Phytogenic nanoparticles: synthesis, characterization, and their roles in physiology and biochemistry of plants.

Researchers are swarming to nanotechnology because of its potentially game-changing applications in medicine, pharmaceuticals, and agriculture. This fast-growing, cutting-edge technology is trying different approaches for synthesizing nanoparticles of specific sizes and shapes. Nanoparticles (NPs) have been successfully synthesized using physical and chemical processes; there is an urgent demand to establish environmentally acceptable and sustainable ways for their synthesis. The green approach of nanoparticle synthesis has emerged as a simple, economical, sustainable, and eco-friendly method. In particular, phytoassisted plant extract synthesis is easy, reliable, and expeditious. Diverse phytochemicals present in the extract of various plant organs such as root, leaf, and flower are used as a source of reducing as well as stabilizing agents during production. Green synthesis is based on principles like prevention/minimization of waste, reduction of derivatives/pollution, and the use of safer (or non-toxic) solvent/auxiliaries as well as renewable feedstock. Being free of harsh operating conditions (high temperature and pressure), hazardous chemicals and the addition of external stabilizing or capping agents makes the nanoparticles produced using green synthesis methods particularly desirable. Different metallic nanomaterials are produced using phytoassisted synthesis methods, such as silver, zinc, gold, copper, titanium, magnesium, and silicon. Due to significant differences in physical and chemical properties between nanoparticles and their micro/macro counterparts, their characterization becomes essential. Various microscopic and spectroscopic techniques have been employed for conformational details of nanoparticles, like shape, size, dispersity, homogeneity, surface structure, and inter-particle interactions. UV-visible spectroscopy is used to examine the optical properties of NPs in solution. XRD analysis confirms the purity and phase of NPs and provides information about crystal size and symmetry. AFM, SEM, and TEM are employed for analyzing the morphological structure and particle size of NPs. The nature and kind of functional groups or bioactive compounds that might account for the reduction and stabilization of NPs are detected by FTIR analysis. The elemental composition of synthesized NPs is determined using EDS analysis. Nanoparticles synthesized by green methods have broad applications and serve as antibacterial and antifungal agents. Various metal and metal oxide NPs such as Silver (Ag), copper (Cu), gold (Au), silicon dioxide (SiO2), zinc oxide (ZnO), titanium dioxide (TiO2), copper oxide (CuO), etc. have been proven to have a positive effect on plant growth and development. They play a potentially important role in the germination of seeds, plant growth, flowering, photosynthesis, and plant yield. The present review highlights the pathways of phytosynthesis of nanoparticles, various techniques used for their characterization, and their possible roles in the physiology of plants.

Microplastics and nanoplastics: Source, behavior, remediation, and multi-level environmental impact.

Plastics introduced into the natural environment persist, degrade, and fragment into smaller particles due to various environmental factors. Microplastics (MPs) (ranging from 1 µm to 5 mm) and nanoplastics (NPs) (less than 1 µm) have emerged as pollutants posing a significant threat to all life forms on Earth. Easily ingested by living organisms, they lead to ongoing bioaccumulation and biomagnification. This review summarizes existing studies on the sources of MPs and NPs in various environments, highlighting their widespread presence in air, water, and soil. It primarily focuses on the sources, fate, degradation, fragmentation, transport, and ecotoxicity of MPs and NPs. The aim is to elucidate their harmful effects on marine organisms, soil biota, plants, mammals, and humans, thereby enhancing the understanding of the complex impacts of plastic particles on the environment. Additionally, this review highlights remediation technologies and global legislative and institutional measures for managing waste associated with MPs and NPs. It also shows that effectively combating plastic pollution requires the synergization of diverse management, monitoring strategies, and regulatory measures into a comprehensive policy framework.

Potential role of tocopherol in protecting crop plants against abiotic stresses.

