Arsenic (As) contamination and bioaccumulation are a serious threat to agricultural plants. To address this issue, we checked the efficacy of As tolerant plant growth promoting bacteria (PGPB), zinc oxide nanoparticles (ZnO NPs) and oxalic acid (OA) in Luffa acutangula grown on As rich soil. The selected most As tolerant PGPB i.e Providencia vermicola exhibited plant growth promoting features i.e solubilzation of phosphate, potassium and siderophores production. Innovatively, we observed the synergistic effects of P. vermicola, ZnO NPs (10 ppm) and OA (100 ppm) in L. acutangula grown on As enriched soil (150 ppm). Our treatments both as alone and in combination alleviated As toxicity exhibited by better plant growth and metabolism. Results revealed significantly enhanced photosynthetic pigments, proline, relative water content, total sugars, proteins and indole acetic acid along with As amelioration in L. acutangula. Furthermore, upregulated plant resistance was manifested with marked reduction in the lipid peroxidation and electrolyte leakage and pronounced antagonism of As and zinc content in leaves under toxic conditions. These treatments also improved level of nutrients, abscisic acid and antioxidants to mitigate As toxicity. This marked improvement in plants’ defense mechanism of treated plants under As stress is confirmed by less damaged leaves cell structures observed through the scanning electron micrographs. We also found substantial decrease in the As bioaccumulation in the L. acutangula shoots and roots by 40 and 58% respectively under the co-application of P. vermicola, ZnO NPs and OA in comparison with control. Moreover, the better activity of soil phosphatase and invertase was assessed under the effect of our application. These results cast a new light on the application of P. vermicola, ZnO NPs and OA in both separate and combined form as a feasible and ecofriendly tool to alleviate As stress in L. acutangula.

Application of nanopesticides may substantially increase surface attachment and internalization of engineered nanoparticles (ENPs) in food crops. This study investigated the role of stomata in the internalization of silver nanoparticles (Ag NPs) using abscisic acid (ABA)-responsive ecotypes (Ler and Col-7) and ABA-insensitive mutants (ost1-2 and scord7) of Arabidopsis thaliana in batch sorption experiments, in combination with microscopic visualization. Compared with those of the ABA-free control, stomatal apertures were significantly smaller for the Ler and Col-7 ecotypes (p ˂ 0.05) but remained unchanged for the ost1-2 and scord7 mutants, after exposure to 10 μM ABA for 1 h. Generally Ag NP sorption to the leaves of the Ler and Col-7 ecotypes treated with 10 μM ABA was lower than that in the ABA-free control, mainly due to ABA-induced stomatal closure. The difference in Ag NP sorption with and without ABA was less pronounced for Col-7 than for Ler, suggesting different sorption behaviors between these two ecotypes. In contrast, there was no significant difference in foliar sorption of Ag NPs by the ost1-2 and scord7 mutants with and without ABA treatment. Ag NPs were widely attached to the Arabidopsis leaf surface, and found at cell membrane, cytoplasm, and plasmodesmata, as revealed by scanning electron microscopy and transmission electron microscopy, respectively. These results highlight the important role of stomata in the internationalization of ENPs in plants and may have broad implications in foliar application of nanopesticides and minimizing contamination of food crops by ENPs.

