Long-term effects of elevated CO2, nighttime warming and drought on plant secondary metabolites in a temperate heath ecosystem.

BACKGROUND AND AIMS: Plant secondary metabolites play critical roles in plant stress tolerance and adaptation, and are known to be influenced by the environment and climate changes, yet the impacts and interactions of multiple climate change components are poorly understood, particularly under natural conditions. METHODS: Accumulation of phenolics and emissions of volatile organic compounds (VOCs) were assessed on heather, Calluna vulgaris, an abundant evergreen dwarf shrub in European heathlands, after 6 years of exposure to elevated CO2, summer drought and nighttime warming. KEY RESULTS: Drought alone had the strongest effects on phenolic concentrations and compositions, with moderate effects of elevated CO2 and temperature. Elevated CO2 exerted the greatest impact on VOC emissions, mainly by increasing monoterpene emissions. The response magnitudes varied among plant tissue types and chemical constituents, and across time. With respect to interactive effects of the studied climate change components, the interaction between drought and elevated CO2 was most apparent. Drought mainly reduced phenolic accumulation and VOC emissions, while elevated CO2 mitigated such effects. CONCLUSIONS: In natural ecosystems, co-occurring climate factors can exert complex impacts on plant secondary metabolite profiles, which may in turn alter ecosystem processes.

Impact of warming, moderate nitrogen addition and bark herbivory on BVOC emissions and growth of Scots pine (Pinus sylvestris L.) seedlings.

The changing climate will expose boreal forests to rising temperatures, increasing soil nitrogen (N) levels and an increasing risk of herbivory. The single and interaction effects of warming (+2 °C increase), moderate N addition (30 kg ha-1 year-1) and bark herbivory by large pine weevil (Hylobius abietis L.) on growth and emissions of biogenic volatile organic compounds (BVOCs) from shoots of Scots pine (Pinus sylvestris L.) seedlings were studied in growth chambers over 175 days. In addition, warming and N addition effects on shoot net photosynthesis (Pn) were measured. Nitrogen addition increased both shoot and root dry weights, whereas warming, in combination with herbivory, reduced stem height growth. Warming together with N addition increased current-year shoot Pn, whereas N effects on previous-year shoot Pn were variable over time. Warming decreased non-oxygenated monoterpene (MT) emissions in June and increased them in July. Of individual MT compounds, α-pinene, δ-3-carene, γ-terpinene and terpinolene were among the most frequently responsive compounds in warming treatments in the May-July period. Sesquiterpene emissions were observed only from warming treatments in July. Moderate N addition increased oxygenated monoterpenes in May, and MTs in June and September. However, N addition effect on MTs in June was clearer without warming than with warming. Bark herbivory tended to increase MT emissions in combination with warming and N addition 3 weeks after the damage caused by weevils. Of individual compounds in other BVOC blends, herbivory increased the emissions of methyl-benzene, benzene and hexanal in July. Hence, though both warming and N addition have a potential to change BVOC emissions from Scots pines, the N effect may also be partly cancelled by warming. Furthermore, herbivory pressure in combination with climate warming and N addition may, at least periodically, increase BVOC release to the atmosphere from young Scots pine seedlings.

Monoterpene emissions in response to long-term night-time warming, elevated CO2 and extended summer drought in a temperate heath ecosystem.

Monoterpenes emitted from plants have an important role in atmospheric chemistry through changing atmospheric oxidative capacity, forming new particles and secondary organic aerosols. The emission rates and patterns can be affected by changing climate. In this study, emission responses to six years of climatic manipulations (elevated CO2, extended summer drought and night-time warming) were investigated in a temperate semi-natural heath ecosystem. Samples for monoterpene analysis were collected in seven campaigns during an entire growing season (April-November, 2011). The results showed that the temperate heath ecosystem was a considerable source of monoterpenes to the atmosphere, with the emission averaged over the 8month measurement period of 21.7±6.8µgm-2groundareah-1 for the untreated heath. Altogether, 16 monoterpenes were detected, of which the most abundant were α-pinene, δ-3-carene and limonene. The emissions of these three compounds were positively correlated with light, chamber temperature and litter abundance, but negatively correlated with soil temperature. Elevated CO2 tended to decrease the average monoterpene emissions by 40% over the whole growing season, and significantly reduced emissions in August. Extended summer drought significantly decreased the emission right after the drought treatment period, but also in the late growing season. Night-time warming significantly increased the total emissions (mainly α-pinene) in April, and tended to mitigate the decrease caused by drought. The inhibition effects of elevated CO2 on emissions were diminished when the treatment was combined with drought or warming. The emission responses to different treatments were not explained by vegetation changes, and the monoterpene emission profile was only moderately related to plant species coverage. The emission responses to these long-term climate manipulations varied over the growing season (with strong correlation with litter abundance) and the observed antagonistic effects in the combined treatments underlie the importance of long-term studies with multiple factors acting in concert.

