H 2 S regulates cold stress. The aim of this study was to explore the molecular mechanism of H 2 S regulating cold tolerance of cucumber seedlings and provide a theoretical basis for the further study of cucumber cultivation and environmental adaptability technology in winter.

Allicin compositions in garlic are used widely as fungicides in modern agriculture, in which diallyl disulfide (DADS) is a major compound. Downy mildew, caused by Pseudoperonospora cubensis ( P. cubensis ), is one of the most destructive diseases and causes severe yield losses in cucumbers. To explore the potential mechanism of DADS-induced cucumber resistance to downy mildew, cucumber seedlings were treated with DADS and then inoculated with P. cubensis at a 10-day interval. Symptom observation showed that DADS significantly induced cucumber resistance to downy mildew. Furthermore, both lignin and H 2 O 2 were significantly increased by DADS treatment to responding P. cubensis infection. Simultaneously, the enzyme activities of peroxidase (POD) in DADS-treated seedlings were significantly promoted. Meanwhile, both the auxin (IAA) and salicylic acid (SA) contents were increased, and their related differentially expressed genes (DEGs) were up-regulated when treated with DADS. Transcriptome profiling showed that many DEGs were involved in the biological processes of defense responses, in which DEGs on the pathways of 'phenylpropanoid biosynthesis', 'phenylalanine metabolism', 'MAPK signaling', and 'plant hormone signal transduction' were significantly up-regulated in DADS-treated cucumbers uninoculated with the pathogen. Based on the results of several physiological indices and transcriptomes, a potential molecular mechanism of DADS-induced cucumber resistance to downy mildew was proposed and discussed. The results of this study might give new insight into the exploration of the induced resistance mechanism of cucumber to downy mildew and provide useful information for the subsequent mining of resistance genes in cucumber.

Plant fungal diseases can lead to yield reduction and quality degradation in crops, which usually cause serious economic losses. Additionally, chemical fungicides used in the prevention and control of plant diseases are increasingly restricted due to resistance development and high toxicity. Therefore, biogenic fungicides such as chitosan with low toxicity and good biocompatibility are receiving increasing attention. This study found that the acid swelling chitosan pretreatment method can accelerate the rate of the specific oxidation of chitosan catalyzed by the TEMPO-NaBr-NaOCl system. This study proved that OCTS induces plant disease resistance, and the control efficiencies achieved in protection and treatment experiments against Botrytis cinerea were 80.6 % and 83.4 %, respectively, at 400 μg/mL OCTS. In addition, OCTS can promote plant growth and enhance plant defense enzyme activities. This research has realized a forward-looking exploration of the application of OCTS in the agricultural field., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Background: TLPs (Tubby-like proteins) are widespread in eukaryotes and highly conserved in plants and animals. TLP is involved in many biological processes, such as growth, development, biotic and abiotic stress responses, while the underlying molecular mechanism remains largely unknown. In this paper we characterized the biological function of cucumber (Cucumis sativus L.) Tubby-like protein 8 (CsTLP8) in Arabidopsis., Results: In cucumber, the expression of the tubby-like protein CsTLP8 was induced by NaCl treatment, but reduced by PEG (Polyethylene Glycol) and ABA (Abscisic Acid) treatment. Subcellular localization and transcriptional activation activity analysis revealed that CsTLP8 possessed two characteristics of classical transcription factors: nuclear localization and trans-activation activity. Yeast two-hybrid assay revealed interactions of CsTLP8 with CsSKP1a and CsSKP1c, suggesting that CsTLP8 might function as a subunit of E3 ubiquitin ligase. The growth activity of yeast with ectopically expressed CsTLP8 was lower than the control under NaCl and mannitol treatments. Under osmotic and salt stresses, overexpression of CsTLP8 inhibited seed germination and the growth of Arabidopsis seedlings, increased the content of MDA (Malondialdehyde), and decreased the activities of SOD (Superoxide Dismutase), POD (Peroxidase) and CAT (Catalase) in Arabidopsis seedlings. Overexpression of CsTLP8 also increased the sensitivity to ABA during seed germination and ABA-mediated stomatal closure., Conclusion: Under osmotic stress, CsTLP8 might inhibit seed germination and seedling growth by affecting antioxidant enzymes activities. CsTLP8 acts as a negative regulator in osmotic stress and its effects may be related to ABA., (© 2021. The Author(s).)

Exogenous application of methyl jasmonate (MeJA) has been extensively used to study jasmonate-dependent signaling events triggered by biotic stresses. MeJA application leads to complex jasmonate-dependent physiological responses, including changes in stomatal openness and induction of emissions of a multitude of volatile compounds. Whether the alterations in stomatal conductance and emissions of MeJA-induced volatiles are quantitatively associated with MeJA dose, and whether the induced volatile emissions are regulated by modifications in stomatal conductance had been poorly known until recently. Our latest studies highlighted a biphasic kinetics of jasmonate-dependent volatile emissions induced by MeJA treatment in the model species cucumber ( Cucumis sativus ), indicating induction of an immediate stress response and subsequent gene-expression level response. Both the immediate and delayed responses were MeJA dose-dependent. The studies further demonstrated that stomata modulated the kinetics of emissions of water-soluble volatiles in a MeJA dose-dependent manner. These studies contribute to understanding of plant short- and long-term responses to different biotic stress severities as simulated by treatments with a range of MeJA doses corresponding to mild to acute stress.

