Monoterpenes and higher terpenes may inhibit enzyme activities in boreal forest soil

Plant secondary compounds, including terpenes, potentially play an important role in controlling the decomposition process in boreal forest soil. However, the role of terpenes is not well understood, and their direct influence on enzyme activity is not well-known. The aim of this study was to examine the possible effects of common monoterpenes and higher terpenes on the activity of enzymes crucial in C, N, P, S cycling, i.e. β-glucosidase, chitinase, protease, acid phosphatase and arylsulfatase. Monoterpenes (α-pinene, carene, myrcene), diterpenes (abietic acid and colophony), and triterpene (β-sitosterol) were used. Studies were done in two environments, in vitro (studies without soil) and in vivo (studies with soil). Soil experiments were conducted using humus layers of two different birch stands, the first N-poor with high organic matter content and the second N-rich with a lower organic matter content. In general, all the terpenes studied showed inhibitory potential against enzymes in in vitro studies. In the soil incubation studies, both of the measured enzymes, chitinase and β-glucosidase, showed some decrease in activity when exposed to different terpenes. Our study suggests that terpenes modify the enzyme machinery in boreal forest soil.     

Factors Responsible for the Co-occurrence of Forested and Unforested Rock Outcrops in the Boreal Forest

Rock outcrops in the boreal forest of Québec can show either of two different states: a forested state with >25% tree cover, and an unforested state (<25% tree cover). We tested three different hypotheses that might explain the co-occurrence of forested and unforested rock outcrops: (1) differences in bedrock geology, with unforested outcrops associated to bedrock types inimical to tree growth; (2) unforested outcrops as recently disturbed sites undergoing secondary succession towards a forested state; (3) unforested outcrops as an alternative stable state to forested outcrops, induced by post-fire regeneration failure. Digitized forest inventory maps were used along with bedrock geology maps and time-since-fire maps to compare forested and unforested outcrops for bedrock geology type and date of the last fire. Field surveys were conducted on 28 outcrops (14 forested, 14 unforested) to gather information regarding tree species composition and site characteristics (thickness of the organic matter layer, percent cover of lichens, mosses and ericaceous shrubs). None of the three hypotheses explain the co-occurrence of forested and unforested rock outcrops in the boreal forest of Québec. Both outcrop types occur on the same bedrock geology types. Unforested outcrops are not recently disturbed sites in early-successional states, as no clear distinction could be made in tree species composition and date of the last fire between the two outcrop types. Forested and unforested outcrops are not alternative stable states, as unforested outcrops are unstable and cannot maintain themselves through time in the prolonged absence of fire. Hence, unforested rock outcrops could be viewed as degraded, diverging post-fire types maintained by the late Holocene disturbance regime, characterized by high fire frequencies.     

Effects of management on coarse woody debris volume and composition in boreal forests in northern Sweden

Forest management practices have led to a reduction in the volume and a change in the composition of coarse woody debris (CWD) in many forest types. This study compared CWD volume and composition in reserves and two types of managed forest in the central boreal zone of Sweden. Ten areas were surveyed, each containing clear-cut, mature managed and old-growth stands, to determine the volume of standing and lying CWD in terms of species composition, decay class and size class. Volumes of CWD on clear-cuts and in mature managed forests were high compared with previous studies. Old-growth forests (72.6 m³ ha^?1) contained a greater volume of CWD than mature managed forests (23.3 m³ ha^?1) and clear-cuts (13.6 m³ ha^?1). Differences were greatest for the larger size classes and intermediate decay stages. Despite stand ages being up to 144 years, CWD volume and composition in managed forests was more similar to clear-cuts than to old-growth forests.     

Effects of forest-fuel harvesting on the amount of deadwood on clear-cuts

Forest residue left after clear-cutting is increasingly being used as biofuel in Sweden. Deadwood is an important habitat for many species and a crucial resource for forest biodiversity. On eight clear-cuts in eastern central Sweden, the amount of Picea abies (L.) Karst. deadwood left on the ground was compared with the amount collected in piles for fuelwood extraction. It was found that 65% of the volume, 77% of the surface area and 84% of the pieces of slash were to be extracted. In another study on 23 clear-cuts in the same region, it was found that 36% of the logs left outside piles after clear-cutting were later removed. These studies indicate that a considerable amount of potential substrate is lost during slash extraction.     

