Metabolic and physiological responses of Mediterranean high‐mountain and Alpine plants to combined abiotic stresses

High-mountain areas provide excellent opportunities to study the effects of combined abiotic stresses on plant physiology given their variety of steep ecological gradients, low anthropogenic disturbance and remarkable levels of taxonomic diversity. Efficient photoprotective and antioxidant scavenging mechanisms are vital for survival in high-mountain plants, having its altitudinal and seasonal variations determined by environmental or ontogenetic factors such as the decrease in mean temperatures and water availability. A number of stress indicators have been described in order to rapidly assess plant fitness in high-mountain environments. For instance, carbon isotope (δ13 C) and proline content as drought and temperature stress indicators, because of their link to water-use efficiency and osmotic adjustment; photosynthetic pigments, related to phenology, nutrient status, light and temperature stress; and non-structural carbohydrate accumulation in response to mild or brief drought conditions. The present review unveils the wide research opportunities available for the study of adaptive responses in high-mountain plants via stress indicators, and calls attention to the substantial knowledge gap existing between alpine zones and other mountainous regions, such as Mediterranean high-mountains. The aim is to grant a more holistic understanding of the physiological mechanisms driving plant life in high altitudes and improve the predictions of the effects of changing environments in these species and across ecological scales.