Combined application of asparagine and thiourea improves tolerance to lead stress in wheat by modulating AsA-GSH cycle, lead detoxification and nitrogen metabolism

Lead (Pb), like other heavy metals, is not essentially required for optimal plant growth; however, plants uptake it from the soil, which poses an adverse effect on growth and yield. Asparagine (Asp) and thiourea (Thi) are known to assuage the negative impacts of heavy metal pollution on plant growth; however, combined application of Asp and Thi has rarely been tested to discern if it could improve wheat yield under Pb stress. Thus, this experimentation tested the role of individual and combined applications of Asp (40 mM) and Thi (400 mg/L) in improving wheat growth under lead (Pb as PbCl2, 0.1 mM) stress. Lead stress significantly reduced plant growth, chlorophyll contents and photosystem system II (PSII) efficiency, whereas it increased Pb accumulation in the leaves and roots, leaf proline contents, phytochelatins, and oxidative stress related attributes. The sole or combined application of Asp and Thi increased the vital antioxidant biomolecules/enzymes, including reduced glutathione (GSH), ascorbic acid (AsA), ascorbate peroxsidase (APX), catalase (CAT), superoxide dismutase (SOD), glutathione S-transferase (GST), dehydroascorbate reductase (DHAR), and glutathione reductase (GR). Furthermore, the sole or the combined application of Asp and Thi modulated nitrogen metabolism by stimulating the activities of nitrate and nitrite reductase, glutamate synthase (GOGAT) and glutamine synthetase (GS). Asp and Thi together led to improve plant growth and vital physiological processes, but lowered down Pb accumulation compared to those by their sole application. The results suggest that Asp and Thi synergistically can improve wheat growth under Pb-toxicity.