Differences in the forest landscape structure along the Finnish–Russian border in southern Karelia

Abstract

This study describes forest landscape fragmentation and connectivity along the Finnish–Russian border near the Karelian Isthmus. The landscape pattern was analysed using classification data based on Landsat ETM+ and Landsat TM images in combination with systematic surveys in Finland (Finnish National Forest Inventory) (n=546) and the authors’ own fieldwork data in Russia (n=101). On the Finnish side the forest patches are significantly smaller than on the Russian side. In addition, the Finnish forests landscape is more scattered and distances between patches of the same forest type are longer. The Russian side is more dominated by broadleaved and mixed forest stands. The disparities are due to differences in forestry policy and traditions of forest practices. The growing conditions of the areas are similar. The habitat fragmentation and habitat connectivity are important issues because the Karelian Isthmus is one of three main corridors and migrating routes connecting large Russian boreal taiga forests and their fauna and flora with Finnish isolated boreal forests.     

Populus tremuloides Michx. postfire stand dynamics in the northern boreal-cordilleran ecoclimatic region of central Yukon Territory, Canada

Postfire vegetation development among 8–185-year-old stand was assessed based on 100 relevés from the northern boreal-cordilleran ecoclimatic region (61–63°N) in the central Yukon Territory, Canada. Vegetation sampling included only stands thought to have originated from postfire Populus tremuloides Michx. regeneration that occurred on well drained and low gradient sites. Seven vegetation types were recognized based on cluster analysis and Kruskal–Wallis testing. Relevé ordination using Detrended Correspondence Analysis (70% explained variance) indicated six of the vegetation types represented a secondary successional chronosequence, based on their juxtaposition and a strong correlation of the primary axis with stand age (r = 0.89, P < 0.001). No correlation (P > 0.05) occurred between stand location and age. The youngest vegetation (8–11 years) had a moderate cover of P. tremuloides and Salix spp. up to 5 m tall, with a ground cover of Ceratodon purpureus (Hedw.) Brid. and Bryum caespiticium Hedw. This vegetation was expected to result in P. tremuloides, mixed P. tremuloides and Picea glauca (Moench) Voss, and P. glauca/Hylocomium splendens forest stands with increasing age, respectively. P. tremuloides//Calamagrostis purpurascens—Arctostaphylos uva-ursi stands formed the mid-seral vegetation. Along the chronosequence, total tree, P. tremuloides, shrub, and herb cover peaked 50–70 years after stand initiation; P. glauca cover, total and nonvascular species richness, and dominance concentration gradually increased (P < 0.001); vascular plant richness decreased; bryophytes had a U-shaped abundance pattern; and total plant cover was constant through time (125%). Richness totalled 113 species with averages of 13–18 per relevé. Coarse woody debris was most abundant (maxima 100–223 m³/ha) during the first 20 years of stand development then declined to <50 m³/ha. Successionally, a stem exclusion stage occurred (years 8–18), but with a delayed peak of 2–4 years and reduced densities (1.47 stems/m²) relative to southern boreal stands. No understory suppression, and therefore, no reinitiation stage occurred. Following stem exclusion, an accelerated canopy transition stage occurred relative to southern boreal forests due to early establishment rather than better height-growth rate of P. glauca relative to P. tremuloides. P. glauca tended to equal the cover of P. tremuloides 95–100 years after stand initiation. The oldest vegetation type in the chronosequence more closely resembled old-growth than a gap dynamic stage of development, possibly because of its youthful average age of 125 years. A modification was proposed for the canopy transition stage (Chen–Popadiouk stand development model) to account for the “forced” replacement of P. tremuloides by P. glauca. Differences in stand development were attributed to the cold northern climate.     