Heavy metal accumulation in agricultural land causes crop production losses worldwide. Metal homeostasis within cells is tightly regulated. However, homeostasis breakdown leads to accumulation of reactive oxygen species (ROS). Overall plant fitness under stressful environment is determined by coordination between roots and shoots. But little is known about organ specific responses to heavy metals, whether it depends on the metal category (redox or non-redox reactive) and if these responses are associated with heavy metal accumulation in each organ or there are driven by other signals. Maize seedlings were subjected to sub-lethal concentrations of four metals (Zn, Ni, Cd and Cu) individually, and were quantified for growth, ABA level, and redox alterations in roots, mature leaves (L1,2) and young leaves (L3,4) at 14 and 21 days after sowing (DAS). The treatments caused significant increase in endogenous metal levels in all organs but to different degrees, where roots showed the highest levels. Biomass was significantly reduced under heavy metal stress. Although old leaves accumulated less heavy metal content than root, the reduction in their biomass (FW) was more pronounced. Metal exposure triggered ABA accumulation and stomatal closure mainly in older leaves, which consequently reduced photosynthesis. Heavy metals induced oxidative stress in the maize organs, but to different degrees. Tocopherols, polyphenols and flavonoids increased specifically in the shoot under Zn, Ni and Cu, while under Cd treatment they played a minor role. Under Cu and Cd stress, superoxide dismutase (SOD) and dehydroascorbate reductase (DHAR) activities were induced in the roots, however ascorbate peroxidase (APX) activity was only increased in the older leaves. Overall, it can be concluded that root and shoot organs specific responses to heavy metal toxicity are not only associated with heavy metal accumulation and they are specialized at the level of antioxidants to cope with.

Tropospheric ozone (O₃) impairs physiological processes of plants while nitrogen (N) deposition may cause imbalances in soil N and other nutrients such as phosphorus (P) suggesting an increase of P demand for plants. However, the combined effect of O₃, soil N and P on isoprene emission from leaves has never been tested. We therefore examined isoprene emission in leaves of Oxford poplar clone exposed to O₃ (ambient, AA [35.0 nmol mol⁻¹ as daily mean]; 1.5 × AA; 2.0 × AA), soil N (0 and 80 kg N ha⁻¹) and soil P (0, 40 and 80 kg P ha⁻¹) in July and September in a Free-Air Controlled Exposure (FACE) facility. We also investigated the response of isoprene emission to foliar N, P and abscisic acid (ABA) contents in September because the 2-C-methylerythritol-5-phosphate (MEP) pathway of isoprenoid biosynthesis produces ABA. We found that O₃ increased isoprene emission in July, which was associated to increased dark respiration, suggesting an activation of metabolism against O₃ stress as an initial response. However, O₃ decreased isoprene emission in September which was associated to reduced net photosynthesis. In September, isoprene emission was positively correlated with leaf N content and negatively correlated with leaf P content in AA. However, no response of isoprene emission to foliar N and P was found in elevated O₃, suggesting that the isoprene responses to foliar N and P depended on the O₃ exposure levels. Isoprene emission rate in 1.5 × AA and 2.0 × AA increased with increasing leaf ABA content, indicating accelerated senescence of injured leaves to favor new leaf growth when high O₃ and nutritional availability in the soil were combined. Even though foliar N and P usually act as a proxy for isoprene emission rate, the impact of recent abiotic factors such as O₃ should be always considered for modeling isoprene emission under climate change.

The present study investigated the beneficial role of selenium-nanoparticles (Se-NPs) in mitigating the adverse effects of soil-salinity on growth and yield of strawberry (Fragaria × ananassa Duch.) plants by maneuvering physiological and biochemical mechanisms. The foliar spray of Se-NPs (10 and 20 mg L⁻¹) improved the growth and yield parameters of strawberry plants grown on non-saline and different saline soils (0, 25, 50 and 75 mM NaCl), which was attributed to their ability to protect photosynthetic pigments. Se-NPs-treated strawberry plants exhibited higher levels of key osmolytes, including total soluble carbohydrates and free proline, compared with untreated plants under saline conditions. Foliar application of Se-NPs improved salinity tolerance in strawberry by reducing stress-induced lipid peroxidation and H₂O₂ content through enhancing activities of antioxidant enzymes like superoxide dismutase and peroxidase. Additionally, Se-NPs-treated strawberry plants showed accumulation of indole-3-acetic acid and abscisic acid, the vital stress signaling molecules, which are involved in regulating different morphological, physiological and molecular responses of plants to salinity. Moreover, the enhanced levels of organic acids (e.g., malic, citric and succinic acids) and sugars (e.g., glucose, fructose and sucrose) in the fruits of Se-NPs-treated strawberry plants under saline conditions indicated the positive impacts of Se-NPs on the improvement of fruit quality and nutritional values. Our results collectively demonstrate the definite roles of Se-NPs in management of soil salinity-induced adverse effects on not only strawberry plants but also other crops.

