Tendry R. Randriamanana

Sex-related differences in growth and carbon allocation to defence in Populus tremula as explained by current plant defence theories

Plant defence theories have recently evolved in such a way that not only the quantity but also the quality of mineral nutrients is expected to influence plant constitutive defence. Recently, an extended prediction derived from the protein competition model (PCM) suggested that nitrogen (N) limitation is more important for the production of phenolic compounds than phosphorus (P). We aimed at studying sexual differences in the patterns of carbon allocation to growth and constitutive defence in relation to N and P availability in Populus tremula L. seedlings. We compared the gender responses in photosynthesis, growth and whole-plant allocation to phenolic compounds at different combination levels of N and P, and studied how they are explained by the main plant defence theories. We found no sexual differences in phenolic concentrations, but interestingly, slow-growing females had higher leaf N concentration than did males, and genders differed in their allocation priority. There was a trade-off between growth and the production of flavonoid-derived phenylpropanoids on one hand, and between the production of salicylates and flavonoid-derived phenylpropanoids on the other. Under limited nutrient conditions, females prioritized mineral nutrient acquisition, flavonoid and condensed tannin (CT) production, while males invested more in above-ground biomass. Salicylate accumulation followed the growth differentiation balance hypothesis as low N mainly decreased the production of leaf and stem salicylate content while the combination of both low N and low P increased the amount of flavonoids and CTs allocated to leaves and to a lesser extent stems, which agrees with the PCM. We suggest that such a discrepancy in the responses of salicylates and flavonoid-derived CTs is linked to their clearly distinct biosynthetic origins and/or their metabolic costs.

Interactive effects of supplemental UV-B and temperature in European aspen seedlings: Implications for growth, leaf traits, phenolic defense and associated organisms

Past studies reveal opposite effects of elevated UV-B and temperature on plant growth and concentrations of UV-B absorbing compounds, yet few studies have dealt with the combined and interactive effects of these two climate change factors on woody dioecious plants. We investigated the interactive effects of UV-B and temperature treatments on growth, leaf traits and phenolic concentrations in Populus tremula L. (European aspen) seedlings. We also considered the consequences of these effects on their associated organisms: herbivorous insects, rust pathogens, the presence of endophytic fungi and whether or not the responses differ between genders and genotypes. Supplemental temperature and UV-B were modulated to +2 °C and +30.77% above ambient conditions, respectively. Warming increased growth, photosynthesis and foliar nitrogen concentration but reduced leaf thickness and phenolic concentrations. On the other hand, supplemental UV-B increased total phenolic glycosides, mainly flavonols and phenolic acids, and partially counteracted the positive effects of warming on growth. Fast growing genotypes were less susceptible to the growth-reducing effect of combined UVB + T, less infected with rust disease and less prone to insect damage probably due to their higher salicylate and lower nitrogen concentrations. Under ambient temperature, the males of European aspen were taller and had bigger leaves than the females, while under elevated temperature, females grew bigger and, under UV-B, had more tremulacin than males. The multiple interactive effects of UV-B and temperature on growth, leaf traits and phenolic compounds, highlight the importance of multifactor experiments as a realistic predictor of plant responses to climate change.

The vegetative buds of Salix myrsinifolia are responsive to elevated UV-B and temperature.

The predicted rise in temperature and variable changes in ultraviolet-B radiation will have marked effects on plant growth and metabolism. Different vegetative parts of trees have been studied to detect the impacts of enhanced temperature and UV-B, but the effects on buds have rarely been considered. In the present study, Salix myrsinifolia clones were subjected to enhanced UV-B and temperature over two growing seasons starting from 2009, and measured springtime bud development and concentrations of phenolic compounds. In 2010 and 2011 the buds under increased temperature were up to 30% longer than those in control plots. On the other hand, UV-B combined with elevated temperature significantly decreased bud length by 4-5% in 2010. This effect was stronger in males than in females. The vegetative buds contained high constitutive amounts of chlorogenic acid derivatives, which may explain the weak increase in hyperin and chlorogenic acid that are usual UV-B sheltering compounds. The elevated temperature treatment significantly increased salicin content (about 18% in males and 22% in females), while triandrin concentration decreased by only 50% in females. Our results indicate that vegetative bud size is highly affected by seasonal temperature, while UV-B induced a weaker and transient effect.