The changing global climate have given rise to abiotic stresses that adversely affect the metabolic activities of plants, limit their growth, and agricultural output posing a serious threat to food production. The abiotic stresses commonly lead to production of reactive oxygen species (ROS) that results in cellular oxidation. Over the course of evolution, plants have devised efficient enzymatic and non-enzymatic anti-oxidative strategies to counteract harmful effects of ROS. Among the emerging non-enzymatic anti-oxidative technologies, the chloroplast lipophilic antioxidant vitamin A (Tocopherol) shows great promise. Working in coordination with the other cellular antioxidant machinery, it scavenges ROS, prevents lipid peroxidation, regulates stable cellular redox conditions, simulates signal cascades, improves membrane stability, confers photoprotection and enhances resistance against abiotic stresses. The amount of tocopherol production varies based on the severity of stress and its proposed mechanism of action involves arresting lipid peroxidation while quenching singlet oxygen species and lipid peroxyl radicals. Additionally, studies have demonstrated its coordination with other cellular antioxidants and phytohormones. Despite its significance, the precise mechanism of tocopherol action and signaling coordination are not yet fully understood. To bridge this knowledge gap, the present review aims to explore and understand the biosynthesis and antioxidant functions of Vitamin E, along with its signal transduction and stress regulation capacities and responses. Furthermore, the review delves into the light harvesting and photoprotection capabilities of tocopherol. By providing insights into these domains, this review offers new opportunities and avenues for using tocopherol in the management of abiotic stresses in agriculture.

Phytoecdysteroids: Distribution, Structural Diversity, Biosynthesis, Activity, and Crosstalk with Phytohormones.

Phytoecdysteroids (PEs) are naturally occurring polyhydroxylated compounds with a structure similar to that of insect molting hormone and the plant hormone brassinosteroids. PEs have a four-ringed skeleton composed of 27, 28, 29, or 30 carbon atoms (derived from plant sterols). The carbon skeleton of ecdysteroid is known as cyclopentanoperhydrophenanthrene and has a ß-sidechain on C-17. Plants produce PEs via the mevalonate pathway with the help of the precursor acetyl-CoA. PEs are found in algae, fungi, ferns, gymnosperms, and angiosperms; more than 500 different PEs are found in over 100 terrestrial plants. 20-hydroxyecdysone is the most common PE. PEs exhibit versatile biological roles in plants, invertebrates, and mammals. These compounds contribute to mitigating biotic and abiotic stresses. In plants, PEs play a potent role in enhancing tolerance against insects and nematodes via their allelochemical activity, which increases plant biological and metabolic responses. PEs promote enzymatic and non-enzymatic antioxidant defense systems, which decrease reactive oxygen species in the form of superoxide radicals and hydroxyl radicals and reduce malondialdehyde content. PEs also induce protein biosynthesis and modulate carbohydrate and lipid synthesis. In humans, PEs display biological, pharmacological, and medicinal properties, such as anti-diabetic, antioxidant, anti-microbial, hepatoprotective, hypoglycemic, anti-cancer, anti-inflammatory, antidepressant, and tissue differentiation activity.

ß-Cyclocitral: Emerging Bioactive Compound in Plants.

ß-cyclocitral (ßCC), a main apocarotenoid of ß-carotene, increases plants' resistance against stresses. It has recently appeared as a novel bioactive composite in a variety of organisms from plants to animals. In plants, ßCC marked as stress signals that accrue under adverse ecological conditions. ßCC regulates nuclear gene expression through several signaling pathways, leading to stress tolerance. In this review, an attempt has been made to summarize the recent findings of the potential role of ßCC. We emphasize the ßCC biosynthesis, signaling, and involvement in the regulation of abiotic stresses. From this review, it is clear that discussing compound has great potential against abiotic stress tolerance and be used as photosynthetic rate enhancer. In conclusion, this review establishes a significant reference base for future research.

Copper: uptake, toxicity and tolerance in plants and management of Cu-contaminated soil.

Copper (Cu) is an essential mineral nutrient for the proper growth and development of plants; it is involved in myriad morphological, physiological, and biochemical processes. Copper acts as a cofactor in various enzymes and performs essential roles in photosynthesis, respiration and the electron transport chain, and is a structural component of defense genes. Excess Cu, however, imparts negative effects on plant growth and productivity. Many studies have summarized the adverse effects of excess Cu on germination, growth, photosynthesis, and antioxidant response in agricultural crops. Its inhibitory influence on mineral nutrition, chlorophyll biosynthesis, and antioxidant enzyme activity has been verified. The current review focuses on the availability and uptake of Cu by plants. The toxic effects of excess Cu on seed germination, plant growth and development, photosynthesis, and antioxidant response in plants are discussed. Plant tolerance mechanisms against Cu stress, and management of Cu-contaminated soils are presented.

Nanoparticles enhances the salinity toxicity tolerance in Linum usitatissimum L. by modulating the antioxidative enzymes, photosynthetic efficiency, redox status and cellular damage.

