BVOC Emissions From a Subarctic Ecosystem, as Controlled by Insect Herbivore Pressure and Temperature

The biogenic volatile organic compounds, BVOCs have a central role in ecosystem-atmosphere interactions. High-latitude ecosystems are facing increasing temperatures and insect herbivore pressure, which may affect their BVOC emission rates, but evidence and predictions of changes remain scattered. We studied the long-term effects of + 3 degrees C warming and reduced insect herbivory (achieved through insecticide sprayings) on mid- and late summer BVOC emissions from field layer vegetation, supplemented with birch saplings, and the underlying soil in Subarctic mountain birch forest in Finland in 2017-2018. Reduced insect herbivory decreased leaf damage by 58-67% and total ecosystem BVOC emissions by 44-72%. Of the BVOC groups, total sesquiterpenes had 70-80% lower emissions with reduced herbivory, and in 2017 the decrease was greater in warmed plots (89% decrease) than in ambient plots (34% decrease). While non-standardized total BVOC, monoterpene, sesquiterpene and GLV emissions showed instant positive responses to increasing chamber air temperature in midsummer samplings, the long-term warming treatment effects on standardized emissions mainly appeared as changes in the compound structure of BVOC blends and varied with compounds and sampling times. Our results suggest that the effects of climate warming on the total quantity of BVOC emissions will in Subarctic ecosystems be, over and above the instant temperature effects, mediated through changes in insect herbivore pressure rather than plant growth. If insect herbivore numbers will increase as predicted under climate warming, our results forecast herbivory-induced increases in the quantity of Subarctic BVOC emissions.Peer reviewe

Living, dead, and absent trees-How do moth outbreaks shape small-scale patterns of soil organic matter stocks and dynamics at the Subarctic mountain birch treeline?

Mountain birch forests (Betula pubescens Ehrh. ssp. czerepanovii) at the subarctic treeline not only benefit from global warming, but are also increasingly affected by caterpillar outbreaks from foliage-feeding geometrid moths. Both of these factors have unknown consequences on soil organic carbon (SOC) stocks and biogeochemical cycles. We measured SOC stocks down to the bedrock under living trees and under two stages of dead trees (12 and 55 years since moth outbreak) and treeless tundra in northern Finland. We also measured in-situ soil respiration, potential SOC decomposability, biological (enzyme activities and microbial biomass), and chemical (N, mineral N, and pH) soil properties. SOC stocks were significantly higher under living trees (4.1 +/- 2.1 kg m(2)) than in the treeless tundra (2.4 +/- 0.6 kg m(2)), and remained at an elevated level even 12 (3.7 +/- 1.7 kg m(2)) and 55 years (4.9 +/- 3.0 kg m(2)) after tree death. Effects of tree status on SOC stocks decreased with increasing distance from the tree and with increasing depth, that is, a significant effect of tree status was found in the organic layer, but not in mineral soil. Soil under living trees was characterized by higher mineral N contents, microbial biomass, microbial activity, and soil respiration compared with the treeless tundra; soils under dead trees were intermediate between these two. The results suggest accelerated organic matter turnover under living trees but a positive net effect on SOC stocks. Slowed organic matter turnover and continuous supply of deadwood may explain why SOC stocks remained elevated under dead trees, despite the heavy decrease in aboveground C stocks. We conclude that the increased occurrence of moth damage with climate change would have minor effects on SOC stocks, but ultimately decrease ecosystem C stocks (49% within 55 years in this area), if the mountain birch forests will not be able to recover from the outbreaks.Peer reviewe

Strong Interactive Effects of Warming and Insect Herbivory on Soil Carbon and Nitrogen Dynamics at Subarctic Tree Line

Warming will likely stimulate Arctic primary production, but also soil C and N mineralization, and it remains uncertain whether the Arctic will become a sink or a source for CO2. Increasing insect herbivory may also dampen the positive response of plant production and soil C input to warming. We conducted an open-air warming experiment with Subarctic field layer vegetation in North Finland to explore the effects of warming (+3°C) and reduced insect herbivory (67% reduction in leaf damage using an insecticide) on soil C and N dynamics. We found that plant root growth, soil C and N concentrations, microbial biomass C, microbial activity, and soil NH4+ availability were increased by both warming and reduced herbivory when applied alone, but not when combined. Soil NO3– availability increased by warming only and in-situ soil respiration by reduced herbivory only. Our results suggest that increasing C input from vegetation under climate warming increases soil C concentration, but also stimulates soil C turnover. On the other hand, it appears that insect herbivores can significantly reduce plant growth. If their abundance increases with warming as predicted, they may curtail the positive effect of warming on soil C concentration. Moreover, our results suggest that temperature and herbivory effects on root growth and soil variables interact strongly, which probably arises from a combination of N demand increasing under lower herbivory and soil mineral N supply increasing under higher temperature. This may further complicate the effects of rising temperatures on Subarctic soil C dynamics.</p

