Alleviative Effect of Exogenous Application of Fulvic Acid on Nitrate Stress in Spinach (Spinacia oleracea L.)

Salt stress could be a significant factor limiting the growth and development of vegetables. In this study, Fulvic Acid (FA) (0.05%, 0.1%, 0.15%, 0.2%, and 0.25%) was applied under nitrate stress (150 mM), with normal Hoagland nutrient solution as a control to investigate the influence of foliar spray FA on spinach growth, photosynthesis, and oxidative stress under nitrate stress. The results showed that nitrate stress significantly inhibited spinach growth, while ROS (reactive oxygen species) accumulation caused photosystem damage, which reduced photosynthetic capacity. Different concentrations of FA alleviated the damage caused by nitrate stress in spinach to varying degrees in a concentration-dependent manner. The F3 treatment (0.15% FA + 150 mM NO3-) exhibited the most significant mitigating effect. FA application promoted the accumulation of biomass in spinach under nitrate stress and increased chlorophyll content, the net photosynthetic rate, the maximum photochemical quantum yield of PSII (Photosystem II) (Fv/Fm), the quantum efficiency of PSII photochemistry [Y(II)], the electron transport rate, and the overall functional activity index of the electron transport chain between the PSII and PSI systems (PItotal); moreover, FA decreased PSII excitation pressure (1 - qP), quantum yields of regulated energy dissipation of PSII [Y(NPQ)], and the relative variable initial slope of fluorescence. FA application increased superoxide dismutase, peroxidase, and catalase activities and decreased malondialdehyde, H2O2, and O2- levels in spinach under nitrate stress. FA can enhance plant resistance to nitrate by accelerating the utilization of light energy in spinach to mitigate excess light energy and ROS-induced photosystem damage and increase photosynthetic efficiency.