Silicon regulates phosphate deficiency through involvement of auxin and nitric oxide in barley roots.

MAIN CONCLUSION: Silicon application mitigates phosphate deficiency in barley through an interplay with auxin and nitric oxide, enhancing growth, photosynthesis, and redox balance, highlighting the potential of silicon as a fertilizer for overcoming nutritional stresses. Silicon (Si) is reported to attenuate nutritional stresses in plants, but studies on the effect of Si application to plants grown under phosphate (Pi) deficiency are still very scarce, especially in barley. Therefore, the present work was undertaken to investigate the potential role of Si in mitigating the adverse impacts of Pi deficiency in barley Hordeum vulgare L. (var. BH902). Further, the involvement of two key regulatory signaling molecules--auxin and nitric oxide (NO)--in Si-induced tolerance against Pi deficiency in barley was tested. Morphological attributes, photosynthetic parameters, oxidative stress markers (O2·-, H2O2, and MDA), antioxidant system (enzymatic--APX, CAT, SOD, GR, DHAR, MDHAR as well as non-enzymatic--AsA and GSH), NO content, and proline metabolism were the key traits that were assessed under different treatments. The P deficiency distinctly declined growth of barley seedlings, which was due to enhancement in oxidative stress leading to inhibition of photosynthesis. These results were also in parallel with an enhancement in antioxidant activity, particularly SOD and CAT, and endogenous proline level and its biosynthetic enzyme (P5CS). The addition of Si exhibited beneficial effects on barley plants grown in Pi-deficient medium as reflected in increased growth, photosynthetic activity, and redox balance through the regulation of antioxidant machinery particularly ascorbate-glutathione cycle. We noticed that auxin and NO were also found to be independently participating in Si-mediated improvement of growth and other parameters in barley roots under Pi deficiency. Data of gene expression analysis for PHOSPHATE TRANSPORTER1 (HvPHT1) indicate that Si helps in increasing Pi uptake as per the need of Pi-deficient barley seedlings, and also auxin and NO both appear to help Si in accomplishing this task probably by inducing lateral root formation. These results are suggestive of possible application of Si as a fertilizer to correct the negative effects of nutritional stresses in plants. Further research at genetic level to understand Si-induced mechanisms for mitigating Pi deficiency can be helpful in the development of new varieties with improved tolerance against Pi deficiency, especially for cultivation in areas with Pi-deficient soils.

Redox regulation by priming agents towards a sustainable agriculture.

Plant are sessile organisms that are often subjected to a multitude of environmental stresses, with the occurrence of these events being further intensified by global climate change. Crop species therefore require specific adaptations to tolerate climatic variability for sustainable food production. Plant stress results in excess accumulation of reactive oxygen species (ROS) leading to oxidative stress, and loss of cellular redox balance in the plant cells. Moreover, enhancement of cellular oxidation as well as oxidative signals have recently been recognized as crucial players in plant growth regulation under stress conditions. Multiple roles of redox regulation in crop production have been well documented, and major emphasis has focused on key redox-regulated proteins and non-protein molecules, such as NAD(P)H, thioredoxins, glutathione, glutaredoxins, peroxiredoxins, ascorbate, and reduced ferredoxin. These have been widely implicated in the regulation of (epi)genetic factors modulating growth and vigor of crop plants, particularly within an agricultural context. In this regard, priming with the employment of chemical and biological agents has emerged as a fascinating approach to improve plant tolerance against various abiotic and biotic stressors. Priming in plants is a physiological process, where prior exposure to specific stressors induces a state of heightened alertness, enabling a more rapid and effective defense response upon subsequent encounters with similar challenges. Priming is reported to play an important role in the regulation of cellular redox homeostasis, maximizing crop productivity under stress conditions and thus achieving yield security. By taking this into consideration, the present review is an up-to-date critical evaluation of promising plant priming technologies and their role in the regulation of redox components towards enhanced plant adaptations to extreme unfavorable environmental conditions. The challenges and opportunities of plant priming are addressed, with the aim to encourage future research in this field towards effective application in crop stress management including horticultural species.

Cytokinin and indole-3-acetic acid crosstalk is indispensable for silicon mediated chromium stress tolerance in roots of wheat seedlings.