BVOC Emissions From a Subarctic Ecosystem, as Controlled by Insect Herbivore Pressure and Temperature

The biogenic volatile organic compounds, BVOCs have a central role in ecosystem-atmosphere interactions. High-latitude ecosystems are facing increasing temperatures and insect herbivore pressure, which may affect their BVOC emission rates, but evidence and predictions of changes remain scattered. We studied the long-term effects of + 3 degrees C warming and reduced insect herbivory (achieved through insecticide sprayings) on mid- and late summer BVOC emissions from field layer vegetation, supplemented with birch saplings, and the underlying soil in Subarctic mountain birch forest in Finland in 2017-2018. Reduced insect herbivory decreased leaf damage by 58-67% and total ecosystem BVOC emissions by 44-72%. Of the BVOC groups, total sesquiterpenes had 70-80% lower emissions with reduced herbivory, and in 2017 the decrease was greater in warmed plots (89% decrease) than in ambient plots (34% decrease). While non-standardized total BVOC, monoterpene, sesquiterpene and GLV emissions showed instant positive responses to increasing chamber air temperature in midsummer samplings, the long-term warming treatment effects on standardized emissions mainly appeared as changes in the compound structure of BVOC blends and varied with compounds and sampling times. Our results suggest that the effects of climate warming on the total quantity of BVOC emissions will in Subarctic ecosystems be, over and above the instant temperature effects, mediated through changes in insect herbivore pressure rather than plant growth. If insect herbivore numbers will increase as predicted under climate warming, our results forecast herbivory-induced increases in the quantity of Subarctic BVOC emissions

Effects of experimental warming on Betula nana epidermal cell growth tested over its maximum climatological growth range

Numerous long-term, free-air plant growth facilities currently explore vegetation responses to the ongoing climate change in northern latitudes. Open top chamber (OTC) experiments as well as the experimental set-ups with active warming focus on many facets of plant growth and performance, but information on morphological alterations of plant cells is still scarce. Here we compare the effects of in-situ warming on leaf epidermal cell expansion in dwarf birch, Betula nana in Finland, Greenland, and Poland. The localities of the three in-situ warming experiments represent contrasting regions of B. nana distribution, with the sites in Finland and Greenland representing the current main distribution in low and high Arctic, respectively, and the continental site in Poland as a B. nana relict Holocene microrefugium. We quantified the epidermal cell lateral expansion by microscopic analysis of B. nana leaf cuticles. The leaves were produced in paired experimental treatment plots with either artificial warming or ambient temperature. At all localities, the leaves were collected in two years at the end of the growing season to facilitate between-site and within-site comparison. The measured parameters included the epidermal cell area and circumference, and using these, the degree of cell wall undulation was calculated as an Undulation Index (UI). We found enhanced leaf epidermal cell expansion under experimental warming, except for the extremely low temperature Greenland site where no significant difference occurred between the treatments. These results demonstrate a strong response of leaf growth at individual cell level to growing season temperature, but also suggest that in harsh conditions other environmental factors may limit this response. Our results provide evidence of the relevance of climate warming for plant leaf maturation and underpin the importance of studies covering large geographical scales.Peer reviewe

Effects of experimental warming on Betula nana epidermal cell growth tested over its maximum climatological growth range

Numerous long-term, free-air plant growth facilities currently explore vegetation responses to the ongoing climate change in northern latitudes. Open top chamber (OTC) experiments as well as the experimental set-ups with active warming focus on many facets of plant growth and performance, but information on morphological alterations of plant cells is still scarce. Here we compare the effects of in-situ warming on leaf epidermal cell expansion in dwarf birch, Betula nana in Finland, Greenland, and Poland. The localities of the three in-situ warming experiments represent contrasting regions of B. nana distribution, with the sites in Finland and Greenland representing the current main distribution in low and high Arctic, respectively, and the continental site in Poland as a B. nana relict Holocene microrefugium. We quantified the epidermal cell lateral expansion by microscopic analysis of B. nana leaf cuticles. The leaves were produced in paired experimental treatment plots with either artificial warming or ambient temperature. At all localities, the leaves were collected in two years at the end of the growing season to facilitate between-site and within-site comparison. The measured parameters included the epidermal cell area and circumference, and using these, the degree of cell wall undulation was calculated as an Undulation Index (UI). We found enhanced leaf epidermal cell expansion under experimental warming, except for the extremely low temperature Greenland site where no significant difference occurred between the treatments. These results demonstrate a strong response of leaf growth at individual cell level to growing season temperature, but also suggest that in harsh conditions other environmental factors may limit this response. Our results provide evidence of the relevance of climate warming for plant leaf maturation and underpin the importance of studies covering large geographical scales.</p