Biocontrol of soil-borne pathogens by recruiting soil microbiota brings forth benefits to soil quality and plant production while lowers environmental impact. Succinoglycan possesses various biological activities, but its role in soil amendment is still elusive. The succinoglycan Riclin was investigated in this study as a polysaccharide-type biocontrol agent for improving the soil suppressiveness on a typical fungal pathogen Fusarium oxysporum f. sp. cucumerinum (FOC). Results demonstrated that addition of Riclin increased the soil microbial carbon and lowered the species richness of soil fungal communities. After addition of 2.5 mg/g Riclin for 90 days, the relative abundance of Actinobacteria and Firmicutes were increased by 76.6% and 193.4%, compared with the control. Meanwhile, Proteobacteria and Ascomycota were decreased by 25.9% and 30.4%. The relative abundance of beneficial genera, namely Nocardioides, Kribbella, Streptomyces, Gaiella, Marmoricola, Bacillus, and Rhizobium, became 1.13, 5.17, 0.87, 0.45, 3.57, 4.53, and 6.30 folds higher than the control, respectively. Antagonism towards soil-borne pathogens was probably enhanced as both hydrolase activity and biosynthesis of bioactive secondary compounds were improved. Importantly, Riclin-treated soil significantly reduced the incidence of Fusarium wilt of cucumber seedlings by suppression of FOC. In conclusion, addition of Riclin was conducive to the improvement of soil suppressiveness., (Copyright © 2021 Elsevier B.V. All rights reserved.)

The unique properties of carbon-based nanomaterials, including fullerenol, have attracted great interest in agricultural and environmental applications. Iron (Fe) is an essential micronutrient for major metabolic processes, for which a shortage causes chlorosis and reduces the yield of many crops cultivated worldwide. In the current study, the metabolic responses of Cucumis sativus (a Strategy I plant) to fullerenol treatments were investigated depending on the Fe status of plants. Cucumber plants were grown hydroponically, either with [+FeII (ferrous) and +FeIII (ferric)] or in Fe-free (-FeII and -FeIII) nutrient solution, with (+F) or without (-F) a fullerenol supply. Iron species-dependent effects were observed in either Fe-fed or Fe-starved plants, with alteration of metabolites involved in the metabolism of carbohydrates, amino acids, organic acids, lipophilic compounds. Metabolic perturbations triggered by fullerenol in the FeIII-treated plants were in the opposite kind from those in the FeII-treated plants. Whereas in the FeIII-fed plants, fullerenol activated the metabolisation of carbohydrates and amino acids, in the FeII-fed plants, fullerenol activated the metabolisation of lipophilic compounds and repressed the metabolisation of carbohydrates and amino acids. In FeIII-deficient plants, fullerenol stimulated the metabolism of C3 carboxylates and lipophilic compounds while repressing the metabolism of amino acids, hexoses and dicarboxylates, while in FeII-deficient plants, activations of the metabolism of amino acids and dicarboxylates and repression of sterol metabolism by fullerenol were observed. The results indicated that the valence state of Fe sources is of importance for re-programming metabolome responses in cucumber to fullerenol either in Fe-sufficient or Fe-deficient conditions. These investigations are significant for understanding fullerenol interactions and risk assessment in plants with different Fe statuses., Competing Interests: The authors have declared that no competing interests exist.

The aim of this study was to evaluate the effect of acetic (AA) or propionic (PA) acid as a cosurfactant on the microemulsion (ME) characteristics of Thymus vulgaris essential oil (TVO). The results showed that addition of propylene glycol to TVO/AA or PA:T80/water MEs gave dilutable systems with particles ~59 nm in diameter. Plain TVO showed the highest antimicrobial activity against E. coli, S. aureus, and S. typhi in in vitro antimicrobial tests, followed closely by AA/PA-MEs. The antimicrobial activity of AA/PA-MEs used as a washing solution on cucumber and strawberry samples was remarkably greater than those of free TVO, TVO nanoemulsions, and chlorhexidine solutions against E. coli and S. aureus. The sensory properties of the samples were not changed by the use of AA/PA-MEs at 0.05 or 0.1% TVO. The results introduce dilutable TVO:AA/PA-MEs for incorporation of TVO in aqueous systems for use as a fruit/vegetable disinfecting agent., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

