Ancient trees are essential elements for high-mountain forest conservation: Linking the longevity of trees to their ecological function.

Mature forests and their extremely old trees are rare and threatened ancient vestiges in remote European high-mountain regions. Here, we analyze the role that extremely long-living trees have in mature forests biodiversity in relation to their singular traits underlying longevity. Tree size and age determine relative growth rates, bud abortion, and the water status of long-living trees. The oldest trees suffer indefectible age-related constraints but possess singular evolutionary traits defined by fitness adaptation, modular autonomy, and a resilient metabolism that allow them to have irreplaceable roles in the ecosystem as biodiversity anchors of vulnerable lichen species like Letharia vulpina. We suggest that the role of ancient trees as unique biodiversity reservoirs is linked to their singular physiological traits associated with longevity. The set of evolutionarily plastic tools that can only be provided by centuries or millennia of longevity helps the oldest trees of mature forests drive singular ecological relationships that are irreplaceable and necessary for ecosystem dynamics.

Biological outliers: essential elements to understand the causes and consequences of reductions in maximum photochemical efficiency of PSII in plants.

MAIN CONCLUSION: By studying Cistus albidus shrubs in their natural habitat, we show that biological outliers can help us to understand the causes and consequences of maximum photochemical efficiency decreases in plants, thus reinforcing the importance of integrating these often-neglected data into scientific practice. Outliers are individuals with exceptional traits that are often excluded of data analysis. However, this may result in very important mistakes not accurately capturing the true trajectory of the population, thereby limiting our understanding of a given biological process. Here, we studied the role of biological outliers in understanding the causes and consequences of maximum photochemical efficiency decreases in plants, using the semi-deciduous shrub C. albidus growing in a Mediterranean-type ecosystem. We assessed interindividual variability in winter, spring and summer maximum PSII photochemical efficiency in a population of C. albidus growing under Mediterranean conditions. A strong correlation was observed between maximum PSII photochemical efficiency (Fv/Fm ratio) and leaf water desiccation. While decreases in maximum PSII photochemical efficiency did not result in any damage at the organ level during winter, reductions in the Fv/Fm ratio were associated to leaf mortality during summer. However, all plants could recover after rainfalls, thus maximum PSII photochemical efficiency decreases did not result in an increased mortality at the organism level, despite extreme water deficit and temperatures exceeding 40ºC during the summer. We conclude that, once methodological outliers are excluded, not only biological outliers must not be excluded from data analysis, but focusing on them is crucial to understand the causes and consequences of maximum PSII photochemical efficiency decreases in plants.

Physiological mechanisms underlying extreme longevity in mountain pine trees.

Ancient trees are life history longevity winners that mostly persist in remote and environmentally harsh mountainous areas. Here, we performed a multifeature analysis in a protected mature mountain pine (Pinus uncinata) forest to identify the morphological and physiological traits that make these trees unique. We compared the physiology of meristematic and somatic tissues (apical buds and needles, respectively) from juvenile, mature young, mature old, and mature ancient trees under cold stress and nonstress conditions. We successfully identified key morphological features of extreme longevity at the organism level, as well as various growth, vigor, stress, and dormancy markers underlying extreme longevity in old and ancient trees. Results indicated that evolution has exerted selective pressure on specific physiological traits that make trees become longevity winners (<0.1% of the tree population were ancient trees, with an average trunk diameter >100 cm and an estimated age of 700 years). Traits entailing longevity not only included apical dominance loss, epicormic growth, and modular senescence, but also an extreme plasticity in both meristematic and somatic tissues (buds and needles, respectively), as shown by various physiological markers. In conclusion, ancient trees are oddities that not only possess a unique ecological value but also show divergent physiological behaviors selected during their evolution to allow them to cope with adversities and attain long life.

English plantain deploys stress tolerance mechanisms at various organization levels across an altitudinal gradient in the Pyrenees.

High-mountain plants must withstand high solar irradiation and low temperatures during winter. Furthermore, climate change is increasing drought events, which pose an additional threat to plants. Here, we studied the stress tolerance mechanisms at various levels of biological organization in English plantain (Plantago lanceolata L.), focusing on photoprotective and antioxidant responses. The response of populations from three different altitudes in the Eastern Pyrenees (1030, 1380, and 1660 m. a.s.l.) was compared during both autumn and winter. Results showed that plants not only suffered from photoinhibition due to very low temperatures at the highest elevation during winter, but also from mild drought stress at the lowest altitude during autumn. Individuals growing at the highest elevation showed reductions in the maximum photochemical efficiency of PSII (Fv /Fm ratio), which might be caused by the lack of an increased induction of tolerance mechanisms at the highest elevation compared to the intermediate one. Although most leaves died at the highest elevation, plants could withstand stress at the organism level by generating new leaves once the stress ceased. Drought at the lowest elevation during autumn caused mild stress with small decreases in the Fv /Fm ratio, along with an increase in abscisic acid and jasmonic acid content. This study underlines the great capacity of English plantain to adapt to high elevation by activating not only photo- and antioxidant protection mechanisms and adjustments in stress-related phytohormones, but also by fully regenerating its aboveground biomass through renewed growth once the stress has ceased.