Strong Interactive Effects of Warming and Insect Herbivory on Soil Carbon and Nitrogen Dynamics at Subarctic Tree Line

Warming will likely stimulate Arctic primary production, but also soil C and N mineralization, and it remains uncertain whether the Arctic will become a sink or a source for CO2. Increasing insect herbivory may also dampen the positive response of plant production and soil C input to warming. We conducted an open-air warming experiment with Subarctic field layer vegetation in North Finland to explore the effects of warming (+3 degrees C) and reduced insect herbivory (67% reduction in leaf damage using an insecticide) on soil C and N dynamics. We found that plant root growth, soil C and N concentrations, microbial biomass C, microbial activity, and soil NH4+ availability were increased by both warming and reduced herbivory when applied alone, but not when combined. Soil NO3- availability increased by warming only and in-situ soil respiration by reduced herbivory only. Our results suggest that increasing C input from vegetation under climate warming increases soil C concentration, but also stimulates soil C turnover. On the other hand, it appears that insect herbivores can significantly reduce plant growth. If their abundance increases with warming as predicted, they may curtail the positive effect of warming on soil C concentration. Moreover, our results suggest that temperature and herbivory effects on root growth and soil variables interact strongly, which probably arises from a combination of N demand increasing under lower herbivory and soil mineral N supply increasing under higher temperature. This may further complicate the effects of rising temperatures on Subarctic soil C dynamics.Peer reviewe

Insect herbivory dampens Subarctic birch forest C sink response to warming

Climate warming is anticipated to make high latitude ecosystems stronger C sinks through increasing plant production. This effect might, however, be dampened by insect herbivores whose damage to plants at their background, non-outbreak densities may more than double under climate warming. Here, using an open-air warming experiment among Subarctic birch forest field layer vegetation, supplemented with birch plantlets, we show that a 2.3 degrees C air and 1.2 degrees C soil temperature increase can advance the growing season by 1-4 days, enhance soil N availability, leaf chlorophyll concentrations and plant growth up to 400%, 160% and 50% respectively, and lead up to 122% greater ecosystem CO2 uptake potential. However, comparable positive effects are also found when insect herbivory is reduced, and the effect of warming on C sink potential is intensified under reduced herbivory. Our results confirm the expected warming-induced increase in high latitude plant growth and CO2 uptake, but also reveal that herbivorous insects may significantly dampen the strengthening of the CO2 sink under climate warming. Warming is expected to increase C sink capacity in high-latitude ecosystems, but plant-herbivore interactions could moderate or offset this effect. Here, Silfver and colleagues test individual and interactive effects of warming and insect herbivory in a field experiment in Subarctic forest, showing that even low intensity insect herbivory strongly reduces C sink potential.Peer reviewe

Living, dead, and absent trees-How do moth outbreaks shape small-scale patterns of soil organic matter stocks and dynamics at the Subarctic mountain birch treeline?

Mountain birch forests (Betula pubescens Ehrh. ssp. czerepanovii) at the subarctic treeline not only benefit from global warming, but are also increasingly affected by caterpillar outbreaks from foliage-feeding geometrid moths. Both of these factors have unknown consequences on soil organic carbon (SOC) stocks and biogeochemical cycles. We measured SOC stocks down to the bedrock under living trees and under two stages of dead trees (12 and 55 years since moth outbreak) and treeless tundra in northern Finland. We also measured in-situ soil respiration, potential SOC decomposability, biological (enzyme activities and microbial biomass), and chemical (N, mineral N, and pH) soil properties. SOC stocks were significantly higher under living trees (4.1 +/- 2.1 kg m2) than in the treeless tundra (2.4 +/- 0.6 kg m2), and remained at an elevated level even 12 (3.7 +/- 1.7 kg m2) and 55 years (4.9 +/- 3.0 kg m2) after tree death. Effects of tree status on SOC stocks decreased with increasing distance from the tree and with increasing depth, that is, a significant effect of tree status was found in the organic layer, but not in mineral soil. Soil under living trees was characterized by higher mineral N contents, microbial biomass, microbial activity, and soil respiration compared with the treeless tundra; soils under dead trees were intermediate between these two. The results suggest accelerated organic matter turnover under living trees but a positive net effect on SOC stocks. Slowed organic matter turnover and continuous supply of deadwood may explain why SOC stocks remained elevated under dead trees, despite the heavy decrease in aboveground C stocks. We conclude that the increased occurrence of moth damage with climate change would have minor effects on SOC stocks, but ultimately decrease ecosystem C stocks (49% within 55 years in this area), if the mountain birch forests will not be able to recover from the outbreaks.</p