Improper application of copper-based fungicides has made copper stress critical in viticulture, necessitating the need to identify substances that can mitigate it. In this study, leaves of ‘Shine Muscat’ (‘SM’) grapevine seedlings were treated with CuSO₄ solution (10 mM/L), CuSO₄ + 5-aminolevulinic acid (ALA) (50 mg/L), and distilled water to explore the mitigation effect of ALA. Physiological assays demonstrated that ALA effectively reduced malondialdehyde accumulation and increased peroxidase and superoxide dismutase activities in grapevine leaves under copper stress. Copper ion absorption, transport pathways, chlorophyll metabolism pathways, photosynthetic system, and antioxidant pathways play key roles in ALA alleviated-copper stress. Moreover, expression changes in genes, such as CHLH, ALAD, RCA, and DHAR, play vital roles in these processes. Furthermore, abscisic acid reduction caused by NCED down-regulation and decreased naringenin, leucopelargonidin, and betaine contents confirmed the alleviating effect of ALA. Taken together, these results reveal how grapevine responds to copper stress and the alleviating effects of ALA, thus providing a novel means of alleviating copper stress in viticulture.

Due to the potential toxicity of polycyclic aromatic hydrocarbons (PAHs) to humans, the uptake and translocation of PAHs in food crops have gained much attention. However, it is still unclear whether phloem participates in the acropetal translocation of PAHs in plants. Herein, the evidence for acropetal translocation of phenanthrene (a model PAH) via phloem is firstly tested. Wheat (Triticum aestivum L.) new leaves contain significantly higher phenanthrene concentration than old leaves (P < 0.05), and the inhibitory effect on phenanthrene translocation is stronger in old leaves after abscisic acid and polyvinyl alcohol (two common transpiration inhibitors) application. Phenanthrene concentration in xylem sap is slightly higher than in phloem sap. Ring-girdling treatment can significantly reduce phenanthrene concentration in castor bean (Ricinus communis L.) leaves. Two-photon fluorescence microscope images indicate a xylem-to-phloem and acropetal phloem translocation of phenanthrene in castor bean stem. Therefore, phloem is involved in the acropetal translocation of phenanthrene in wheat seedlings, especially when the xylem is not mature enough in scattered vascular bundle plants. Our results provide a deeper understanding of PAH translocation in plants, which have significant implications for food safety and phytoremediation enhancement of PAH-contaminated soil and water.

Treated wastewater is increasingly used to meet agriculture's water needs; however, treated wastewater contains numerous contaminants of emerging concern (CECs). With exposure and uptake of CECs, phytotoxicity and health of crop plants is of concern, but is poorly understood. This study evaluated the effect of low-dose, chronic exposure to a mixture of 10 CECs, including 4 antibiotics, 3 anti-inflammatory drugs, 1 antiepileptic, 1 beta-blocker, and 1 antimicrobial, on lettuce (Lactuca sativa) and cucumber (Cucumis sativa L.) plants. The CEC mixture was added in nutrient media at 1 to 20X of their typical levels in treated wastewater effluents. Biological endpoints including germination, growth, phytohormone homeostasis, and CEC bioaccumulation were determined. Exposure to the CEC mixture did not affect the germination rate of lettuce seeds, but stimulated root elongation and increased the root-to-shoot biomass ratio during a 7 d cultivation. A dose-dependent decrease in biomass was observed in cucumber seedling after a 30 d exposure, with the highest rate CEC treatment resulting in decreases of 51.2 ± 20.9, 26.3 ± 34.1, and 33.2 ± 41.7% in the below-ground, above-ground, and total biomass, respectively. Levels of abscisic acid were significantly elevated (p < 0.05) in the leaves, but decreased (p < 0.05) in the roots. The dose-response of auxin was characterized by a hormesis effect. A significant 6-fold increase in the stem auxin level was observed at the 1X CEC rate, followed by a decrease to 2-fold the control at the 20X rate. Leaf auxin concentrations also significantly increased at the 1X CEC rate to 16-fold, followed by a decrease at the highest CEC rate. The results of this study suggeste that chronic exposure to low levels of CEC mixtures may compromise the fitness of plants, and the impairments are underlined by alterations in hormone balances.

