Transient photoinhibition and photo-oxidative stress as an integral part of stress acclimation and plant development in a dioecious tree adapted to Mediterranean ecosystems.

Mastic trees (Pistacia lentiscus L.) are dioecious perennial plants that are highly adapted to Mediterranean climates but display a high sensitivity to winter periods. In order to understand how sex, leaf phenology and ecological context could condition sensitivity to winter and associated mechanisms to acclimate to these conditions, photoinhibition and photo-oxidative stress markers were examined in mastic trees (P. lentiscus) from a natural population growing in the Garraf Natural Park for a consecutive 12-month period (seasonal study), as well as in three populations naturally growing in the Montseny Natural Park, including the highest altitudes described for this species, during winter (altitudinal study). Results from these studies indicate that both the winter period and higher elevation influenced the degree of photoinhibition, but this was not conditioned by sex. In fact, winter photoinhibition occurred transiently even though it was accompanied by chlorophyll loss and malondialdehyde contents. Stress acclimation was achieved through biochemical adjustments in chloroplasts, characterized by anthocyanin shielding, increased de-epoxidation state of the xanthophyll cycle as well as tocopherol accumulation, and phenological adaptations, the latter allowing a complete resetting of the physiological performance of leaves. Moreover, although females showed higher lipid peroxidation than males during the coldest winter months, at the highest elevation and during flowering in spring, this oxidative stress was mild and transient with no negative consequences for the physiology of plants. It is concluded that evergreen mastic trees acclimate to winter conditions and higher elevations by activation of antioxidant defenses together with phenological adjustments, altogether playing a crucial role in plant survival. Sexual dimorphism in mastic trees appears as a relevant factor when considering sensitivity to photo-oxidative stress in winter and altitudinal conditions.

Strategies for severe drought survival and recovery in a Pyrenean relict species.

In the context of future climate change new habitats will be threatened and unique species will be forced to develop different strategies to survive. Saxifraga longifolia Lapeyr. is an endemic species from the Pyrenees with a very particular habitat. We explored the capacity and strategies of S. longifolia plants to face different severities of drought stress under both natural conditions and controlled water stress followed by a re-watering period of 20 days. Our results showed a role for abscisic acid (ABA), salicylic acid (SA) and cytokinins (CKs) in plant survival from drought stress, and as the stress increased, ABA lost significance and SA appeared to be more associated with the response mechanisms. Moreover, photo-oxidative stress markers revealed that both xanthophyll cycles played a photoprotection role with a stronger participation of the lutein epoxide cycle as the stress was more intense. Severe drought decreased the maximum efficiency of photosystem II (Fv /Fm ) below 0.45, being this the limit to survive upon rewatering. Overall, our results proved different strategies of S. longifolia plants to cope with drought stress and suggested a Fv /Fm threshold to predict plant survival in high-mountain environments.

Plasticity in the growth habit prolongs survival at no physiological cost in a monocarpic perennial at high altitudes.

BACKGROUND AND AIMS: Monocarpic plants are those that flower, produce seeds and then die. Although most monocarpic plants are annual or biennial, some of them are perennial. However, relatively little is known regarding the biology of monocarpic perennials. Pyrenean saxifrage (Saxifraga longifolia) is a monocarpic perennial that is well adapted to high-mountain ecosystems. Here, we evaluated altitudinal changes in clonality in various populations growing in their natural habitat with particular emphasis on the physiological costs of clonal growth. METHODS: We assessed the percentage of clonal plants in nine populations growing in their natural habitat, as well as the plant stress response of clonal and non-clonal plants, in terms of photoprotection and accumulation of stress-related phytohormones, in a 3-year study at Las Blancas (2100 m a.s.l.). We also evaluated the influence of plant size on the activation of defensive responses to biotic and abiotic stresses. KEY RESULTS: We found that 12 % of Pyrenean saxifrage plants growing at the highest altitudes (2100 m a.s.l.) produced lateral rosettes which survived the flowering of the main rosette and shared the same axonomorphic root, thus escaping monocarpic senescence. This clonal growth did not worsen the physiological performance of plants growing at this altitude. Furthermore, increased plant size did not negatively affect the physiology of plants, despite adjustments in endogenous stress-related phytohormones. In contrast, maturity led to rapid physiological deterioration of the rosette, which was associated with monocarpic senescence. CONCLUSIONS: This study shows that the evolution of clonality has allowed Pyrenean saxifrage to survive harsh environmental conditions and it provides evidence that harsh environments push plant species to their limits in terms of life form and longevity.