The contribution of nanoparticles (NPs) in physiology of the plants became the new area of interest for the physiologists; as it is very much cost effective compared to the phytohormones. Our present investigation was also based on this interest in which the same doses (50 mg/L) of four different NPs were sprayed on stressed and non-stressed foliage. The experiment was conducted to assess the impact of four NPs viz., zinc oxide (ZnO), silicon dioxide (SiO2), titanium dioxide (TiO2), and ferric oxide (Fe2O3) on the morphology and physiology of linseed in the presence of sodium chloride (NaCl). Plants responded positively to all the treated NPs and improved the growth, carbon and nutrient assimilation, while salt stress increased the content of proline, hydrogen peroxide and superoxide anion. Application of NPs over the stressed plants further increased the antioxidant enzymatic system and other physiochemical reactions. Results indicate that application of NPs increased the growth and physiology of the plant and also increased the salt tolerance capacity of the plant.

Phytoremediation of Cadmium Contaminated Soil Using Brassica juncea: Influence on PSII Activity, Leaf Gaseous Exchange, Carbohydrate Metabolism, Redox and Elemental Status.

Phytoremediation is an ecologically and economically feasible technique to remove heavy metal from soil. The aim of the study was to examine cadmium (Cd) toxicity and phytoremediation aptitude of Brassica juncea. In the present study, plants survived when exposed to different levels of Cd (0, 25, 50 and 100 mg/kg soil) and accumulated a large amount of Cd in its root and shoot. Translocation factor (TF) of Cd from root to shoot was > 1 at both 45 and 60-day stage of growth suggesting that B. juncea is a hyperaccumulator and strong candidate for phytoextraction of Cd. Alongside, Cd impaired photolysis of water, PSII activity, nutrient uptake, photosynthesis and sugar accumulation in the plant. Cd-generated oxidative stress restricts the growth of B. juncea. The toxic effect of Cd was more pronounced at 45-day stage of growth signifying the drifting of plant towards acquirement of exclusion strategy.

Foliar spray of Auxin/IAA modulates photosynthesis, elemental composition, ROS localization and antioxidant machinery to promote growth of Brassica juncea.

Auxins (Aux) are primary growth regulators that regulate almost every aspect of growth and development in plants. It plays a vital role in various plant processes besides controlling the key aspects of cell division, cell expansion, and cell differentiation. Considering the significance of Aux, and its potential applications, a study was conducted to observe the impact of indole acetic acid (IAA), a most active and abundant form of Aux on Brassica juncea plants growing under natural environmental conditions. Different concentrations (0, 10-10, 10-8, 10-6 M) of IAA were applied once in a day at 25-day stage of growth for 5 days, consecutively. Various parameters (growth, photosynthetic, biochemical, oxidative biomarkers and nutrient composition) were assessed at different days after sowing (DAS). Scanning electron microscopy (SEM) of leaf stomata, reactive oxygen species (ROS) localization in leaf and roots, and confocal microscopy were also conducted. The results revealed that all the IAA concentrations were effective in growth promotion and ROS reduction, however, the 10-8 M of IAA exhibited the maximum improvement in all the above mentioned parameters as compared to the control.

Nitric oxide-mediated integrative alterations in plant metabolism to confer abiotic stress tolerance, NO crosstalk with phytohormones and NO-mediated post translational modifications in modulating diverse plant stress.

Nitric oxide (NO) is a major signaling biomolecule associated with signal transduction in plants. The beneficial role of NO in plants, exposed to several abiotic stresses shifted our understanding as it being not only free radical, released from the toxic byproducts of oxidative metabolism but also helps in plant sustenance. An explosion of research in plant NO biology during the last two decades has revealed that NO is a key signal associated with plant growth, germination, photosynthesis, leaf senescence, pollen growth and reorientation. NO is beneficial as well as harmful to plants in a dose-dependent manner. Exogenous application of NO at lower concentrations promotes seed germination, hypocotyl elongation, pollen development, flowering and delays senescence but at higher concentrations it causes nitrosative damage to plants. However, this review concentrates on the beneficial impact of NO in lower concentrations in the plants and also highlights the NO crosstalk of NO with other plant hormones, such as auxins, gibberellins, abscisic acid, cytokinins, ethylene, salicylic acid and jasmonic acid, under diverse stresses. While concentrating on the multidimensional role of NO, an attempt has been made to cover the role of NO-mediated genes associated with plant developmental processes, metal uptake, and plant defense responses as well as stress-related genes. More recently, several NO-mediated post translational modifications, such as S-nitrosylation, N-end rule pathway operates under hypoxia and tyrosine nitration also occurs to modulate plant physiology.