In an effort to discover new agents with good fungicidal activities against CDM (cucumber downy mildew), a series of tetrazole derivatives containing phenyloxadiazole moieties were designed and synthesized. The EC 50 values for fungicidal activities against CDM were determined. Bioassay results indicated that most synthesized compounds exhibited potential in vivo fungicidal activity against CDM. A CoMFA (comparative molecular field analysis) model based on the bioactivity was developed to identify some primary structural quality for the efficiency. The values of q 2 and r 2 for the established model were 0.791 and 0.982 respectively, which reliability and predict abilities were verified. Three analogues (q3, q4, q5) were designed and synthesized based on the model. All these compounds exhibited significant fungicidal activity on CDM with the EC 50 of 1.43, 1.52, 1.77 mg·L -1 . This work could provide a useful instruction for the further structure optimization., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Salicylic acid (SA) is an important signal molecule, regulating oxidative stress response in plants. In this study, we evaluated the influences of SA (1 mg L -1 , 10 mg L -1 and 50 mg L -1 ) on the accumulation of clothianidin (CLO), dinotefuran (DFN) and difenoconazole (DFZ) (5 mg L -1 ) and pesticide-induced (CLO-10 mg L -1 , DFN-20 mg L -1 , and DFZ-10 mg L -1 ) oxidative stress in cucumber plants. Exogenous SA at 10 mg L -1 significantly reduced the half-lives of three pesticides in nutrient solution and prevented the accumulation of pesticides in roots and leaves. And the role of SA in reducing residues was related to the major accumulation sites of pesticides. By calculating the root concentration factor (RCF) and translocation factor (TF), we found that SA at 10 mg L -1 reduced the ability of roots to absorb pesticides and enhanced the translocation ability from roots to leaves. Roots exposed to high concentrations of three pesticides could reduce biomass, low chlorophyll content, increase the accumulation of reactive oxygen species (ROS) and proline, promote lipid peroxidation, and alter the activities of a range of antioxidant enzymes, respectively. Exogenous SA at low concentrations (1 mg L -1 and 10 mg L -1 ) significantly mitigated these negative effects. Hence, application of exogenous SA at 10 mg L -1 could effectively alleviate the accumulation of pesticides and induce stress tolerance in cucumber planting systems., (Copyright © 2020 The Authors. Published by Elsevier Inc. All rights reserved.)

Dimethyl disulfide (DMDS), a promising alternative fumigant, has been highly desirable for excellent management of soil pests and diseases. However, high volatility and moderate toxicity of this sulfide limit its application. To address these issues, a novel controlled release formulation of DMDS was proposed employing multiple emulsions and polyurea microcapsules (DMDS@MEs-MCs). The successful combination of the two technologies was revealed by confocal laser scanning microscopy, scanning electron microscopy, thermogravimetric analysis, and Fourier transform infrared. According to the multiple encapsulation structure, the encapsulation efficiency decreased by only 3.13% after thermal storage, compared with a 15.21% decrease of microcapsules made with only a monolayer film. DMDS@MEs-MCs could effectively control the release of active ingredient, which increased applicator and environmental safety during application. Moreover, it could be facilely used by spraying and drip irrigation instead of a special fumigation device. The innovative formulation exhibited better control efficacy on soil pathogens ( Fusarium spp. and Phytophthora spp.) and root-knot nematodes ( Meloidogyne spp.) than DMDS technical concentration (DMDS TC). In addition, it did not inhibit seed germination after 10 days when the plastic film was removed from the fumigated soil. This method appears to be of broad interest for the development of safe and handy fumigant application.

Due to the deliberate use of cupric fungicides in the last century for crop-defence programs, copper (Cu) has considerably accumulated in the soil. The concentrations of Cu often exceed the safety limits of risk assessment for Cu in soil and this may cause toxicity in plants. Copper toxicity induces nutritional imbalances in plants and constraints to plants growth. These aspects might be of paramount importance in the case of phosphorus (P), which is an essential plant macronutrient. In this work, hydroponically grown cucumber plants were used to investigate the influence of the exposure to different Cu concentrations (0.2, 5, 25 and 50 μM) on i) the phenotypic traits of plants, particularly at root level, ii) the nutrient content in both roots and shoots, and iii) the P uptake mechanisms, considering both the biochemical and molecular aspects. At high Cu concentrations (i.e. above 25 μM), the shoot and root growth resulted stunted and the P influx rate diminished. Furthermore, two P transporter genes (i.e. CsPT1.4 and CsPT1.9) were upregulated at the highest Cu concentration, albeit with different induction kinetics. Overall, these results confirm that high Cu concentrations can limit the root acquisition of P, most likely via a direct action on the uptake mechanisms (e.g. transporters). However, the alteration of root plasma membrane permeability induced by Cu toxicity might also play a pivotal role in the observed phenomenon., (Copyright © 2020 Elsevier Masson SAS. All rights reserved.)

TCP proteins are plant-specific transcription factors widely implicated in leaf morphogenesis and senescence, flowering, lateral branching, hormone crosstalk, and stress responses. However, the relationship between the transcription pattern of TCPs and organ development in cucumber has not been systematically studied. In this study, we performed a genome-wide identification of putative TCP genes and analyzed their chromosomal location, gene structure, conserved motif, and transcript expression. A total of 27 putative TCP genes were identified and characterized in cucumber. All 27 putative CsTCP genes were classified into class I and class II. Class I comprised 12 CsTCPs and Class II contained 15 CsTCPs . The 27 putative CsTCP genes were randomly distributed in five of seven chromosomes in cucumber. Four putative CsTCP genes were found to contain putative miR319 target sites. Quantitative RT-PCR revealed that 27 putative CsTCP genes exhibited different expression patterns in cucumber tissues and floral organ development. Transcript expression and phenotype analysis showed that the putative CsTCP genes responded to temperature and photoperiod and were induced by gibberellin (GA)and ethylene treatment, which suggested that CsTCP genes may regulate the lateral branching by involving in multiple signal pathways. These results lay the foundation for studying the function of cucumber TCP genes in the future.

