RESUMEN
Tolerance mechanisms employed by plants under environmental stresses can protect them against other co-occurring stresses. In this study, the effect of pre-exposure and simultaneous salt treatment on nickel (Ni) toxicity tolerance in one halophyte (L. sativum) and one glycophyte (L. latifolium) Lepidium species in hydroponics was investigated. In order to compare the species independent from their salt and Ni tolerance level, the glycophyte was subjected to lower salt and Ni concentrations and for a shorter period of time than the halophyte. Salt (NaCl) was applied at 50 and 100 mM concentrations and Ni was provided at an equal free Ni2+ activity by adding 100 and 200 µM Ni as single stresses, but 130 and 300 µM Ni for the treatment of its combination with salt in the glycophyte and halophyte, respectively. Temporal analyses of signaling molecules revealed that the halophyte is characteristically different from the glycophyte in that it exhibits a higher constitutive level of nitric oxide and hydrogen peroxide, a longer duration of response to Ni, and its augmentation by salt. In addition to higher biomass and less Ni accumulation in salt-treated plants, the concentrations of free thiol groups, leaf pigments, proline, free and cell wall-bound phenolics contents, and the activity of phenolic metabolizing enzymes were higher in L. latifolium under the combined salt and Ni treatments than under the single Ni stress. In contrast, the biomass and most biochemical parameters of Ni-stressed L. sativum plants were not enhanced by salt treatment but rather decreased. Our findings shed light on cross-tolerance mechanisms in halophytes and uncovered halophyte survival strategies under multiple stresses.