Heavy metals are natural non-biodegradable constituents of the Earth's crust that accumulate and persist indefinitely in the ecosystem as a result of human activities. Since the industrial revolution, the concentration of cadmium, arsenic, lead, mercury and zinc, amongst others, have increasingly contaminated soil and water resources, leading to significant yield losses in plants. These issues have become an important concern of scientific interest. Understanding the molecular and physiological responses of plants to heavy metal stress is critical in order to maximize their productivity. Recent research has extended our view of how plant hormones can regulate and integrate growth responses to various environmental cues in order to sustain life. In the present review we discuss current knowledge about the role of the plant growth hormones abscisic acid, auxin, brassinosteroid and ethylene in signaling pathways, defense mechanisms and alleviation of heavy metal toxicity.

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Interactions between plant hormones and heavy metals responses.

The sleep disorder narcolepsy is linked to the HLA-DQB1*0602 haplotype and dysregulation of the hypocretin ligand-hypocretin receptor pathway. Narcolepsy was associated with Pandemrix vaccination (an adjuvanted, influenza pandemic vaccine) and also with infection by influenza virus during the 2009 A(H1N1) influenza pandemic. In contrast, very few cases were reported after Focetria vaccination (a differently manufactured adjuvanted influenza pandemic vaccine). We hypothesized that differences between these vaccines (which are derived from inactivated influenza viral proteins) explain the association of narcolepsy with Pandemrix-vaccinated subjects. A mimic peptide was identified from a surface-exposed region of influenza nucleoprotein A that shared protein residues in common with a fragment of the first extracellular domain of hypocretin receptor 2. A significant proportion of sera from HLA-DQB1*0602 haplotype-positive narcoleptic Finnish patients with a history of Pandemrix vaccination (vaccine-associated narcolepsy) contained antibodies to hypocretin receptor 2 compared to sera from nonnarcoleptic individuals with either 2009 A(H1N1) pandemic influenza infection or history of Focetria vaccination. Antibodies from vaccine-associated narcolepsy sera cross-reacted with both influenza nucleoprotein and hypocretin receptor 2, which was demonstrated by competitive binding using 21-mer peptide (containing the identified nucleoprotein mimic) and 55-mer recombinant peptide (first extracellular domain of hypocretin receptor 2) on cell lines expressing human hypocretin receptor 2. Mass spectrometry indicated that relative to Pandemrix, Focetria contained 72.7% less influenza nucleoprotein. In accord, no durable antibody responses to nucleoprotein were detected in sera from Focetria-vaccinated nonnarcoleptic subjects. Thus, differences in vaccine nucleoprotein content and respective immune response may explain the narcolepsy association with Pandemrix.

Antibodies to influenza nucleoprotein cross-react with human hypocretin receptor 2

Plants confront multifarious environmental stresses widely divided into abiotic and biotic stresses, of which heavy metal stress represents one of the most damaging abiotic stresses Heavy metals cause toxicity by targeting crucial molecules and vital processes in the plant cell One of the approaches by which heavy metals act in plants is by over production of reactive oxygen species (ROS) either directly or indirectly Plants act against such overdose of metal in the environment by boosting the defense responses like metal chelation, sequestration into vacuole, regulation of metal intake by transporters, and intensification of antioxidative mechanisms This response shown by plants is the result of intricate signaling networks functioning in the cell in order to transmit the extracellular stimuli into an intracellular response The crucial signaling components involved are calcium signaling, hormone signaling, and mitogen activated protein kinase (MAPK) signaling that are discussed in this review Apart from signaling components other regulators like microRNAs and transcription factors also have a major contribution in regulating heavy metal stress This review demonstrates the key role of MAPKs in synchronously controlling the other signaling components and regulators in metal stress Further, attempts have been made to focus on metal transporters and chelators that are regulated by MAPK signaling

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Traversing the Links between Heavy Metal Stress and Plant Signaling.