Background: Hydrogen sulfide (H 2 S) is a gas signal molecule involved in regulating plants tolerance to heavy metals stress. In this study, we investigated the role of H 2 S in cadmium-(Cd-) induced cell death of root tips of cucumber seedlings., Results: The results showed that the application of 200 μM Cd caused cell death, increased the content of reactive oxygen species (ROS), chromatin condensation, the release of Cytochrome c (Cyt c) from mitochondria and activated caspase-3-like protease. Pretreatment of seedlings with 100 μM sodium hydrogen sulfide (NaHS, a H 2 S donor) effectively alleviated the growth inhibition and reduced cell death of root tips caused by Cd stress. Additionally, NaHS + Cd treatment could decrease the ROS level and enhanced antioxidant enzyme activity. Pretreatment with NaHS also inhibited the release of Cyt c from the mitochondria, the opening of the mitochondrial permeability transition pore (MPTP), and the activity of caspase-3-like protease in the root tips of cucumber seedling under Cd stress., Conclusion: H 2 S inhibited Cd-induced cell death in cucumber root tips by reducing ROS accumulation, activating the antioxidant system, inhibiting mitochondrial Cyt c release and reducing the opening of the MPTP. The results suggest that H 2 S is a negative regulator of Cd-induced cell death in the root tips of cucumber seedling.

Background: Metallic nanoparticles (NPs) are highly exploited in manufacturing and medical processes in a broad spectrum of industrial applications and in the academic sectors. Several studies have suggested that many metallic nanomaterials including those derived by silver (Ag) are entering the ecosystem to cause significant toxic consequences in cell culture and animal models. However, ecotoxicity studies are still receiving limited attention when designing functionalized and non.-functionalized AgNPs., Objective: This study aimed to investigate different ecotoxicological profiles of AgNPs, which were analyzed in two different states: in pristine form uncoated AgNPs and coated AgNPs with the antimicrobial peptide indolicidin. These two types of AgNPs are exploited for a set of different tests using Daphnia magna and Raphidocelis subcapitata , which are representatives of two different levels of the aquatic trophic chain, and seeds of Lepidium sativum, Cucumis sativus and Lactuca sativa ., Results: Ecotoxicological studies showed that the most sensitive organism to AgNPs was crustacean D. magna, followed by R. subcapitata and plant seeds, while AgNPs coated with indolicidin (IndAgNPs) showed a dose-dependent decreased toxicity for all three., Conclusion: The obtained results demonstrate that high ecotoxicity induced by AgNPs is strongly dependent on the surface chemistry, thus the presence of the antimicrobial peptide. This finding opens new avenues to design and fabricate the next generation of metallic nanoparticles to ensure the biosafety and risk of using engineered nanoparticles in consumer products., Competing Interests: The authors report no conflicts of interest in this work., (© 2020 Falanga et al.)

Key Message: CsGPA1 interacts with CsTIP1.1 (a member of CsAQPs) and suppression of CsGPA1 results the reverse expression of CsAQPs in leaves and roots, resulting in declining water content of cucumber seedlings under salt stress. Salt stress seriously affects cucumber growth and development. Whether the G-protein alpha subunit functions in cucumber during salt stress and its regulation mechanism remains unknown. We interrogated CsGPA1-RNAi lines to identify the role of CsGPA1 during salt stress. Phenotypically, compared with wild type, leaves were severely withered, and root cells showed signs of senescence under salt stress for RNAi lines. Compared with WT, SOD and CAT activity, soluble protein and proline contents all decreased in RNAi lines, while malondialdehyde and relative electrical conductivity increased. Through screening the yeast two-hybrid library and combined with yeast two-hybrid and GST pull-down, the interaction of CsGPA1 with CsTIP1.1 was found the first time in a plant. Then, the expression of aquaporin (AQP) family genes was detected. The expression of CsAQP genes in leaves and roots was primarily up-regulated in WT under salt stress. However, interference by CsGPA1 resulted in enhanced expression of CsAQPs except for CsTIP3.2 in leaves, but reduced expression of some CsAQPs in roots under salt stress. Furthermore, principal component analysis of CsAQP expression profiles and linear regression analysis between CsGPA1 and CsAQPs revealed that CsGPA1 reversely regulated the expression of CsAQPs in leaves and roots under salt stress. Moreover, the water content in leaves and roots of RNAi seedlings significantly decreased compared with WT under salt stress. Overall, CsGPA1 interacts with CsTIP1.1 and suppression of CsGPA1 results in opposite patterns of expression of CsAQPs in leaves and roots, resulting in declining water content of cucumber under salt stress.