Plant growth regulators (PGRs) assisted phytoextraction was investigated as a viable phytoremediation technology to increase the phytoextraction efficiency in contaminated soils. This study aimed to evaluate the cadimum (Cd)/lead (Pb)/zinc (Zn) phytoextraction efficiency by a hyperaccumulator Sedum alfredii Hance (S. alfredii) treated with 9 PGRs, including indole-3-acetic acid (IAA), gibberellin (GA₃), cytokinin (CKs), abscisic acid (ABA), ethylene (ETH), brassinosteroid (BR), salicylic acid (SA), strigolactones (SL) and jasmonic acid (JA), in slightly or heavily contaminated (SC and HC, respectively) soil. Results demonstrated that PGRs were able to improve S. alfredii biomass, the most significant increases were observed in GA₃ and SL for HC soil, while for SC soil, IAA and BR exhibited positive effects. The levels of Cd, Pb and Zn in the shoots of S. alfredii treated with ABA and SL were noticeably greater than in the CK treatment in HC soil, while the uptake of metals were increased by IAA and CKs in SC soil. Combined with the results of biomass and metal contents in shoots, we found that ABA showed the highest Cd removal efficiency and the maximum Pb and Zn removal efficiency was observed with GA₃, which was 62.99%, 269.23%, and 41.18%, respectively higher than the control in HC soil. Meanwhile, compared to control, the maximum removal efficiency of Cd by IAA treatment (52.80%), Pb by JA treatment (165.1%), and Zn by BR treatment (44.97%) in the SC soil. Overall, our results suggested that these PGRs, especially, ABA, SL, IAA, BR and GA₃ had great potential in improving phytoremediation efficiency of S. alfredii grown in contaminated soils.

Silicon is absorbed as uncharged mono-silicic acid by plant roots through passive absorption of Lsi1, an influx transporter belonging to the aquaporin protein family. Lsi2 then actively effluxes silicon from root cells towards the xylem from where it is exported by Lsi6 for silicon distribution and accumulation to other parts. Recently, it was proposed that silicon nanoparticles (SiNPs) might share a similar route for their uptake and transport. SiNPs then initiate a cascade of morphophysiological adjustments that improve the plant physiology through regulating the expression of many photosynthetic genes and proteins along with photosystem I (PSI) and PSII assemblies. Subsequent improvement in photosynthetic performance and stomatal behaviour correspond to higher growth, development, and productivity. On many occasions, SiNPs have demonstrated a protective role during stressful environments by improving plant-water status, source-sink potential, reactive oxygen species (ROS) metabolism, and enzymatic profile. The present review comprehensively discusses the crop improvement potential of SiNPs stretching their role during optimal and abiotic stress conditions including salinity, drought, temperature, heavy metals, and ultraviolet (UV) radiation. Moreover, in the later section of this review, we offered the understanding that most of these upgrades can be explained by SiNPs intricate correspondence with phytohormones, antioxidants, and signalling molecules. SiNPs can modulate the endogenous phytohormones level such as abscisic acid (ABA), auxins (IAAs), cytokinins (CKs), ethylene (ET), gibberellins (GAs), and jasmonic acid (JA). Altered phytohormones level affects plant growth, development, and productivity at various organ and tissue levels. Similarly, SiNPs regulate the activities of catalase (CAT), ascorbate peroxidase (APX), superoxide dismutase (SOD), and ascorbate-glutathione (AsA-GSH) cycle leading to an upgraded defence system. At the cellular and subcellular levels, SiNPs crosstalk with various signalling molecules such as Ca²⁺, K⁺, Na⁺, nitric oxide (NO), ROS, soluble sugars, and transcription factors (TFs) was also explained.