Doubled volatile organic compound emissions from subarctic tundra under simulated climate warming.

*Biogenic volatile organic compound (BVOC) emissions from arctic ecosystems are important in view of their role in global atmospheric chemistry and unknown feedbacks to global warming. These cold ecosystems are hotspots of climate warming, which will be more severe here than averaged over the globe. We assess the effects of climatic warming on non-methane BVOC emissions from a subarctic heath. *We performed ecosystem-based chamber measurements and gas chromatography-mass spectrometry (GC-MS) analyses of the BVOCs collected on adsorbent over two growing seasons at a wet subarctic tundra heath hosting a long-term warming and mountain birch (Betula pubescens ssp. czerepanovii) litter addition experiment. *The relatively low emissions of monoterpenes and sesquiterpenes were doubled in response to an air temperature increment of only 1.9-2.5 degrees C, while litter addition had a minor influence. BVOC emissions were seasonal, and warming combined with litter addition triggered emissions of specific compounds. *The unexpectedly high rate of release of BVOCs measured in this conservative warming scenario is far above the estimates produced by the current models, which underlines the importance of a focus on BVOC emissions during climate change. The observed changes have implications for ecological interactions and feedback effects on climate change via impacts on aerosol formation and indirect greenhouse effects.

Climatic warming increases isoprene emission from a subarctic heath.

Emissions of isoprene, a reactive hydrocarbon, from Subarctic vegetation are not well documented. However, the Arctic is likely to experience the most pronounced effects of climatic warming, which may increase temperature-dependent isoprene emission. Here, we assessed isoprene emission from a Subarctic heath subjected to a 3-4 degrees C increase in air temperature and mountain birch (Betula pubescens ssp. czerepanovii) litter addition for 7-8 yr, simulating climatic warming and the subsequent expansion of deciduous shrub species and migration of the treeline. The measurements were performed using the dynamic chamber method on a wet heath with a mixture of shrubs, herbs and graminoids. Isoprene emissions averaged across the treatments were 36 +/- 5 microg m(-2) h(-1) in 2006 and 58 +/- 7 microg m(-2) h(-1) in 2007. The experimental warming increased the emissions by 83% in 2007 (P = 0.021) and by 56% in 2006 (P = 0.056), while litter addition had no significant effects. The net ecosystem CO(2) exchange was significantly decreased by warming in 2007. These results show that isoprene emissions from Subarctic heaths are comparable to those from Subarctic peatlands. Climatic warming will increase the emissions, and the amount of carbon lost as isoprene, from Subarctic heath ecosystems.

Isoprene emission from a subarctic peatland under enhanced UV-B radiation.

Isoprene is a reactive hydrocarbon with an important role in atmospheric chemistry, and emissions from vegetation contribute to atmospheric carbon fluxes. The magnitude of isoprene emissions from arctic peatlands is not known, and it may be altered by increasing UV-B radiation. Isoprene emission was measured with the dynamic chamber method from a subarctic peatland under long-term enhancement of UV-B radiation targeted to correspond to a 20% loss in the stratospheric ozone layer. The site type of the peatland was a flark fen dominated by the moss Warnstorfia exannulata and sedges Eriophorum russeolum and Carex limosa. The relationship between species densities and the emission was also assessed. Isoprene emissions were significantly increased by enhanced UV-B radiation during the second (2004) and the fourth (2006) growing seasons under the UV-B exposure. Emissions were related to the density of E. russeolum. The dominant moss, W. exannulata, proved to emit small amounts of isoprene in a laboratory trial. Subarctic fens, even without Sphagnum moss, are a significant source of isoprene to the atmosphere, especially under periods of warm weather. Warming of the Arctic together with enhanced UV-B radiation may substantially increase the emissions.