The increasing application of various surfactants nowadays, may lead to the contamination of the natural environment and represent potential threat to terrestrial higher plants. In this article, the effect of 13 surfactants, with dodecyl alkyl chain and various aromatic (imidazolium, pyridinium, thiazolium) and aliphatic (guanidinium, ammonium, thiosemicarbazidium) polar heads, on germination, development and growth of wheat and cucumber was investigated. The study aimed to prove how changes in lipophilicity of surfactants and their various structural modifications (existence of the aliphatic or aromatic polar group, the introduction of oxygen and sulfur) influence toxicity towards investigated plants. The calculated lipophilic parameter (AlogP) is shown to be a useful parameter for predicting potential toxicity of the compound. The strategy of using surfactants with aliphatic polar heads instead of aromatic prove to be a promising strategy in reducing harmful effect, as well as the introduction of polar groups in the structure of cation. From all investigated compounds, surfactants with imidazolium polar head displayed the most harmful effect towards wheat and cucumber. The cucumber seeds were more sensitive to the addition of surfactants comparing to wheat. All obtained experimental results were additionally investigated using computational methods, simulating the transport of surfactants through a lipid bilayer. The influence of cation tendency to fit in lipid bilayer structure was correlated with toxicity. For the first time, it is concluded that cation ability to mimic the structure of bilayer have less harmful effect on plant development., 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. The authors declare the following financial interests/personal relationships which may be considered as potential competing interests: Aleksandar Tot., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Microplastics pollution in farmlands has become a major concern. However, few studies have assessed the effects of microplastics on higher plants. In this study, we investigated the influence of polystyrene nanoplastics (PSNPs, 50 mg L -1 ), with four different particle sizes (100, 300, 500, and 700 nm), on the physiological and biochemical indexes of cucumber leaves. The biomass of cucumber plants significantly decreased after exposure to 300 nm PSNPs. Similarly, the chlorophyll a, chlorophyll b, soluble sugar, carotenoid, and proline content, as well as the fluorescence of cucumber leaves were significantly reduced by 100 nm PSNPs. Malondialdehyde, proline, peroxidase gene expression and enzyme activity, and hydrogen peroxide content significantly increased in cucumber leaves exposed to 700 nm PSNPs. In addition, increasing PSNPs particle size led to decreased relative expression levels and activities of the major antioxidant enzymes superoxide dismutase and catalase, while vitamin C and soluble protein content significantly increased. Overall, our results indicated that PSNPs affect the photosynthetic, antioxidant, and sugar metabolism systems of cucumber leaves, with the latter clearly affecting the total biomass of cucumber plants. The benzene ring resulting from the degradation of PSNPs in cucumber leaves may be the main factor affecting chlorophyll metabolism and sugar metabolism. Our findings provide a scientific basis for the risk assessment of PSNPs exposure in soil-plant systems., Competing Interests: Declaration of competing interests Authors declare no conflict of interest., (Published by Elsevier Ltd.)

Main Conclusion: NO was involved in H 2 -induced adventitious rooting by regulating the protein and gene expressions of PM H + -ATPase and 14-3-3. Simultaneously, the interaction of PM H + -ATPase and 14-3-3 protein was also involved in this process. Hydrogen gas (H 2 ) and nitric oxide (NO) have been shown to be involved in plant growth and development. The results in this study revealed that NO was involved in H 2 -induced adventitious root formation. Western blot (WB) analysis showed that the protein abundances of plasma membrane H + -ATPase (PM H + -ATPase) and 14-3-3 protein were increased after H 2 , NO, H 2 plus NO treatments, whereas their protein abundances were down regulated when NO scavenger carboxy-2-(4-carboxyphenyl)-4,4,5,5-tetramethylimidazoline-1-oxyl-3-oxide (cPTI O) was added. Moreover, the mRNA abundances of the HA3 and 14-3-3(7) gene as well as the activities of PM H + -ATPase (EC 3.6.1.35) and H + pump were in full agreement with the changes of protein abundance. Phosphorylation of PM H + -ATPase and the interaction of PM H + -ATPase and 14-3-3 protein were detected by co-immunoprecipitation analysis. H 2 and NO significantly up regulated the phosphorylation of PM H + -ATPase and the interaction of PM H + -ATPase and 14-3-3 protein. Conversely, the stimulation of PM H + -ATPase phosphorylation and protein interaction were significantly diminished by cPTIO. Protein interaction activator fusicoccin (FC) and inhibitor adenosine monophosphate (AMP) of PM H + -ATPase and 14-3-3 were used in this study, and the results showed that FC significantly increased the abundances of PM H + -ATPase and 14-3-3, while AMP showed opposite trends. We further proved the critical roles of PM H + -ATPase and 14-3-3 protein interaction in NO-H 2 -induced adventitious root formation. Taken together, our results suggested that NO might be involved in H 2 -induced adventitious rooting by regulating the expression and the interaction of PM H + -ATPase and 14-3-3 protein.

Waterlogging is a severe environmental stress that causes severe crop productivity losses. Cucumber (Cucumis sativus L.) survives waterlogging by producing adventitious roots (ARs) that enhance gas exchange. Little is known about the role of light and sugars in the waterlogging-induced production of ARs. The role of these factors in AR production was therefore studied in cucumber seedlings grown in the absence or presence of waterlogging and different light conditions. The effect of photosynthesis was studied by removing the shoots of the seedlings and replacing them with exogenous applications of sucrose or stachyose. Shoot removal inhibited AR emergence and elongation. However, the exogenous application of sugars fully restored AR emergence and partially restored root elongation. The exogenous application of a synthetic auxin restored AR emergence but not AR elongation. Transcriptome profiling analysis was used to determine the effects of light on gene expression in the hypocotyls under these conditions. The levels of transcripts encoding proteins involved in auxin transport and signalling were higher in the light and following the exogenous application of sucrose and stachyose. These results show that the waterlogging-induced emergence of ARs is regulated by the interaction between sugars and auxin, whereas AR elongation depends only on sugars alone., (© 2020 John Wiley & Sons Ltd.)

