Under elevated atmospheric CO(2) concentrations, soil carbon (C) inputs are typically enhanced, suggesting larger soil C sequestration potential. However, soil C losses also increase and progressive nitrogen (N) limitation to plant growth may reduce the CO(2) effect on soil C inputs with time. We compiled a data set from 131 manipulation experiments, and used meta-analysis to test the hypotheses that: (1) elevated atmospheric CO(2) stimulates soil C inputs more than C losses, resulting in increasing soil C stocks; and (2) that these responses are modulated by N. Our results confirm that elevated CO(2) induces a C allocation shift towards below-ground biomass compartments. However, the increased soil C inputs were offset by increased heterotrophic respiration (Rh), such that soil C content was not affected by elevated CO(2). Soil N concentration strongly interacted with CO(2) fumigation: the effect of elevated CO(2) on fine root biomass and -production and on microbial activity increased with increasing soil N concentration, while the effect on soil C content decreased with increasing soil N concentration. These results suggest that both plant growth and microbial activity responses to elevated CO(2) are modulated by N availability, and that it is essential to account for soil N concentration in C cycling analyses.

Living trees are the main source of biogenic volatile organic compounds (BVOCs) in forest ecosystems, but substantial emissions originate from leaf and wood litter, the rhizosphere and from microorganisms. This review focuses on temperate and boreal forest ecosystems and the roles of BVOCs in ecosystem function, from the leaf to the forest canopy and from the forest soil to the atmosphere level. Moreover, emphasis is given to the question of how BVOCs will help forests adapt to environmental stress, particularly biotic stress related to climate change. Trees use their vascular system and emissions of BVOCs in internal communication, but emitted BVOCs have extended the communication to tree population and whole community levels and beyond. Future forestry practices should consider the importance of BVOCs in attraction and repulsion of attacking bark beetles, but also take an advantage of herbivore-induced BVOCs to improve the efficiency of natural enemies of herbivores. BVOCs are extensively involved in ecosystem services provided by forests including the positive effects on human health. BVOCs have a key role in ozone formation but also in ozone quenching. Oxidation products form secondary organic aerosols that disperse sunlight deeper into the forest canopy, and affect cloud formation and ultimately the climate. We also discuss the technical side of reliable BVOC sampling of forest trees for future interdisciplinary studies that should bridge the gaps between the forest sciences, health sciences, chemical ecology, conservation biology, tree physiology and atmospheric science.

Field-growing silver birch (Betula pendula Roth) clones (clone 4 and 80) were exposed to elevated CO 2 and O 3 in open-top chambers for three consecutive growing seasons (1999)(2000)(2001). At the beginning of the OTC experiment, all trees were 7 years old. We studied the single and interaction effects of CO 2 and O 3 on silver birch below-ground carbon pools (i.e. effects on fine roots and mycorrhizas, soil microbial communities and sporocarp production) and also assessed whether there are any clonal differences in these belowground CO 2 and O 3 responses. The total mycorrhizal infection level of both clones was stimulated by elevated CO 2 alone and elevated O 3 alone, but not when elevated CO 2 was used in fumigation in combination with elevated O 3 . In both clones, elevated CO 2 affected negatively light brown/orange mycorrhizas, while its effect on other mycorrhizal morphotypes was negligible. Elevated O 3 , instead, clearly decreased the proportions of black and liver-brown mycorrhizas and increased that of light brown/ orange mycorrhizas. Elevated O 3 had a tendency to decrease standing fine root mass and sporocarp production as well, both of these O 3 effects mainly manifesting in clone 4 trees. CO 2 and O 3 treatment effects on soil microbial community composition (PLFA, 2-and 3-OH-FA profiles) were negligible, but quantitative PLFA data showed that in 2001 the PLFA fungi : bacteria-ratio of clone 80 trees was marginally increased because of elevated CO 2 treatments. This study shows that O 3 effects were most clearly visible at the mycorrhizal root level and that some clonal differences in CO 2 and O 3 responses were observable in the below-ground carbon pools. In conclusion, the present data suggests that CO 2 effects were minor, whereas increasing tropospheric O 3 levels can be an important stress factor in northern birch forests, as they might alter mycorrhizal morphotype assemblages, mycorrhizal infection rates and sporocarp production.

Climate change in the boreal forests include, e.g., warming, increased tropospheric ozone concentration, higher nitrogen (N) deposition and increased risk of insect outbreaks. Climate change influences emissions of biogenic volatile organic compounds (BVOCs) affecting plant defense, communication and atmospheric feedbacks. We studied the effects of elevated temperature (ca. 1°C), elevated ozone (ca. 1.5 9 ambient), two soil N availability levels (prevailing and 120 kg N ha -1 a -1 ) and herbivory on BVOC emission rates, net photosynthesis and resin canals (BVOCs storage), of Scots pine (Pinus sylvestris) seedlings in an open-field exposure in central Finland. Shoot BVOCs were collected in July 2012 within a few days after feeding by larvae of pine-sawfly Acantholyda posticalis, a month later in August, and in May 2013. Elevated temperature caused twofold to fourfold increases in total emissions of non-oxygenated monoterpenes (MTs), oxygenated MTs and sesquiterpenes (SQTs) and several reactive compounds, and higher N enhanced some of these changes. Ozone and higher N together increased emissions of several MTs and total SQTs. Higher number of resin canals and higher net photosynthesis might have contributed to BVOC increases. Herbivory had the strongest effect on SQT emissions (threefold increase) shortly after feeding. In the following spring, herbivory reduced emission rates of some MTs, but also synergistically increased MTs emissions with temperature but suppressed the increase caused by ozone. Results suggest that warming and ozone, particularly in areas with increased soil N availability, can increase BVOC emissions from young boreal forests in the near future, and herbivory may modify these responses.