Drought stress memory in the photosynthetic mechanisms of an invasive CAM species, Aptenia cordifolia.

Plants are known for their high capacity to acclimatise to fluctuating environmental conditions. A wide range of environmental conditions can lead to suboptimal physiological efficiency. However, recent studies have shown that plants can withstand repeated periods of stress. To find out how they do it, we studied photosynthetic adjustments to repeated water stress in Aptenia cordifolia: a facultative, invasive CAM species. Plants were subjected to three cycles of water deficit, and photosynthetic parameters and chloroplast antioxidants were quantified to gain an understanding of the mechanisms by which they cope with repeated stress periods. Significant modification of the photosystems' antenna and reaction centres was observed in plants subjected to previous water stress cycles, and this led to higher PSII efficiency than in plants challenged with drought for the first time. These findings underline the biological significance of stress memory and show how plants can adjust their photosynthetic apparatus to fluctuating environmental conditions and thus optimise photosynthesis and photoprotection under drought conditions.

Adaptation of the Long-Lived Monocarpic Perennial Saxifraga longifolia to High Altitude.

Global change is exerting a major effect on plant communities, altering their potential capacity for adaptation. Here, we aimed at unveiling mechanisms of adaptation to high altitude in an endemic long-lived monocarpic, Saxifraga longifolia, by combining demographic and physiological approaches. Plants from three altitudes (570, 1100, and 2100 m above sea level [a.s.l.]) were investigated in terms of leaf water and pigment contents, and activation of stress defense mechanisms. The influence of plant size on physiological performance and mortality was also investigated. Levels of photoprotective molecules (α-tocopherol, carotenoids, and anthocyanins) increased in response to high altitude (1100 relative to 570 m a.s.l.), which was paralleled by reduced soil and leaf water contents and increased ABA levels. The more demanding effect of high altitude on photoprotection was, however, partly abolished at very high altitudes (2100 m a.s.l.) due to improved soil water contents, with the exception of α-tocopherol accumulation. α-Tocopherol levels increased progressively at increasing altitudes, which paralleled with reductions in lipid peroxidation, thus suggesting plants from the highest altitude effectively withstood high light stress. Furthermore, mortality of juveniles was highest at the intermediate population, suggesting that drought stress was the main environmental driver of mortality of juveniles in this rocky plant species. Population structure and vital rates in the high population evidenced lower recruitment and mortality in juveniles, activation of clonal growth, and absence of plant size-dependent mortality. We conclude that, despite S. longifolia has evolved complex mechanisms of adaptation to altitude at the cellular, whole-plant and population levels, drought events may drive increased mortality in the framework of global change.

Sex-related differences in photoinhibition, photo-oxidative stress and photoprotection in stinging nettle (Urtica dioica L.) exposed to drought and nutrient deficiency.

Dimorphic plant species can show distinct nutrient needs due to sex-related differences in nutrient allocation to reproductive structures, which can potentially affect their sensitivity to photoinhibition and photo-oxidative stress. Here, we investigated sex-related differences in the extent of photo-oxidative stress in male and female individuals of U. dioica exposed to a combination of severe drought and nutrient starvation. Male and female individuals of U. dioica subject to severe drought stress were exposed to various levels of nutrient availability. First, a set of plants grown under field conditions and exposed to summer drought was used to test the effects of nutrient supply (given as NPK fertilizer). Secondly, the effects of various phosphate concentrations in the nutrient solution were tested in drought-stressed potted plants. The Fv/Fm ratio (maximum efficiency of PSII photochemistry), photoprotection capacity (levels of carotenoids, including the xanthophyll cycle, and vitamins C and E), and the extent of lipid peroxidation (hydroperoxide levels) were measured. Results showed that an application of the NPK fertilizer to the soil had a positive effect on drought-stressed plants, reducing the extent of lipid peroxidation in both males and females. P deficiency led to residual photoinhibition, as indicated by significant reductions in the Fv/Fm ratio, and enhanced lipid peroxidation in females, but not in males. We conclude that (i) increased nutrient availability in the soil can alleviate photo-oxidative stress in drought-stressed U. dioica plants, and (ii) U. dioica plants show sexual secondary dimorphism in terms of photoinhibition and photo-oxidative stress, but this is only apparent when stress infringed on plants is very severe.