Seasonal ionomic and metabolic changes in Aleppo pines growing on mine tailings under Mediterranean semi-arid climate

Aleppo pine is the most abundant conifer species in Mediterranean basin. Knowledge of adaptive mechanisms to cope with different environmental stresses simultaneously is necessary to improve its resilience to the predicted climatic changes and anthropogenic stressors, such as heavy metal/metal(loid)s (HMMs) pollution. Here, one year-old needles and rhizosphere soil samples from five mining and non-mining (NM) populations of Aleppo pines grown spontaneously in SE Spain were sampled in two consecutive years during spring and summer. Quantitative determination of a wide suite of edaphic, biochemical, and physiological parameters was performed, including soil physicochemical properties, ionome profile, foliar redox components, primary and secondary metabolites. Mining rhizosphere soils were characterized by elevated contents of HMMs, particularly lead and zinc, and low carbon, nitrogen and potassium levels. Multivariate data analysis based on needle ionome and antioxidative/oxidative parameters revealed a clear distinction between seasons irrespective of the population considered. Spring needles were characterized by higher levels of HMMs, sulfur, glutathione (GSH), proanthocyanidins (PAs), and soluble phenols (TPC), whereas reduced chlorophylls and increased levels of carotenoids, relative water content and K⁺, Na⁺ and Cl^- typified summer needles. In general mining populations had higher levels of ascorbate, and TPC, and exhibited higher antioxidant activities than the NM population. This could contribute to prevent oxidative injury induced by HMMs. Taken together, results suggest that seasonal factors have a more marked effect on the metabolism of the Aleppo pine populations studied than that exerted by soil conditions. This effect could be mediated by water availability in surface soil layers. If this conclusion is right, predicted rainfall reduction and temperature increase in the Mediterranean basin associated to global climate change would lead to pine needle metabolism to express the summer pattern for more prolonged periods. This, in turn, could negatively affect the performance of Aleppo pine populations.