Effect of climate change on bud phenology of young aspen plants (Populus tremula. L)

Abstract Boreal tree species are excellent tools for studying tolerance to climate change. Bud phenology is a trait, which is highly sensitive to environmental fluctuations and thus useful for climate change investigations. However, experimental studies of bud phenology under simulated climate change outdoors are deficient. We conducted a multifactorial field experiment with single (T, UVA, UVB) and combined treatments (UVA+T, UVB+T) of elevated temperature (T, +2°C) and ultraviolet‐B radiation (+30% UVB) in order to examine their impact on both male and female genotypes of aspen (Populus tremula L.). This study focuses on the effect of the treatments in years 2 and 3 after planting (2013, 2014) and follows how bud phenology is adapting in year 4 (2015), when the treatments were discontinued. Moreover, the effect of bud removal was recorded. We found that elevated temperature played a key role in delaying bud set and forcing bud break in intact individuals, as well as slightly delaying bud break in bud‐removed individuals. UVB delayed the bud break in bud‐removed males. In addition, both UVA and UVB interacted with temperature in year 3 and even in year 4, when the treatments were off, but only in male individuals. Axillary bud removal forced both bud break and bud set under combined treatments (UVA+T, UVB+T) and delayed both under individual treatments (T, UVB). In conclusion, male aspens were more responsive to the treatments than females and that effect of elevated temperature and UV radiation on bud set and bud break of aspen is not disappearing over 4‐year study period.

Effects of long-term UV-exposure and plant sex on the leaf phenoloxidase activities and phenolic concentrations of Salix myrsinifolia (Salisb.)

The accumulation of flavonoids on the leaf surface is a well-characterized protective mechanism against UV-B radiation. Other protective mechanisms, such as the induction of antioxidative enzymes and peroxidase-mediated lignification may also be important. The effects of UV-B radiation have mainly been considered in short-term studies, whereas ecologically more relevant long-term field studies are still rare. Here we examined the effects of long-term exposure to enhanced UV-B radiation on the activities of two antioxidative enzymes, polyphenol oxidase (PPO; EC 1.10.2.2 and EC 1.14.18.1) and guaiacol peroxidase (POD; EC 1.11.1.7), as well as the phenolic concentrations in two sexes of the dioecious species, Salix myrsinifolia. After three consecutive growth seasons with enhanced UV-B radiation, we found that PPO activity was decreased by UV radiation in male plants, which might explain their lower UV-B tolerance when compared to female plants. In addition, male plants had higher specific activity than did female plants under ambient conditions, supporting the idea that males of S. myrsinifolia are generally more growth-oriented than females. By contrast, neither UV treatment nor sex had significant effects on the POD activities of willows. Gender differences in the concentrations of phenolic compounds are in line with the general concept that males are less well defended than females. We suggest that the inability to increase PPO and POD activity, along with lower accumulation of UV-B absorbing compounds under UV-B exposure, might be one of the reasons why males had thinner leaves and were less tolerant of UV-B than were females.

Interannual variation in UV-B and temperature effects on bud phenology and growth in Populus tremula

Warming affects phenological processes such as spring bud break and autumnal bud set, and also growth rates of trees. Recently, it has been shown that these physiological processes also may be influenced by the ultraviolet-B (UV-B) part of the solar spectrum, and there are reasons to expect that the two environmental factors induce interactive effects when acting in concert. In this study, our aim was to elucidate how experimental enhancements in temperature and UV-B, alone and in combination, affect growth and seasonal phenology of Eurasian aspen (Populus tremula) over several growing seasons (three years). Moreover, we tested how environmentally induced changes in phenology affect the growth achieved over each season, that is, the importance of a prolonged growing season for growth yield. The plants grew in an outdoor experiment with modulated enhancements of temperature and UV-B during the growing season. Both UV-B and temperature enhancement affected bud set dates, while bud break dates were only affected by temperature enhancement. Temperature delayed bud set in all years, but gradually less over years, while UV-B yielded earlier bud set the first year but showed a delayed response the following years. Bud break was always earlier under temperature enhancement. The experimentally induced extension of the growing season in both ends had a positive effect of growth throughout the three-year period. However, the reduced responsiveness of bud set to both enhancement treatments suggest that the plants gradually acclimated to the modified climate, a finding that should also be investigated for other tree species.