Heavy metals and arsenic stress in food crops: Elucidating antioxidative defense mechanisms in hyperaccumulators for food security, agricultural sustainability, and human health

Abstract

The spread of heavy metal(loid)s at soil-food crop interfaces has become a threat to sustainable agricultural productiv-ity, food security, and human health. The eco-toxic effects of heavy metals on food crops can be manifested through reactive oxygen species that have the potential to disturb seed germination, normal growth, photosynthesis, cellular Abbreviations: ABA, abscisic acid; ABC, ATP-binding cassette family of transporters; ACBP, acyl-CoA-binding protein; ACC, 1-aminocyclopropane-1-carboxylic acid deaminase; APX, ascorbate peroxidase; AqpS, aquaglyceroporin; ARC, International Agency for Research on Cancer; AsA-GSH, ascorbate glutathione; AsA, ascorbate; ATSDR, US Agency for Toxic substances and Disease Registry; BS, brassinosteroids; CAT, catalase; CDF, cation diffusion facilitator; COPT, copper cation protein transporters; CRISPR-Cas9, clustered regularly interspaced short palindromic repeats nucleases; DHAR, dehydroascorbate reductase; EBS, 24-epibrassinolide; EDCs, endocrine disrupting chemicals; EPS, extracellular polymeric substances; ETS, electron transport system; GA, gibberellic acid; GPX, guaiacol peroxidase; GR, glucocorticoid receptor; GR, glutathione reductase; GS, glutathione synthetase; GSH, glutathione; GST, glutathione S-transferase; H2O2, hydrogen peroxide; HM-As, heavy metals and arsenic; HM, heavy metals; HMAs, heavy metal ATPases; HMW, high molecular weight; IAA, indole acetic acid; JA, jasmonic acid; LMW, low molecular weight; MAPK, mitogen-activated protein kinases; MAS, marker-assisted selection; MDA, malondialdehyde; MDHAR, monodehydroascorbate reductase; MFS, major facilitator superfamily; miRNAs, microRNAs; MPs, microplastics; MTP, metal transport protein; NGS, next generation sequencing; NIPs, nodulin 26-like intrinsic proteins; NMs, nanomaterials; NRAMPs, natural resistance associ-ated macrophage proteins; PAL, phenylalanine ammonia-lyase; PC, phytochelatin; PCD, programmed cell death; PCS, phytochelatin synthase; PGPR, plant growth promoting rhizobacteria; PHT, phosphate transporters; POD, peroxidase; PPCPs, pharmaceutical and personal care products; PUFA, polyunsaturated fatty acids; RNS, reactive nitrogen species; ROS, reactive oxygen species; SA, salicylic acid; SBPs, SQUAMOSA promoter binding proteins; Se, selenium; SH, sulf-hydryl; Si, silicon; SMRT, single-molecular real-time; SOD, superoxide dismutase; SSNs, sequence-specific nu-cleases; TALENs, transcription activator-like effector nucleases; TBARS, thiobarbituric acid-reactive substances; TFs, transcription factors; TIR1, transport inhibitor response 1 gene; VIT, vacuolar iron transporter; YCF, yeast cadmium factor; YSL, yellow stripe like; ZFNs, zinc finger nucleases; ZIF, zinc induced facilitator; ZIP, zinc regulated transporter and iron regulated transporter; gamma-ECS, glutamyl-cysteinyl synthetase.