Cucumber mosaic cucumovirus (CMV) is a deadly plant virus that results in crop-yield losses with serious economic consequences. In recent years, environmentally friendly components have been developed to manage crop diseases as alternatives to chemical pesticides, including the use of natural compounds such as glycine betaine (GB) and chitosan (CHT), either alone or in combination. In the present study, the leaves of the cucumber plants were foliar-sprayed with GB and CHT-either alone or in combination-to evaluate their ability to induce resistance against CMV. The results showed a significant reduction in disease severity and CMV accumulation in plants treated with GB and CHT, either alone or in combination, compared to untreated plants (challenge control). In every treatment, growth indices, leaf chlorophylls content, phytohormones (i.e., indole acetic acid, gibberellic acid, salicylic acid and jasmonic acid), endogenous osmoprotectants (i.e., proline, soluble sugars and glycine betaine), non-enzymatic antioxidants (i.e., ascorbic acid, glutathione and phenols) and enzymatic antioxidants (i.e., superoxide dismutase, peroxidase, polyphenol oxidase, catalase, lipoxygenase, ascorbate peroxidase, glutathione reductase, chitinase and β-1,3 glucanase) of virus-infected plants were significantly increased. On the other hand, malondialdehyde and abscisic acid contents have been significantly reduced. Based on a gene expression study, all treated plants exhibited increased expression levels of some regulatory defense genes such as PR1 and PAL1 . In conclusion, the combination of GB and CHT is the most effective treatment in alleviated virus infection. To our knowledge, this is the first report to demonstrate the induction of systemic resistance against CMV by using GB., Competing Interests: The authors declare no conflict of interest.

Fusaric acid (FA), the fungal toxin produced by Fusarium oxysporum , plays a predominant role in the virulence and symptom development of Fusarium wilt disease. As mineral nutrients can be protective agents against Fusarium wilt, hydroponic experiments employing zinc (Zn) and copper (Cu) followed by FA treatment were conducted in a glasshouse. FA exhibited strong phytotoxicity on cucumber plants, which was reversed by the addition of Zn or Cu. Thus, Zn or Cu dramatically reduced the wilt index, alleviated the leaf or root cell membrane injury and mitigated against the FA inhibition of plant growth and photosynthesis. Cucumber plants grown with Zn exhibited decreased FA transportation to shoots and a 17% increase in toxicity mitigation and showed minimal hydrogen peroxide, lipid peroxidation level with the increased of antioxidant enzymes activity in both roots and leaves. Cucumber grown with additional Cu absorbed less FA but showed more toxicity mitigation at 20% compared to with additional Zn and exhibited decreased hydrogen peroxide level and increased antioxidant enzymes activity. Thus, adding Zn or Cu can decrease the toxicity of the FA by affecting the absorption or transportation of the FA in plants and mitigate toxicity possibly through chelation. Zn and Cu modify the antioxidant system to scavenge hydrogen peroxide for suppressing FA induction of oxidative damage. Our experiments could provide a theoretical basis for the direct application of micro-fertilizer as protective agents in farming.

The water-soluble fullerenols are novel carbon-based nanomaterials with unique properties, which afford them with wide agricultural applications. Iron (Fe) deficiency is the most common and widespread nutrition disorder affecting plants. Foliar Fe treatments of plants have been carried out with solutions devoid of fullerenol. In this study, the role of fullerenol [C60(OH)22-24] in alleviation of Fe deficiency in Cucumis sativus (a Strategy I plant) via foliar fertilization was investigated. Cucumber plants were grown hydroponically, either with (Fe) or in Fe-free (-Fe) nutrient solution. The following foliar spray treatments were applied: fullerenol at final concentrations of 1 (F1) and 10 (F10) mg L-1; Fe(II)SO4·7H2O; Fe(II)-EDTA (ethylenediaminetetraacetic acid); and Fe(II)-F1 and Fe(II)-F10. The best used compound was a combination of Fe(II)-sulfate with fullerenol, especially Fe-F1. The addition of fullerenol to Fe(II)-sulfate solutions significantly increased leaf-active Fe (extracted by an Fe(II) chelator) and re-greening at the site of application. The fullerenol-induced mutual influences did not appear when fullerenol was sprayed alone, suggesting a beneficial role of Fe(II)-fullerenol interactions in the penetration of Fe(II) in the leaves and re-greening under Fe-limited conditions. The results are of importance to enhancing the potential of foliar Fe fertilization as the commonly used strategy for ameliorating Fe deficiency and improving crop yield and quality., Competing Interests: The authors have declared that no competing interests exist.