The rising heavy metal contamination of soils imposes toxic impacts on plants as well as other life forms. One such highly toxic and carcinogenic heavy metal is hexavalent chromium [Cr(VI)] that has been reported to prominently retard the plant growth. The present study investigated the potential of silicon (Si, 10 µM) to alleviate the toxicity of Cr(VI) (25 µM) on roots of wheat (Triticum aestivum L.) seedlings. Application of Si to Cr(VI)-stressed wheat seedlings improved their overall growth parameters. This study also reveals the involvement of two phytohormones, namely auxin and cytokinin and their crosstalk in Si-mediated mitigation of the toxic impacts of Cr(VI) in wheat seedlings. The application of cytokinin alone to wheat seedlings under Cr(VI) stress reduced the intensity of toxic effects of Cr(VI). In combination with Si, cytokinin application to Cr(VI)-stressed wheat seedlings significantly minimized the decrease induced by Cr(VI) in different parameters such as root-shoot length (10.8% and 13%, respectively), root-shoot fresh mass (11.3% and 10.1%, respectively), and total chlorophyll and carotenoids content (13.4% and 6.8%, respectively) with respect to the control. This treatment also maintained the regulation of proline metabolism (proline content, and P5CS and PDH activities), ascorbate-glutathione (AsA-GSH) cycle and nutrient homeostasis. The protective effect of Si and cytokinin against Cr(VI) stress was minimized upon supplementation of an inhibitor of polar auxin transport- 2,3,5-triiodobenzoic acid (TIBA) which suggested a potential involvement of auxin in Si and cytokinin-mediated mitigation of Cr(VI) toxicity. The exogenous addition of a natural auxin - indole-3-acetic acid (IAA) confirmed auxin is an active member of a signaling cascade along with cytokinin that aids in Si-mediated Cr(VI) toxicity alleviation as IAA application reversed the negative impacts of TIBA on wheat roots treated with Cr(VI), cytokinin and Si. The results of this research are also confirmed by the gene expression analysis conducted for nutrient transporters (Lsi1, CCaMK, MHX, SULT1 and ZIP1) and enzymes involved in the AsA-GSH cycle (APX, GR, DHAR and MDHAR). The overall results of this research indicate towards possible induction of a crosstalk between cytokinin and IAA upon Si supplementation which in turn stimulates physiological, biochemical and molecular changes to exhibit protective effects against Cr(VI) stress. Further, the information obtained suggests probable employment of Si, cytokinin and IAA alone or combined in agriculture to maintain plant productivity under Cr(VI) stress and data regarding expression of key genes can be used to develop new crop varieties with enhanced resistance against Cr(VI) stress together with its reduced load in seedlings.

Encapsulated nanopesticides application in plant protection: Quo vadis?

The increased global food insecurity due to the growing population can be addressed with precision and sustainable agricultural practices. To tackle the issues regarding food insecurity, farmers used different agrochemicals that improved plant growth and protection. Among these agrochemicals, synthetic pesticides used for plant protection in the agricultural field have various disadvantages. Conventional applications of synthetic pesticides have drawbacks such as rapid degradation, poor solubility, and non-target effects, as well as increased pesticide runoff that pollutes the environment. Nanotechnology has evolved as a potential solution to increase agricultural productivity through the development of different nanoforms of agrochemicals such as nanopesticides, nano-fabricated fertilizers, nanocapsules, nanospheres, nanogels, nanofibers, nanomicelles, and nano-based growth promoters. Encapsulation of these pesticides inside the nanomaterials has provided good biocompatibility over conventional application by inhibiting the early degradation of active ingredients (AI), increasing the uptake and adhesion of pesticides, improving the stability, solubility, and permeability of the pesticides, and decreasing the environmental impacts due to the pesticide runoff. In this review, different nanoforms of encapsulated pesticides and their smart delivery systems; nanocarriers in RNA interference (RNAi) based pesticides; environmental fate, practical implications, management of nanopesticides; and future perspectives are discussed.

Zinc induced regulation of PCR1 gene for cadmium stress resistance in rice roots.

