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Cobalt (Co) commonly co-exists with copper (Cu) in natural soils, but the information about their combined effects on plants is poorly available. In this study, we hydroponically investigated the combined effects of Co and Cu on two barley genotypes differing in Co toxicity tolerance to reveal the interaction pattern of these two metals. The results showed that single treatment of Co or Cu at the dose of 100 μM led to a significant decrease of growth and photosynthetic rate, and a significant increase of lipid peroxidation, ROS radicals as well as anti-oxidative enzyme (SOD, CAT and GR) activities and glutathione content, with the extent of effect being less in Yan66 than Ea52. The combined treatment of Co and Cu alleviated the toxicity of both metals in comparison with each metal treatment alone, as reflected by improved growth and photosynthesis, and much slight oxidative stress. The alleviation of metal toxicity upon combined treatment is mainly attributed to a drastic reduction of Co uptake and its translocation from roots to shoots. It may be suggested that interaction of Co and Cu on their uptake and movement in plants is antagonistic. • The interaction of Co and Cu is antagonistic. • The combined treatment of Cu and Co alleviated Co toxicity relative to Co treatment alone. • The alleviation of Co toxicity by combined treatment is attributed to the uptake and transportation of Co in plants. [ABSTRACT FROM AUTHOR]

Cobalt (Co) contamination in soils is becoming a severe issue in environment safety and crop production. Calcium (Ca) , as a macro-nutrient element, shows the antagonism with many divalent heavy metals and the capacity of alleviating oxidative stress in plants. In this study, the protective role of Ca in alleviating Co stress was hydroponically investigated using two barley genotypes differing in Co toxicity tolerance. Barley seedlings exposed to 100 µM Co showed the significant reduction in growth and photosynthetic rate, and the dramatic increase in the contents of reactive oxygen species (ROS), malondialdehyde (MDA), reduced glutathione (GSH) and oxidized glutathione (GSSG), and the activities of anti-oxidative enzymes, with Ea52 (Co-sensitive) being much more affected than Yan66 (Co-tolerant). Addition of Ca in growth medium alleviated Co toxicity by reducing Co uptake and enhancing the antioxidant capacity. The effect of Ca in alleviating Co toxicity was much greater in Yan66 than in Ea52. The results indicate that the alleviation of Co toxicity in barley plants by Ca is attributed to the reduced Co uptake and enhanced antioxidant capacity. [ABSTRACT FROM AUTHOR]

The combined effects of cobalt (Co) and copper (Cu) in their toxicity to plants is poorly studied although these two metals co-exist commonly in soil. In this study, a hydroponic experiment was carried out to investigate the effect of longer exposure of two barley genotypes differing in Co tolerance to the combined Co and Cu stress. The results confirmed the previous findings that Co accumulation in plant tissues was reduced by Cu presence, while Cu accumulation was stimulated by Co presence. Moreover, both single and combined treatments of Co and Cu reduced the mineral (Mn, Zn and K) uptake. Co and Cu applied alone or in combination at rate of 50 μM resulted in the significant reduction of plant growth and increase of oxidative stress (ROS and MDA), and meanwhile the capacity of scavenging active oxygen species (AOS) was enhanced, reflected by increased phytochelatin (PC) and glutathione (GSH and GSSG) content, as well as expression of the related genes (HvPCS1 and HvGR1). Yan66, a Co tolerant genotype was less affected in oxidative stress, and had higher AOS scavenging capacity in comparison with Ea52, a Co sensitive one. Among three HvSOD isoforms, only HvFeSOD expression was up-regulated in the combined treatment relative to control as well as the treatment of Co or Cu alone, while HvCuZnSOD and HvMnSOD were down-regulated and unaffected, respectively. In addition, the expressions of metal transporter genes (HvHMA2 , HvHMA3 and HvHMA5) varied with genotype and metal treatments, with the extent being greater in Yan66 on the whole. The results suggest that upon longer exposure to the combined stress of Co and Cu, the greater phyto-toxicity than each element alone is associated with more Cu accumulation stimulated by Co and that, the higher regulation of transporter genes observed in Yan66 could in part explain for its higher metal tolerance ability. • Co accumulation in plant tissues was reduced by Cu presence, while Cu accumulation was stimulated by Co presence. • Yan66 had higher AOS scavenging capacity than Ea52, a Co sensitive one. • The expressions of metal transporter genes varied with genotype and metal treatments. [ABSTRACT FROM AUTHOR]

Highlights • Tibetan wild barley (Hordeum spontaneum) accession XZ16 is higher salt tolerant than the cultivar Golden Promise. • In roots, miR393a, miR156d and miR172b regulating HvAFB2 / HvTIR1 , UGTs and HvAP2 are responsible for salt tolerance. • In shoots, miR319a/miR396e module, miR159a and miR172b regulating GRFs , MYB33 and HvAP2 might contribute to salt tolerance. Abstract A deep understanding of the mechanisms underlying salt tolerance should be helpful for breeding of salt-tolerant crop cultivars. Our previous study identified some wild barley accessions with high salt tolerance. In this study, small RNA and degradome sequencing was performed to identify salt-responsive microRNAs (miRNAs) and their target genes in a Tibetan wild barley accession XZ16 and a cultivar Golden Promise (GP), differing in salt tolerance. Under salt stress, XZ16 showed higher salt tolerance than GP in terms of biomass and shoot Na+ accumulation. A total of 278 and 320 miRNAs were found in roots and shoots of both genotypes, respectively. Among them, 40 and 51 miRNAs were salt-responsive in roots and shoots, respectively. In roots, miR156d, miR164a, miR393a, miR319a and miR172b targeting UGTs , NAC079 , HvAFB2 / HvTIR1 , TCP4 and HvAP2 , respectively, are probably responsible for salt tolerance. In shoots, miR159a, miR169i, miR319a/miR396e module and miR172b regulating MYB33 , NHX1/LEA7 , TCP4 , GRFs and HvAP2 , respectively, might contribute to salt tolerance. Compared with GP, XZ16 showed differential expression of miR156d, miR164a, miR169i, miR172b, miR319a, miR393a and a novel miRNA PC-miR124. We proposed that these differentially expressed miRNAs could account for the difference in salt tolerance of the two genotypes and drawn up a putative regulatory network of miRNAs to reveal molecular mechanisms of salt tolerance in barley. [ABSTRACT FROM AUTHOR]