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.