In this study, the interaction between zinc (Zn) and cadmium (Cd) was investigated in rice roots to evaluate how Zn can protect the plants from Cd stress. Rice seedlings were treated with Cd (100 µM) and Zn (100 µM) in different combinations (Cd alone, Zn alone, Zn+ Cd, Zn+ Cd+ L-NAME, Zn+ Cd+ L-NAME+ SNP). Rice roots treated with only Zn also displayed similar toxic effects, however when combined with Cd exhibited improved growth. Treating the plant with Zn along with Cd distinctly reduced Cd concentration in roots while increasing its own accumulation due to modulation in expression of Zinc-Regulated Transporter (ZRT)-/IRT-Like Protein (OsZIP1) and Plant Cadmium Resistance1 (OsPCR1). Cd reduced plant biomass, cell viability, pigments, photosynthesis and causing oxidative stress due to inhibition in ascorbate-glutathione cycle. L-NAME (NG-nitro L-arginine methyl ester), prominently suppressed the beneficial impacts of Zn against Cd stress, whereas the presence of a NO donor, sodium nitroprusside (SNP), significantly reversed this effect of L-NAME. Collectively, results point that NO signalling is essential for Zn- mediated cross-tolerance against Cd stress via by modulating uptake of Cd and Zn and expression of OsZIP1 and OsPCR1, and ROS homeostasis due to fine tuning of ascorbate-glutathione cycle which finally lessened oxidative stress in rice roots. The results of this study can be utilized to develop new varieties of rice through genetic modifications which will be of great significance for maintaining crop productivity in Cd-contaminated areas throughout the world.

Bacterial community and root endodermis: a complementary relationship.

There are feedforward and feedback loops along the microbiota-root-shoot axis to maintain plant growth or defense under environmental stresses. Here, we highlight a reciprocal interaction between the endodermis and the plant-bacterial community, which stabilizes the diffusion barriers to maintain nutrient homeostasis under nutritional stress.

Nanocarrier spray: a nontransgenic approach for crop engineering.

Genetic modification allows engineering of important traits in crops through expensive and tedious procedures to alter their genetic background. Recently, Thagun et al. developed a nanocarrier-based foliar spray method to translocate bioactive molecules of interest into plant cells to engineer important traits without introducing a transgene.

Nitric oxide overcomes copper and copper oxide nanoparticle-induced toxicity in Sorghum vulgare seedlings through regulation of ROS and proline metabolism.

This study aimed to investigate the phytotoxic effect of copper (Cu) and copper nanoparticles (CuONPs) and ameliorative potential of nitric oxide (NO) against these toxic materials in Sorghum vulgare Pers. seedlings. Data suggested that exposure of Cu and CuONPs significantly reduced growth, chlorophyll, carotenoids and protein in root and shoot, which coincided with increased Cu accumulation. However, addition of sodium nitroprusside (SNP, a donor of NO) lowered Cu and CuONPs mediated toxicity through restricting Cu accumulation and improving photosynthetic pigments and total soluble protein contents. Data further suggested that exposure of Cu and CuONPs significantly increased hydrogen peroxide (H2 O2 ), superoxide radicals (O2 •- ), and malondialdehyde (MDA) contents. Enhanced level of oxidative stress severely inhibited the enzymatic activities of glutathione reductase (GR), ascorbate peroxidase (APX), dehydroascorbate reductase (DHAR) and monodehydroascorbate reductase (MDHAR) but enhanced superoxide dismutase (SOD) and catalase (CAT) activity. However, addition of SNP positively regulated antioxidants enzymes activity, particularly the enzymes involved in the ascorbate-glutathione cycle to overcome Cu- and CuONPs-induced stress in Sorghum seedlings. Further, Cu and CuONPs enhanced accumulation of free proline through inducing Δ1 -pyrroline-5-carboxylate synthetase (P5CS) activity while lowering the proline dehydrogenase (PDH) activity. However, addition of SNP reversed these responses. Therefore, overall results revealed that SNP has enough potential of reducing the toxicity of Cu and CuONPs in Sorghum seedlings through regulation of proline metabolism and activity of enzymes of the ascorbate-glutathione cycle. These findings can be employed in developing new resistant varieties of Sorghum having enhanced tolerance against Cu or CuONP stress and improved productivity.