Regulation of growth and photosynthetic parameters by salicylic acid and calcium in Brassica juncea under cadmium stress.

Cadmium, a non-essential and toxic metal, negatively affects plant growth and productivity, and alters the plant's physiological processes necessary for its survival. The present study was designed to explore the individual and combined effects of calcium and salicylic acid (SA) on the morphology and physiology of Brassica juncea L. cv. Varuna under cadmium stress. The application of calcium (2 mM) through the soil and/or SA (10-5 M) as foliar spray enhanced the growth, photosynthetic parameters, and proline content determined after 45 days of treatment. The application of cadmium (6 mg kg-1) through the soil was toxic and decreased both growth and the photosynthetic parameters. The application of calcium and SA in combination was most effective in alleviating the harmful effects of cadmium on growth and photosynthesis. Calcium and SA clearly induced plant protection mechanisms by enhancing proline and chlorophyll accumulation in the leaves.

Arsenic-induced plant stress: Mitigation strategies and omics approaches to alleviate toxicity.

Arsenic (As) is a metalloid pollutant that is extensively distributed in the biosphere. As is among the most prevalent and toxic elements in the environment; it induces adverse effects even at low concentrations. Due to its toxic nature and bioavailability, the presence of As in soil and water has prompted numerous agricultural, environmental, and health concerns. As accumulation is detrimental to plant growth, development, and productivity. Toxicity of As to plants is a function of As speciation, plant species, and soil properties. As inhibits root proliferation and reduces leaf number. It is associated with defoliation, reduced biomass, nutrient uptake, and photosynthesis, chlorophyll degradation, generation of reactive oxygen species, membrane damage, electrolyte leakage, lipid peroxidation and genotoxicity. Plants respond to As stress by upregulating genes involved in detoxification. Different species have adopted avoidance and tolerance responses for As detoxification. Plants also activate phytohormonal signaling to mitigate the stressful impacts of As. This review addresses As speciation, uptake, and accumulation by plants. It describes plant morpho-physiological, biochemical, and molecular changes and how phytohormones respond to As stress. The review closes with a discussion of omic approaches for alleviating As toxicity in plants.

Quercetin-mediated alteration in photosynthetic efficiency, sugar metabolism, elemental status, yield, and redox potential in two varieties of okra.

Quercetin is a bioactive natural compound with an antioxidative property that can potentially modify plant physiology. The current investigation aimed to gauge the effect of different concentrations of foliar spray of quercetin (0, 0.5, 1, 1.5, 2.0 mM) on several morphological and physio-biochemical performances of Abelmoschus esculentus L. (Moench.) plants under normal environmental conditions. The foliar spray on the plant leaves was applied 25 days after sowing (DAS) and continued up to 30 DAS once each day. The plants were sampled at 30 and 45 DAS to monitor several parameters. The foliar treatments of quercetin significantly upgraded all the studied parameters. The results direct that most of the traits such as growth, nutrient uptake, photosynthetic, and enzyme activities were promoted in a dose-dependent way. Quercetin application lowered the reactive oxygen species (ROS) buildup by increasing the antioxidant enzyme activities. Microscopic investigations further revealed a significant enhancement in the stomatal aperture under quercetin application. Out of several doses tested, 1 mM of quercetin proved best and can be used for further investigations.

Insights into salicylic acid-mediated redox homeostasis, carbohydrate metabolism and secondary metabolite involvement in improvement of photosynthetic performance, enzyme activities, ionomics, and yield in different varieties of Abelmoschus esculentus.

