High Dose of Nickel Unbalances Carbon Metabolism and Nitrogen Assimilation in Barley (Hordeum vulgare L.)

Pollution from heavy metals represents one of the most important threats to crops. Among these, Nickel (Ni) represents a dangerous element, strictly related to anthropic activity and easily accumulated in plants. In this study, effects of high levels (1 mM) of Ni²⁺ were investigated in barley (<i>Hordeum vulgare</i> L. cv. Nure) grown hydroponically, inducing a severe reduction in plant growth, as well as genotoxic damage. Moreover, stress affects photosynthesis, inducing a decrease in F<i>v</i>/F<i>m</i> and ΦPSII and an increase in D1 protein and RuBisCO (RbcL) abundance to compensate for the loss of photosynthetic efficiency. Changes were observed in carbon metabolism, with increases in phosphofructokinase, glyceraldehyde-3P dehydrogenase-NAD⁺, and pyruvate kinase expression confirmed by increased proteins and activities. Notably, there was an evident rise in PEP carboxylase activity, presence, and expression. This increase boosts the TCA cycle (increased fumarase) and supports photorespiration. Evident rises were observed also for glucose-6P dehydrogenase activity and presence. Ni²⁺ stress induced an evident increase in enzymes involved in nitrogen metabolism: particularly, the chloroplastic GS2/Fd-GOGAT cycle and N assimilation through the cytosolic glutamate dehydrogenase reaction were enhanced. These results design a specific stress-responsive metabolism by diverting the synthesis of N-compounds through alternative C/N assimilation pathways to counteract the effects of Ni²⁺ toxicity. This study depicts a diversion of the main C/N metabolism network towards an increase in leaf N assimilation, using carbon skeletons from dark CO₂ fixation under high Ni²⁺ stress. These results may provide possible targets for the improvement of heavy metal tolerance in cereals.