There is a need to incorporate the effects of herbivore damage into future models of plant volatile organic compound (VOC) emissions at leaf or canopy levels. Short-term (a few seconds to 48 h) changes in shoot VOC emissions of silver birch (Betula pendula Roth) in response to feeding by geometrid moths (Erannis defoliaria Hübner) were monitored online by proton transfer reaction time-of-flight mass spectrometry (PTR-TOF-MS). In addition, two separate field experiments were established to study the effects of long-term foliage herbivory (FH, 30-32 days of feeding by geometrids Agriopis aurantiaria (Clerck) and E. defoliaria in two consecutive years) and bark herbivory (BH, 21 days of feeding by the pine weevil (Hylobius abietis L.) in the first year) on shoot and rhizosphere VOC emissions of three silver birch genotypes (gt14, gt15 and Hausjärvi provenance). Online monitoring of VOCs emitted from foliage damaged by geometrid larvae showed rapid bursts of green leaf volatiles (GLVs) immediately after feeding activity, whereas terpenoid emissions had a tendency to gradually increase during the monitoring period. Long-term FH caused transient increases in total monoterpene (MT) emissions from gt14 and sesquiterpene (SQT) emissions from Hausjärvi provenance, mainly in the last experimental season. In the BH experiment, genotype effects were detected, with gt14 trees having significantly higher total MT emissions compared with other genotypes. Only MTs were detected in the rhizosphere samples of both field experiments, but their emission rates were unaffected by genotype or herbivory. The results suggest that silver birch shows a rapid VOC emission response to short-term foliage herbivory, whereas the response to long-term foliage herbivory and bark herbivory is less pronounced and variable at different time points.

We assessed the effects of ambient solar ultraviolet (UV) radiation on below-ground parameters in an arctic heath in north-eastern Greenland. We hypothesized that the current UV fluxes would reduce root biomass and mycorrhizal colonization and that these changes would lead to lower soil microbial biomass and altered microbial community composition. These hypotheses were tested on cored soil samples from a UV reduction experiment with three filter treatments (Mylar, 60% UV-B reduction; Lexan, up to 90% UV-B reduction 1 UV-A reduction; UV transparent Teflon, filter control) and an open control treatment in two study sites after 3 years' manipulation. Reduction of both UV-A and UV-B radiation caused over 30% increase in the root biomass of Vaccinium uliginosum, which was the dominant plant species. UV reduction had contrasting effects on ericoid mycorrhizal colonization of V. uliginosum roots in the two sites, while it had no clear effects on fungal (ergosterol) or microbial biomass (measured both with fumigation-extraction and quantitative lipid biomarker analysis) in soil. However, principal component analysis of lipid biomarkers (phospholipid and glycolipid fatty acid profiles) showed that microbial community composition was altered by UV reduction. Although the UV responses were slight considering the large dose difference between the treatments (from near-ambient to up to 90% UV-B reduction), we cannot rule out the possibility that the recovery of ozone layer would change the below-ground functioning of arctic ecosystems.

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Summary Young Scots pine trees naturally established at a pine heath were exposed to two concentrations of CO2 (ambient and doubled ambient) and two O3 regimes (ambient and doubled ambient) and their combination in open‐top field chambers during growing seasons 1994, 1995 and 1996 (late May to 15 September). Filtered ozone treatment and chamberless control trees were also included in the treatment comparisons. Root ingrowth cores were inserted to the undisturbed soil below the branch projection of each tree at the beginning of the fumigation period in 1994 and were harvested at the end of the fumigation periods in 1995 and 1996. Root biomasses were determined from different soil layers in the ingrowth cores, and the infection levels of different mycorrhizal types were calculated. Elevated O3 and CO2 did not have significant effects on the biomass production of Scots pine coarse (Ø > 2 mm) or fine roots (Ø < 2 mm) and roots of grasses and dwarf shrubs. Elevated O3 caused a transient stimulation, observable in 1995, in the proportion of tuber‐like mycorrhizas, total mycorrhizas and total short roots but this stimulation disappeared during the last study year. Elevated CO2 did not enhance carbon allocation to root growth or mycorrhiza formation, although a diminishing trend in the mycorrhiza formation was observed. In the combination treatment increased CO2 inhibited the transient stimulating effect of ozone, and a significant increase of old mycorrhizas was observed. Our conclusion is that doubled CO2 is not able to increase carbon allocation to growth of fine roots or mycorrhizas in nutrient poor forest sites and realistically elevated ozone does not cause a measurable limitation to roots within a period of three exposure years.