Plant Growth Hormones and Nanomaterial Interface: Exploring the connection from development to defense.

The global increase in nanotechnology applications has been unprecedented and has now moved into the area of agriculture and food production. Applications with promising potential in sustainable agriculture include nanobiosensors, nanofertilizers, nanopesticides, nano-mediated remediation strategies for contaminated soils and nanoscale strategies to increase crop production and protection. Given this, the impact of nanomaterials/nanoparticles (NPs) on plant species needs to be thoroughly evaluated as this represents a critical interface between the biosphere and the environment. Importantly, phytohormones represent a critical class of biomolecules to plant health and productivity; however, the impact of NPs on these molecules is poorly understood. In addition, phytohormones, and associated pathways, are widely explored in agriculture to influence several biological processes for the improvement of plant growth and productivity under natural as well as stressed conditions. However, the impact of exogenous applications of phytohormones on NP-treated plants has not been explored. The importance of hormone signaling and cross-talk with other metabolic systems makes these biomolecules ideal candidates for a thorough assessment of NP impacts on plant species. This article presents a critical evaluation of the existing yet limited literature available on NP-phytohormone interactions in plants. In addition, the developing strategy of nano-enabled precision delivery of phytohormones via nanocarriers will be explored. Finally, directions for future research and critical knowledge gaps will be identified for this important aspect of nano-enabled agriculture.

Silicon Palliates Chromium Toxicity through the Formation of Root Hairs in Rice (Oryza sativa) Mediated by GSH and IAA.

Along with the rapidly increasing environmental contamination by heavy metals, the exposure of plants to chromium has also magnified, resulting in a declined productivity. Hexavalent chromium [Cr(VI)], the most toxic form of Cr, brings about changes in plant processes at morpho-physiological and biochemical levels. However, silicon (Si) is known to mitigate the impact of abiotic stresses in plants. Here, we demonstrate Si-mediated alleviation of Cr(VI) toxicity and its effects on root hair formation in rice seedlings. Reduced glutathione (GSH) and indole-3 acetic acid (IAA, an important auxin) were assessed for their involvement in root hair formation after the application of Si to Cr(VI)-stressed plants, and our results confirmed their crucial significance in such developmental processes. The expression analysis of genes involved in GSH biosynthesis (OsGS2) and regeneration (OsGR1), and auxin biosynthesis (OsTAA1 and OsYUCCA1) and transport (OsAUX1 and OsPIN1) corroborated their positive role in Si-mediated root hair formation in Cr(VI)-stressed rice seedlings. Moreover, the results indicated that nitric oxide (NO) seems a probable but not fundamental component in Si-mediated formation of roots in rice during exposure to Cr(VI) stress. In this study, the indispensable role of GSH and IAA, redox homeostasis of GSH and IAA biosynthesis and transport are discussed with regard to Si-mediated formation of root hairs in rice under Cr(VI) stress. The results of the study suggest that Si is a protective agent against Cr(VI) stress in rice, and the findings can be used to develop Cr(VI) stress-tolerant varieties of rice with enhanced productivity.

Ethylene Renders Silver Nanoparticles Stress Tolerance in Rice Seedlings by Regulating Endogenous Nitric Oxide Accumulation.