Microcystins (MCs) have become an important global environmental issue, causing oxidative stress, which is an important toxic mechanism for MCs in plants. However, the regulating mechanism of antioxidative enzymes in plants in adapting to MCs stress remains unclear. We studied the dynamic effects of MCs at different concentrations (5, 10, 50 and 100 μg/L) in rice and cucumber seedlings on relative growth rate (RGR), and reactive oxygen species and malondialdehyde (MDA) content, and antioxidative enzyme activities, during a stress period (MCs exposed for 1, 3, 5 and 7 d) and recovery period (7 d). During the stress period, MCs at 5 μg/L inhibited RGR in cucumber and promoted RGR in rice. The contents of superoxide anion (O 2 · - ), hydrogen peroxide (H 2 O 2 ) and MDA increased and RGR declined in both crops with time and intensity of MCs stress. For cucumber, all these parameters responded earlier to MCs stress, and O 2 · - , MDA and RGR were more responsive to MCs stress than in rice. Moreover, catalase (CAT) and peroxidase (POD), and the relative expressions of CAT genes increased in both crops at 5-100 μg/L MCs, whereas relative expression of POD genes increased only in cucumber. Diversely, superoxide dismutase (SOD) response to MCs in cucumber leaves was later than for rice. MCs at 100 μg/L decreased the relative expression of SOD genes in cucumber but did not change SOD activity. During the recovery period, all the above indicators in both crops were higher than the control and lower than in the stress period. Conversely, RGR was lower than in the control and higher than in the stress period, except for cucumber which was lower, and MDA content higher than the stress period at 100 μg/L MCs. Overall, these results indicated that cucumber was more sensitive to MCs than rice, and SOD, CAT and POD play an important role in plant response to MCs stress., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Iron deficiency severely affects crop yield and quality. Gamma-aminobutyric acid (GABA) plays a vital role in plant responses to multifarious stresses. However, the role of GABA in Fe deficiency responses and the potential mechanisms remain largely unknown in cucumber. Here, we found that Fe deficiency raised the GABA levels in leaves and roots of cucumber. To probe the role of GABA in Fe deficiency, the seedlings were subjected to five levels of GABA concentrations (0, 5, 10, 20 and 40 mmol L -1 ) for 7 days under Fe deficiency. The results demonstrated that 20 mM GABA in alleviating the Fe deficiency-induced stress was the most effective. GABA pretreatment reduced the Fe deficiency-induced chlorosis and inhibition of photosynthesis and growth, and significantly enhanced the contents of iron in shoots and roots. Exogenous GABA significantly decreased the pH of nutrient solution and increased ferric-chelate reductase (FCR) activity induced by Fe deficiency and the transcript levels of Fe uptake-related genes HA1, FRO2 and IRT1 in roots. GABA also increased the content of auxin (IAA) and expression of auxin biosynthesis (YUC4), response (IAA1), and transport (PIN1) genes under Fe deficiency. Furthermore, exogenous the auxin transport inhibitor 1-naphthylphthalamic acid (NPA) application abolished the GABA-induced changes in Fe deficiency. In summary, we found that GABA improves tolerance to iron deficiency via an auxin-dependent mechanism in cucumber., Competing Interests: Declaration of competing interest The authors have no conflicts of interest to report., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Background: Brassinolide (BR), as a new type of plant hormones, is involved in the processes of plant growth and stress response. Previous studies have reported the roles of BR in regulating plant developmental processes and also response tolerance to abiotic stresses in plants. The main purpose of our study was to explore whether nitric oxide (NO) plays a role in the process of BR-induced adventitious root formation in cucumber (Cucumis sativus L.)., Results: Exogenous application of 1 μM BR significantly promoted adventitious rooting, while high concentrations of BR (2-8 μM) effectively inhibited adventitious rooting. NO donor (S-nitroso-N-acerylpenicillamine, SNAP) promoted the occurrence of adventitious roots. Simultaneously, BR and SNAP applied together significantly promoted adventitious rooting and the combined effect was superior to the application of BR or SNAP alone. Moreover, NO scavenger (c-PTIO) and inhibitors (L-NAME and Tungstate) inhibited the positive effects of BR on adventitious rooting. BR at 1 μM also increased endogenous NO content, NO synthase (NOS-like) and Nitrate reductase (NR) activities, while BRz (a specific BR biosynthesis inhibitor) decreased these effects. In addition, the relative expression level of NR was up-regulated by BR and SNAP, whereas BRz down-regulated it. The application of NO inhibitor (Tungstate) in BR also inhibited the up-regulation of NR., Conclusion: BR promoted the formation of adventitious roots by inducing the production of endogenous NO in cucumber.

Micronutrients provide a potentially interesting alternative to fungicides for the protection of crops against fungal pathogens. Here we studied the effect of foliar-applied manganese (Mn) in the form of MnSO 4 on severity of anthracnose disease, caused by Colletotrichum lagenarium in cucumber (Cucumis sativus L.) plant. The study was done aimed to characterize the optimum dose and application time of Mn fertilizer on disease suppression as well as to identify the defense mechanisms by which Mn-treated plants resist to fungal disease. In preliminary tests, Mn was applied at different concentrations (1.8, 4.5 and 7.2 mM) and various time points (three days before or two hours before inoculation, or three days after inoculation). Results showed that application of Mn either before or after inoculation suppressed the fungal infection in leaves and cotyledons, with a higher efficiency when applied three days prior to inoculation. However, all applied concentrations of Mn equally reduced the disease severity. Mn treatment in the absence of the pathogen promoted lignification and reactive oxygen species (ROS) accumulation. Also, pre-inoculation Mn treatment enhanced pathogen-induced lignification, callose or ROS production and reduced pathogen-induced cell death. The increase of lignin, callose and ROS induction by Mn application were 34, 30 and 31 % compared to control, respectively. Together, the results suggested the effectiveness of Mn treatments on anthracnose alleviation in cucumber plants. The findings here have a practical importance in plant physiology studies to identify the resistance-relevant mechanisms to pathogens and in sustainable agriculture to control the fungal diseases by a safe method., Competing Interests: Declaration of Competing Interest The authors declare that they have no conflict of interest., (Copyright © 2020 Elsevier GmbH. All rights reserved.)