Salicylic acid (SA) is a well-known signaling molecule and phenolic plant hormone. However, the optimal concentration of SA required for beneficial effects may vary across different plant species. The objective of this study was to investigate the effects of salicylic acid (SA) on two different varieties of Abelmoschus esculentus (Sakata-713 and Neelam) in order to determine the optimal concentration of SA and its impact on the growth, physiology, and biochemical processes of the plants. We conducted an experiment applying different SA concentrations (0, 10-4, 10-5, 10-6, 10-7 M) at 25 days after sowing (DAS) and evaluated various plant parameters at different stages. To evaluate various parameters sampling was performed at 30 and 45 DAS; yield traits were calculated at 60 DAS. The results indicate that SA application increased cell division, trichome number, chlorophyll content, photosynthesis, gas exchange traits, and elemental status which further boosted plants growth and yield traits. SA application stimulated activity of several enzymes that participate in carboxylation/decarboxylation homeostasis (carbonic anhydrase), nitrogen metabolism (nitrate reductase), Calvin cycle (Rubisco), TCA cycle (succinate dehydrogenase and fumarase) and secondary metabolism (phenylalanine lyase). A gradual increase in the production of secondary metabolites (total phenol, total flavonoid, anthocyanin) and carbon metabolism (total reducing sugars, starch, glucose, fructose, sucrose) was observed. Notably, SA treatment also played a vital role in maintaining a balanced equilibrium between reactive oxygen species (ROS) and the scavenging system (catalase, peroxidase, superoxide dismutase). Based on our results, the optimal concentration of SA was determined to be 10-5 M, as it yielded the most favourable outcomes among the different concentrations tested. Moreover, when comparing the two varieties of okra, Sakata-713 exhibited a more promising response to SA treatment compared to Neelam.

CuO Nanoparticle-Mediated Seed Priming Improves Physio-Biochemical and Enzymatic Activities of Brassica juncea.

The use of nanoparticles (NPs) in agricultural fields has risen to a level where people are considering NPs as an alternative to commercial fertilizers. The input of copper oxide NPs (CuO NPs) as seed primers was investigated in this study, and the growth indices of Brassica juncea such as phenotypic parameters, photosynthetic attributes, and biochemical parameters were measured during maximum vegetative growth stage, i.e., at 45 days after sowing. Surface sterilized seeds were soaked in varying concentrations (0, 2, 4, 8 and 16 mg/L) of CuO NPs for 15, 30, and/or 45 min. After those priming periods, the seeds were planted in pots and allowed to grow naturally. Among the different tested concentrations of CuO NPs, 4 mg/L of CuO NPs for 30 min seed priming proved to be best, and considerably increased the, shoot length (30%), root length (27%), net photosynthetic rate (30%), internal CO2 concentration (28%), and proline content (41%). Besides, the performance of the antioxidant enzymes, viz, superoxide dismutase, catalase, peroxidase, and biochemical parameters such as nitrate reductase and carbonic anhydrase were also increased by several folds after the application of CuO NPs in B. juncea. The present study suggests that CuO NPs can be effectively used to increase the performance of B. juncea and may also be suitable for testing on other crop species.

Nanobionics: A Sustainable Agricultural Approach towards Understanding Plant Response to Heavy Metals, Drought, and Salt Stress.

In the current scenario, the rising concentration of heavy metals (HMs) due to anthropogenic activities is a severe problem. Plants are very much affected by HM pollution as well as other abiotic stress such as salinity and drought. It is very important to fulfil the nutritional demands of an ever-growing population in these adverse environmental conditions and/or stresses. Remediation of HM in contaminated soil is executed through physical and chemical processes which are costly, time-consuming, and non-sustainable. The application of nanobionics in crop resilience with enhanced stress tolerance may be the safe and sustainable strategy to increase crop yield. Thus, this review emphasizes the impact of nanobionics on the physiological traits and growth indices of plants. Major concerns and stress tolerance associated with the use of nanobionics are also deliberated concisely. The nanobionic approach to plant physiological traits and stress tolerance would lead to an epoch of plant research at the frontier of nanotechnology and plant biology.

Auxin regulates growth, photosynthetic efficiency and mitigates copper induced toxicity via modulation of nutrient status, sugar metabolism and antioxidant potential in Brassica juncea.

The involvement of auxin (IAA) in growth and development of plants is well known, but its role in the mitigation of metal stress, especially copper (Cu), is not fully understood; therefore, it is time to explore its involvement in minimizing the stress. A pot experiment was conducted to assess the protective function of IAA, applied to the foliage, on photosynthetic machinery, carbohydrate metabolism, and growth of Brassica juncea, grown with Cu (30 or 60 mg kg-1 of soil). Among the different concentrations (10-10, 10-8, or 10-6 M), 10-8 M of IAA alone enhanced the photosynthetic characteristics, sugar accumulation and vegetative growth with minimal cellular oxidative stress level. Moreover, the same concentration of auxin was most effective in decreasing the stress levels generated by Cu and maintained it nearly to that of the control in terms of photosynthetic attributes, gas exchange parameters, PSII activity, electron transport rate, and growth attributes. Auxin also maintained the membrane stability and ultrastructure of chloroplast, stomatal morphology with a reduction in malondialdehyde (MDA), electrolyte leakage (EL) and cell death in test plants even under Cu stress. IAA also improved the translocation of Cu from root to the aerial parts, thus enhanced the Cu-reclamation in metal contaminated soils. Our findings suggest that the application of 10-8 M of IAA maintains the overall growth of plants and may be used as an effective agent to improve growth, photosynthesis and phyto-remediation potential of B. juncea in Cu contaminated soil.