Nitric oxide (NO) is a free radical molecule involved in an array of functions under physiological and adverse environmental conditions As other free radical molecules, NO biological action depends on its cellular concentration, acting as a signal molecule when produced at low concentration or resulting in cellular damage when produced at sufficiently high levels to trigger nitro-oxidative stress Over the last decade, significant progress has been made in characterizing NO metabolism and action mechanism, revealing that diverse biosynthetic routes can generate this free radical in plants and its action mainly occurs through posttranslational modification (nitration and S-nitrosylation) of target proteins Intricate crosstalk networks between NO and other signaling molecules have been described involving phytohormones, other second messengers, and key transcription factors This review will focus on our current understanding of NO interplay with phytohormones and other plant growth regulators under abiotic stress conditions

Nitric oxide signaling and its crosstalk with other plant growth regulators in plant responses to abiotic stress.

Roots are subjected to a range of abiotic stresses as they forage for water and nutrients. Cytosolic free calcium is a common second messenger in the signaling of abiotic stress. In addition, roots take up calcium both as a nutrient and to stimulate exocytosis in growth. For calcium to fulfill its multiple roles must require strict spatio-temporal regulation of its uptake and efflux across the plasma membrane, its buffering in the cytosol and its sequestration or release from internal stores. This prompts the question of how specificity of signaling output can be achieved against the background of calcium's other uses. Threats to agriculture such as salinity, water availability and hypoxia are signaled through calcium. Nutrient deficiency is also emerging as a stress that is signaled through cytosolic free calcium, with progress in potassium, nitrate and boron deficiency signaling now being made. Heavy metals have the capacity to trigger or modulate root calcium signaling depending on their dose and their capacity to catalyze production of hydroxyl radicals. Mechanical stress and cold stress can both trigger an increase in root cytosolic free calcium, with the possibility of membrane deformation playing a part in initiating the calcium signal. This review addresses progress in identifying the calcium transporting proteins (particularly channels such as annexins and cyclic nucleotide-gated channels) that effect stress-induced calcium increases in roots and explores links to reactive oxygen species, lipid signaling, and the unfolded protein response.

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Calcium-Mediated Abiotic Stress Signaling in Roots

The root is the first plant organ to get in contact with the toxin cadmium (Cd), which is a widespread soil contaminant. Cd inhibits the growth of the primary root, but the mechanisms underlying this inhibition remain elusive. In this study, we used physiological, pharmacological and genetic approaches to investigate the roles of nitric oxide (NO) and auxin in Cd-mediated inhibition of Arabidopsis thaliana root meristem growth. Our study demonstrated that in the first 12 h of exposure, Cd inhibits primary root elongation through a decrease in the sizes of both the elongation and meristematic zones. Following Cd exposure, a decrease in auxin levels is associated with reduced PIN1/3/7 protein accumulation, but not with reduced PIN1/3/7 transcript levels. Additionally, Cd stabilized AXR3/IAA17 protein to repress auxin signalling in this Cd-mediated process. Furthermore, decreasing Cd-induced NO accumulation with either NO-specific scavenger 2-(4-carboxyphenyl)-4,4,5,5-tetramethylimidazoline-1-oxyl-3-oxide (cPTIO) or NO synthase inhibitor Nω-nitro-l-Arg-methylester (l-NAME) compromised the Cd-mediated inhibition of root meristem development, reduction in auxin and PIN1/3/7 accumulation, as well as stabilization of AXR3/IAA17, indicating that NO participates in Cd-mediated inhibition of root meristem growth. Taken together, our data suggest that Cd inhibits root meristem growth by NO-mediated repression of auxin accumulation and signalling in Arabidopsis.

https://www.onlinelibrary.wiley.com/doi/pdf/10.1111/pce.12597

Xi Huang

Papers

Inhibition of root meristem growth by cadmium involves nitric oxide-mediated repression of auxin accumulation and signalling in Arabidopsis.