Developments in the field of nanotechnology over the past few years have increased the prevalence of silver nanoparticles (AgNPs) in the environment, resulting in increased exposure of plants to AgNPs. Recently, various studies have reported the effect of AgNPs on plant growth at different concentrations. However, identifying the mechanisms and signaling molecules involved in plant responses against AgNPs stress is crucial to find an effective way to deal with the phytotoxic impacts of AgNPs on plant growth and development. Therefore, this study was envisaged to investigate the participation of ethylene in mediating the activation of AgNPs stress tolerance in rice (Oryza sativa L.) through a switch that regulates endogenous nitric oxide (NO) accumulation. Treatment of AgNPs alone hampered the growth of rice seedlings due to severe oxidative stress as a result of decline in sulfur assimilation, glutathione (GSH) biosynthesis and alteration in the redox status of GSH. These results are also accompanied by the higher endogenous NO level. However, addition of ethephon (a donor of ethylene) reversed the AgNP-induced effects. Though the application of silicon nanoparticles (SiNPs) alone promoted the growth of rice seedlings but, interestingly their application in combination with AgNPs enhanced the AgNP-induced toxicity in the seedlings through the same routes as exhibited in the case of AgNPs alone treatment. Interestingly, addition of ethephon reversed the negative effects of SiNPs under AgNPs stress. These results suggest that ethylene might act as a switch to regulate the level of endogenous NO, which in turn could be associated with AgNPs stress tolerance in rice. Furthermore, the results also indicated that addition of l-NG-nitro arginine methyl ester (l-NAME) (an inhibitor of endogenous NO synthesis) also reversed the toxic effects of SiNPs together with AgNPs, further suggesting that the low level of endogenous NO was associated with AgNPs stress tolerance. Overall, the results indicate that the low level of endogenous NO triggers AgNPs stress tolerance, while high level leads to AgNPs toxicity by regulating sulfur assimilation, GSH biosynthesis, redox status of GSH and oxidative stress markers. The results revealed that ethylene might act as a switch for regulating AgNPs stress in rice seedlings by controlling endogenous NO accumulation.

Auxin Crosstalk with Reactive Oxygen and Nitrogen Species in Plant Development and Abiotic Stress.

The phytohormone auxin acts as an important signaling molecule having regulatory functions during the growth and development of plants. Reactive oxygen species (ROS) are also known to perform signaling functions at low concentrations; however, over-accumulation of ROS due to various environmental stresses damages the biomolecules and cell structures and leads to cell death, and therefore, it can be said that ROS act as a double-edged sword. Nitric oxide (NO), a gaseous signaling molecule, performs a wide range of favorable roles in plants. NO displays its positive role in photomorphogenesis, root growth, leaf expansion, seed germination, stomatal closure, senescence, fruit maturation, mitochondrial activity and metabolism of iron. Studies have revealed the early existence of these crucial molecules during evolution. Moreover, auxin, ROS and NO together show their involvement in various developmental processes and abiotic stress tolerance. Redox signaling is a primary response during exposure of plants to stresses and shows a link with auxin signaling. This review provides updated information related to crosstalk between auxin, ROS and NO starting from their evolution during early Earth periods and their interaction in plant growth and developmental processes as well as in the case of abiotic stresses to plants.

Silicon nanoforms in crop improvement and stress management.

Although, silicon - the second most abundant element in the earth crust could not supersede carbon (C) in the competition of being the building block of life during evolution, yet its presence has been reported in some life forms. In case of the plants, silicon has been reported widely to promote the plant growth under normal as well as stressful situations. Nanoform of silicon is now being explored for its potential to improve plant productivity and its tolerance against various stresses. Silicon nanoparticles (SiNPs) in the form of nanofertilizers, nanoherbicides, nanopesticides, nanosensors and targeted delivery systems, find great utilization in the field of agriculture. However, the mechanisms underlying their uptake by plants need to be deciphered in detail. Silicon nanoformss are reported to enhance plant growth, majorly by improving photosynthesis rate, elevating nutrient uptake and mitigating reactive oxygen species (ROS)-induced oxidative stress. Various studies have reported their ability to provide tolerance against a range of stresses by upregulating plant defense responses. Moreover, they are proclaimed not to have any detrimental impacts on environment yet. This review includes the up-to-date information in context of the eminent role of silicon nanoforms in crop improvement and stress management, supplemented with suggestions for future research in this field.

Ca2+ sensor-mediated ROS homeostasis: defense without yield penalty.

Constitutively active immunity against pathogen infection affects the growth and yield of plants. Gao and colleagues recently demonstrated that a Ca2+ sensor, RESISTANCE OF RICE TO DISEASES1 (ROD1), regulates an immune suppression network that consolidates reactive oxygen species (ROS) metabolism and Ca2+-sensing to equilibrate both defense responses and optimal growth in plants.

Heavy metal induced regulation of plant biology: Recent insights.

