Volatile responses of dwarf birch to mimicked insect herbivory and experimental warming at two elevations in Greenlandic tundra.

Plants release a complex blend of volatile organic compounds (VOCs) in response to stressors. VOC emissions vary between contrasting environments and increase with insect herbivory and rising temperatures. However, the joint effects of herbivory and warming on plant VOC emissions are understudied, particularly in high latitudes, which are warming fast and facing increasing herbivore pressure. We assessed the individual and combined effects of chemically mimicked insect herbivory, warming, and elevation on dwarf birch (Betula glandulosa) VOC emissions in high-latitude tundra ecosystems in Narsarsuaq, South Greenland. We hypothesized that VOC emissions and compositions would respond synergistically to warming and herbivory, with the magnitude differing between elevations. Warming increased emissions of green leaf volatiles (GLVs) and isoprene. Herbivory increased the homoterpene, (E)-4,8-dimethyl-1,3,7-nonatriene, emissions, and the response was stronger at high elevation. Warming and herbivory had synergistic effects on GLV emissions. Dwarf birch emitted VOCs at similar rates at both elevations, but the VOC blends differed between elevations. Several herbivory-associated VOC groups did not respond to herbivory. Harsher abiotic conditions at high elevations might not limit VOC emissions from dwarf birch, and high-elevation plants might be better at herbivory defense than assumed. The complexity of VOC responses to experimental warming, elevation, and herbivory are challenging our understanding and predictions of future VOC emissions from dwarf birch-dominated ecosystems.

Synergistic effects of insect herbivory and changing climate on plant volatile emissions in the subarctic tundra.

Climate change increases the insect abundance, especially in Arctic ecosystems. Insect herbivory also significantly increases plant emissions of volatile organic compounds (VOCs), which are highly reactive in the atmosphere and play a crucial role in atmospheric chemistry and physics. However, it is unclear how the effects of insect herbivory on VOC emissions interact with climatic changes, such as warming and increased cloudiness. We assessed how experimental manipulations of temperature and light availability in subarctic tundra, that had been maintained for 30 years at the time of the measurements, affect the VOC emissions from a widespread dwarf birch (Betula nana) when subjected to herbivory by local geometrid moth larvae, the autumnal moth (Epirrita autumnata) and the winter moth (Operophtera brumata). Warming and insect herbivory on B. nana stimulated VOC emission rates and altered the VOC blend. The herbivory-induced increase in sesquiterpene and homoterpene emissions were climate-treatment-dependent. Many herbivory-associated VOCs were more strongly induced in the shading treatment than in other treatments. We showed generally enhanced tundra VOC emissions upon insect herbivory and synergistic effects on the emissions of some VOC groups in a changing climate, which can have positive feedbacks on cloud formation. Furthermore, the acclimation of plants to long-term climate treatments affects VOC emissions and strongly interacts with plant responses to herbivory. Such acclimation complicates predictions of how climate change, together with interacting biotic stresses, affects VOC emissions in the high latitudes.

Seasonal and elevational variability in the induction of specialized compounds from mountain birch (Betula pubescens var. pumila) by winter moth larvae (Operophtera brumata).

The mountain birch [Betula pubescens var. pumila (L.)] forest in the Subarctic is periodically exposed to insect outbreaks, which are expected to intensify due to climate change. To mitigate abiotic and biotic stresses, plants have evolved chemical defenses, including volatile organic compounds (VOCs) and non-volatile specialized compounds (NVSCs). Constitutive and induced production of these compounds, however, are poorly studied in Subarctic populations of mountain birch. Here, we assessed the joint effects of insect herbivory, elevation and season on foliar VOC emissions and NVSC contents of mountain birch. The VOCs were sampled in situ by an enclosure technique and analyzed by gas chromatography-mass spectrometry. NVSCs were analyzed by liquid chromatography-mass spectrometry using an untargeted approach. At low elevation, experimental herbivory by winter moth larvae (Operophtera brumata) increased emissions of monoterpenes and homoterpenes over the 3-week feeding period, and sesquiterpenes and green leaf volatiles at the end of the feeding period. At high elevation, however, herbivory augmented only homoterpene emissions. The more pronounced herbivory effects at low elevation were likely due to higher herbivory intensity. Of the individual compounds, linalool, ocimene, 4,8-dimethylnona-1,3,7-triene, 2-methyl butanenitrile and benzyl nitrile were among the most responsive compounds in herbivory treatments. Herbivory also altered foliar NVSC profiles at both low and high elevations, with the most responsive compounds likely belonging to fatty acyl glycosides and terpene glycosides. Additionally, VOC emissions from non-infested branches were higher at high than low elevation, particularly during the early season, which was mainly driven by phenological differences. The VOC emissions varied substantially over the season, largely reflecting the seasonal variations in temperature and light levels. Our results suggest that if insect herbivory pressure continues to rise in the mountain birch forest with ongoing climate change, it will significantly increase VOC emissions with important consequences for local trophic interactions and climate.

Insect Herbivory Strongly Modifies Mountain Birch Volatile Emissions.

Insect herbivory is known to augment emissions of biogenic volatile organic compounds (BVOCs). Yet few studies have quantified BVOC responses to insect herbivory in natural populations in pan-Arctic regions. Here, we assess how quantitative and qualitative BVOC emissions change with increasing herbivore feeding intensity in the Subarctic mountain birch (Betula pubescens var pumila (L.)) forest. We conducted three field experiments in which we manipulated the larval density of geometrid moths (Operophtera brumata and Epirrita autumnata), on branches of mountain birch and measured BVOC emissions using the branch enclosure method and gas chromatography-mass spectrometry. Our study showed that herbivory significantly increased BVOC emissions from the branches damaged by larvae. BVOC emissions increased due to insect herbivory at relatively low larvae densities, causing up to 10% of leaf area loss. Insect herbivory also changed the blend composition of BVOCs, with damaged plants producing less intercorrelated BVOC blends than undamaged ones. Our results provide a quantitative understanding of the relationship between the severity of insect herbivore damage and emissions of BVOCs at larvae densities corresponding to background herbivory levels in the Subarctic mountain birch. The results have important and practical implications for modeling induced and constitutive BVOC emissions and their feedbacks to atmospheric chemistry.