Cadmium level is continuously increasing in agricultural soils mainly due to anthropogenic activities. Cadmium is one of the most phytotoxic metals in the soils. The present study investigates the possible role of 2-hydroxymelatonin (2-OHMT) in assuagement of Cd-toxicity in cucumber (Cucumis sativus L.) plants. 2-OHMT is an important metabolite produced through interaction of melatonin with oxygenated compounds. Cadmium stress decreased the activity of antioxidant enzymes and polyamines. However, exogenously applied 2-OHMT enhanced plant growth attributes including photosynthetic rate, intercellular CO 2 concentration, stomatal conductance and transpiration rate in treated plants. In addition, 2-OHMT induced enhancement of the activity of PAs biosynthesizing enzymes (putrescine, spermidine and spermine) in conjunction with reduction in activity of polyamine oxidase (PAO). 2-OHMT mitigated Cd stress through up-regulation in expression of stress related CS-ERS gene along with the amplified activity of superoxide dismutase (SOD), catalase (CAT) and ascorbate peroxidase (APX) in treated seedlings. The improved activity of antioxidant scavengers played central role in reduction of hydrogen peroxide (H 2 O 2 ), electrolyte leakage (EL) and malondialdehyde (MDA) in plants under Cd stress. Recent findings also advocate the positive correlation between PAs and ethylene, as both possess common precursor. The current study reveals that priming seeds with 2-OHMT reduces Cd-toxicity and makes it possible to cultivate cucumber in Cd-contaminated areas. Future experiments will perhaps help in elucidation of 2-OHMT intervened stress mitigation procedure in C. sativus crop. Furthermore, research with reference to potential of 2-OHMT for stress alleviation in other horticultural and agronomic crops will assist in enhancement of crop productivity., Competing Interests: Declaration of competing interest The authors declare no conflict of interest., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Sulfate reducing bacteria (SRB) biostabilization has attracted particular attention due to its ability to prevent and control heavy metal pollution. In this study, biotic experiments (immobilisation test of Daphnia (D.) magna, germination experiment of cucumber seeds, and in vitro experiment using gut juices of Sipunculus (S.) nudus) and diffusive gradients in thin films (DGT) technique were performed to investigate the biosafety of cadmium (Cd) contaminated sediments after being treated by indigenous SRB. Results showed that SRB treatment reduced Cd bioaccessibility of sediment to S. nudus, Cd levels in the overlying water and Cd bioavailability to D. magna. However, the treatment increased the biotoxicity of overlying water due to significant reduction in the root length and germination index of cucumber seeds. DGT results confirmed that SRB treatment increased Cd stability in sediment, and reduced its release from the sediment into the overlying water. The biotoxicity of overlying water was not caused by Cd, but possibly by the added culture medium, SRB itself, or its metabolites. More attention is required to assess the safety of SRB treatment when it is used to remediate environmental matrix contaminated by heavy metals., (Copyright © 2019 Elsevier B.V. All rights reserved.)

Treatment by volatile plant hormone methyl jasmonate (MeJA) leads to release of methanol and volatiles of lipoxygenase pathway (LOX volatiles) in a dose-dependent manner, but how the dose dependence is affected by stomatal openness is poorly known. We studied the rapid (0-60 min after treatment) response of stomatal conductance ( G s ), net assimilation rate ( A ), and LOX and methanol emissions to varying MeJA concentrations (0.2-50 mM) in cucumber ( Cucumis sativus ) leaves with partly open stomata and in leaves with reduced G s due to drought and darkness. Exposure to MeJA led to initial opening of stomata due to an osmotic shock, followed by MeJA concentration-dependent reduction in G s , whereas A initially decreased, followed by recovery for lower MeJA concentrations and time-dependent decline for higher MeJA concentrations. Methanol and LOX emissions were elicited in a MeJA concentration-dependent manner, whereas the peak methanol emissions (15-20 min after MeJA application) preceded LOX emissions (20-60 min after application). Furthermore, peak methanol emissions occurred earlier in treatments with higher MeJA concentration, while the opposite was observed for LOX emissions. This difference reflected the circumstance where the rise of methanol release partly coincided with MeJA-dependent stomatal opening, while stronger stomatal closure at higher MeJA concentrations progressively delayed peak LOX emissions. We further observed that drought-dependent reduction in G s ameliorated MeJA effects on foliage physiological characteristics, underscoring that MeJA primarily penetrates through the stomata. However, despite reduced G s , dark pretreatment amplified stress-volatile release upon MeJA treatment, suggesting that increased leaf oxidative status due to sudden illumination can potentiate the MeJA response. Taken together, these results collectively demonstrate that the MeJA dose response of volatile emission is controlled by stomata that alter MeJA uptake and volatile release kinetics and by leaf oxidative status in a complex manner., Competing Interests: The authors declare no competing financial interest.