Perspective of Melatonin-Mediated Stress Resilience and Cu Remediation Efficiency of Brassica juncea in Cu-Contaminated Soils.

The present study evaluated the influence of melatonin (MEL) on copper toxicity in terms of morphophysiological, microscopic, histochemical, and stress resilience responses in Brassica juncea. Different levels of Cu (0, 30, and 60 mg kg-1) were given in air-dried soil, and 25 days after sowing (DAS), plants were sprayed with 30, 40, or 50 µM of MEL. The results demonstrated that under Cu stress, a significant amount of Cu accumulated in plant tissues, particularly in roots than in upper ground tissues, thereby suppressing the overall growth as evidenced by decrease in tolerance index and photosynthesis and increase in oxidative stress biomarkers (reactive oxygen species, malondialdehyde, and electrolyte leakage content) and cell death. Interestingly, the follow-up treatment of MEL, mainly 40 µM, efficiently improved the physio-biochemical and growth parameters, sugar accumulation, and metabolism. The potential of MEL in modulating Cu stress is attributed to its involvement in enriching the level of nutrient and improving chloroplast and stomatal organization besides lowering oxidative stress via enhanced levels of antioxidants. MEL improved the Cu reclamation potential in plants by enhancing Cu uptake and its translocation to aerial tissues. Principal component analysis showed that most of the morphophysiological and growth attributes were positively linked with MEL and negatively related to Cu levels, whereas all the stress-enhancing attributes showed a strong relationship with excessive Cu levels in soils. The present study suggested that MEL has the potential to improve growth and photosynthesis resulting in improved stress resilience under Cu stress along with increased remediation capability of mustard for remediation of Cu-contaminated soils.

Specific Roles of Lipoxygenases in Development and Responses to Stress in Plants.

Lipoxygenases (LOXs), naturally occurring enzymes, are widely distributed in plants and animals. LOXs can be non-sulfur iron, non-heme iron, or manganese-containing dioxygenase redox enzymes. LOXs catalyze the oxidation of polyunsaturated fatty acids into fatty acid hydroperoxides. Linolenic acid, a precursor in the jasmonic acid (JA) biosynthesis, is converted to 12-oxo-phytodienoic acid through oxygenation with LOX, allene oxide synthase, and allene oxide cyclase. Moreover, JA participates in seed germination, fruit ripening, senescence, and many other physio-biochemical processes. LOXs also play crucial roles in defense responses against biotic stress, i.e., insects, pests, pathogenic attacks, and abiotic stress, such as wounding, UV-rays, extreme temperature, oxidative stress, and drought.

Protective response of 28-homobrassinolide in cultivars of Triticum aestivum with different levels of nickel.

Seeds of five wheat (Triticum aestivum) cultivars (PBW-373, UP-2338, DL-LOK-01, DL-373, and HD-2338) were sown in earthen pots and 10-day-old seedlings were exposed to 0, 50, and 100 µM of nickel (Ni) in the form of nickel chloride. At the 20-day stage, seedlings were sprayed with 0.01 µM of 28-homobrassinolide (HBL). The results of the experiment at the 30-day stage revealed a decline in the dry mass per plant, leaf area, leaf water potential, and net photosynthetic rate with concomitant decline in the activities of various enzymes (viz. carbonic anhydrase and nitrate reductase) with an increasing concentration of Ni. However, an increase in proline content and the activities of catalase, peroxidase, and superoxide dismutase was observed as a result of an increase in Ni concentration. Moreover, the treatment of these stressed plants with HBL enhanced the activities of carbonic anhydrase and nitrate reductase, catalase, peroxidase, and superoxide dismutase. The proline content in the leaves also increased, which is known to act as an osmolyte and reactive oxygen species scavenger. The toxic effects generated by Ni were ameliorated by HBL through an improved antioxidant system and osmolyte. Moreover, improvement of photosynthetic parameters and growth characteristics further strengthen our belief that HBL acted as a potent stress alleviator.