Agricultural soil degradation is occurring at unprecedented rates, not only as an indirect effect of climate change (CC) but also due to intensified agricultural practices which affect soil properties and biodiversity. Therefore, understanding the impacts of CC and soil degradation on plant physiology is crucial for the sustainable development of mitigation strategies to prevent crop productivity losses. The amino acid proline has long been recognized for playing distinct roles in plant cells undergoing osmotic stress. Due to its osmoprotectant and redox-buffering ability, a positive correlation between proline accumulation and plants’ tolerance to abiotic stress has been pointed out in numerous reviews. Indeed, proline quantification is used systematically by plant physiologists as an indicator of the degree of tolerance and a measurement of the antioxidant potential in plants under stressful conditions. Moreover, the exogenous application of proline has been shown to increase resilience to several stress factors, including those related to soil degradation such as salinity and exposure to metals and xenobiotics. However, recent data from several studies often refer to proline accumulation as a signal of stress sensitivity with no clear correlation with improved antioxidant activity or higher stress tolerance, including when proline is used exogenously as a stress reliever. Nevertheless, endogenous proline levels are strongly modified by these stresses, proving its involvement in plant responses. Hence, one main question arises—is proline augmentation always a sign of improved stress resilience? From this perspective, the present review aims to provide a more comprehensive understanding of the implications of proline accumulation in plants under abiotic stress induced by soil degradation factors, reinforcing the idea that proline quantification should not be employed as a sole indicator of stress sensitivity or resilience but rather complemented with further biochemical and physiological endpoints.
Solanum nigrum L. plants were exposed for 28 days to 100 and 200 μmol/L copper (Cu) in a hydroponic system to analyze the antioxidant defense response. A dose‐dependent reduction in growth (fresh mass of root and shoot, shoot height, and root elongation) with increasing concentration of Cu was observed, whereas Cu treatments did funt affect total chlorophyll and carotefunids content. An enhanced lipid peroxidation, in terms of malondialdehyde (MDA) content, was quantified in shoots when the plants were subjected to the highest Cu level, while in roots MDA levels showed a dose‐dependent increase along the increasing Cu concentrations applied. An increase of proline in roots of plants exposed to 200 μmol/L Cu was found. Antioxidant enzymes superoxide dismutase (SOD) and catalase (CAT) did funt show significant changes with respect to control, in both roots and shoots, despite mRNA‐specific accumulations varied between Cu levels and organs. Ascorbate peroxidase (APX) was negatively affected in shoots by the highest Cu level. Gene expression of the subtype 2d metallothioneins (MT) revealed to be Cu‐enhanced throughout the plant body and correlated with Cu tissue levels, with the other MT1 and MT2 gene members downregulated in roots and upregulated in shoots, contributing more as antioxidants in the latter organs than in Cu homeostasis. MT3s are not involved in Cu homeostasis and phytochelatin (PC) production was enhanced in roots of plants exposed to 200 μmol/L Cu, contributing to a higher Cu accumulation in these organs.