The presence of different forms of heavy metals in the earth crust is very primitive and probably associated with the origin of plant life. However, since the beginning of human civilisation, heavy metal use and its contamination to all living systems on earth have significantly increased due to human anthropogenic activities. Heavy metals are nonbiodegradable, which directly or indirectly impact photosynthesis, antioxidant system, mineral nutrition status, phytohormones and amino acid-derived molecules. Due to the toxic behaviour of some heavy metals, the endogenous status of chemical messengers like phytohormones may get significantly influenced, leading to harmful impacts on plant growth, development and overall yield of the plants. It has been noticed that exogenous application of phytohormones, that is, abscisic acid, salicylic acid, auxins, brassinosteroids, cytokinins, ethylene and gibberellins can positively regulate the heavy metal toxicity in plants through the regulation of the ascorbate-glutathione cycle, nitrogen metabolism, proline metabolisms, transpiration rate, and cell division. Furthermore, it may also restrict the entry of heavy metals into the plant cells, which aids in the recovery of plant growth and productivity. Besides these, some defence molecules also assist the plant in dealing with heavy metal toxicity. Therefore, the present review aims to bridge the knowledge gap in this context and present outstanding discoveries related to plant life supportive processes during stressful conditions including phytohormones and heavy metal crosstalk along with suggestions for future research in this field.

Nanoparticles as potential hallmarks of drought stress tolerance in plants.

Plants are inevitably exposed to drought stress limiting their growth and causing yield loss, thus inciting food crises across the world. Nanoparticles (NPs) are regarded as effective and promising tools for modulation of crop yield to overcome current and future constraints in sustainable agricultural production by upgrading the plant tolerance mechanism under abiotic stress conditions, including drought. NPs exhibit alleviating effects against drought stress via induction of physiological and biochemical readjustments accompanied by modulation of gene expression involved in drought response/tolerance. NPs ameliorate drought-induced reduction in carbon assimilation via increasing the photosynthetic activity. The improved root growth, upregulation of aquaporins, modification of intracellular water metabolism, accumulation of compatible solutes and ion homeostasis are the major mechanisms used by NPs to mitigate the osmotic stress caused by water deficit. NPs reduce water loss from leaves through stomatal closure due to fostered abscisic acid (ABA) accumulation and ameliorate oxidative stress damage by reducing reactive oxygen species and activating the antioxidant defense system. This review provides an evolutionary foundation regarding drought stress in plant life and summarizes the interactions between NPs and plants under drought. The subsequent impact of NPs on plant development and productivity and recent nanobiotechnological approaches to improve drought stress resilience are presented. On the whole, this review highlights the significance of NPs in dealing with the global problem of water scarcity faced by farmers.

Arsenite: the umpire of arsenate perception and responses in plants.

Arsenite regulates the uptake and detoxification of arsenate in plants under low-phosphate conditions by governing the stability of PHOSPHATE STARVATION RESPONSE 1, as reported in a recent study by Navarro and colleagues. This finding opens new opportunities for research into developing mitigation strategies to deal with arsenic toxicity in plants.

Nanoparticles as a potential protective agent for arsenic toxicity alleviation in plants.

Aggrandized technological and industrial progression in past decades have occasioned immense depreciation in the quality of environment and ecosystem, majorly due to augmentation in the number of obnoxious pollutants incessantly being released in soil, water or air. Arsenic (As) is one such hazardous metalloid contaminating the environment which has the potential to detrimentally affect the life on earth. Even in minute quantity, As is known to cause various critical diseases in humans and toxicity in plants. Recent studies on nanoparticles (NPs) approve of their ability to qualify the criterion of becoming a potent tool for mitigating As-induced phytotoxicity. Nanoparticles are reported to promote plant growth under As-stress by stimulating various alterations at physiological, biochemical, and molecular levels. In this review, we provide an up-to-date compilation of research that has been carried out in comprehending the mechanisms utilized by nanoparticles including controlled As uptake and distribution in plants, maintenance of ROS homeostasis during stress and chelation and vacuolar sequestration of As so as to reduce the severity of toxicity induced by As, and potential areas of research in this field will also be indicated for future perspectives.

Silica nanoparticles: the rising star in plant disease protection.

When applied exogenously, silica (Si) can have a beneficial impact on plants under biotic stress conditions, as revealed by its recent application in the form of nanoparticles (SiO2NPs) to induce pathogen resistance (El-Shetehy et al.). This opens up a new window of research into combating the devastating effects of biotic stresses.