Effects of forest restoration treatments on the abundance of bark beetles in Norway spruce forests of southern Finland

Restoration of protected areas in boreal forests frequently includes creating substantial volumes of dead wood. While this benefits a wide range of dead wood dependent invertebrate species, some of these are regarded as forest pests. Therefore, the risk of elevated levels of tree mortality in surrounding commercial forests must be considered. In a large-scale field experiment in southern Finland, we studied the effects of restoration treatments on the abundance of bark beetles within and in the vicinity of restored areas, in particular focusing on Ips typographus and Pityogenes chalcographus. The treatments applied to managed Norway spruce forests were controlled burning and partial harvesting combined with retaining 5, 30 or 60 m³/ha of cut down wood. We found that the abundance of bark beetles increased by both burning and harvesting with down wood retention, being highest where burning and harvesting had been combined. The actual volume of down wood retention had no significant effect. The effect of burning on the number of bark beetles along host tree boles was negative which suggests that burnt spruces provided a less suitable resource for bark beetles than unburnt dead spruces. The abundance of bark beetles along host trees also decreased with increasing volume of down wood retention. The abundance of P. chalographus was slightly elevated up to 50 m outside restored areas but the abundance was very low compared to that within the areas. The abundance of I. typographus was extremely low outside restored areas. We conclude that restoration treatments increase the abundance of bark beetles via increased availability of resources, but that the effect of burning is likely to be counteracted by decreased resource quality. Thus, burning might be the “safest” way to produce large quantities of dead wood. Furthermore, the fact that only few beetles were collected in adjacent areas suggests that restored areas pose little threat of serving as refugia in which bark beetle populations increase in sufficient numbers to attack live trees in adjacent forests. However, restoration actions repeated at consecutive years within a small area might enable the populations to grow to outbreak levels.     

Establishment and growth of white spruce on a boreal forest floodplain: Interactions between microclimate and mammalian herbivory

White spruce (Picea glauca (Moench) Voss) is a dominant species in late-successional ecosystems along the Tanana River, interior Alaska, and the most important commercial timber species in these boreal floodplain forests. Whereas white spruce commonly seed in on young terraces in early primary succession, the species does not become a conspicuous component of the vegetation until after 60–80 years. To address what abiotic and/or biotic factors may explain the paucity of spruce in earlier stages of succession, we examined germination and growth of planted white spruce seedlings across an environmental gradient that included variation in soil physico-chemical properties in the presence and absence of mammal browsing. The effect of browsing pressure over the first four years after planting was most noticeable on the older terraces. Likewise, direct effects of hare browsing on spruce seedling mortality were only manifested at the oldest sites. Spruce germination and survival was inversely proportional to soil cation concentrations, which was largely controlled by temperature-driven evapotranspiration. High light intensities and high air temperatures significantly reduced seedling growth, whereas variation in soil moisture only explained a significant amount of variation in seedling survival. Temperatures within the needle clusters on terminal shoots reached values that adversely affect photosynthesis (>32 °C) on multiple occasions over the growing season. We conclude that the direct (temperature) and indirect (soil chemistry) effects of high insolation are major factors constraining spruce performance on early successional terraces, and that these effects can be significantly exacerbated by mammal browsing on associated deciduous vegetation.     

Does the mutualism between wood ants (Formica rufa group) and Cinara aphids affect Norway spruce growth?

Ant–aphid mutualisms, in which ants tend aphids, which in turn provide honeydew to the ants, are widespread and have been shown to affect plant growth. In boreal forests the effect of ant–aphid mutualism on tree growth can vary with stand age, because forest clear-cutting harms the ecologically most dominant ant partner in such mutualisms, wood ants (Formica rufa group). We studied whether the mutualism between wood ants and Cinara aphids affects the growth of boreal Norway spruces (Picea abies L. Karst.) in stands of different ages. In boreal forests, conifers, unlike deciduous trees, have only few defoliating insects, and therefore we expected the growth loss of conifers due to sap sucking by aphids not to be compensated by reduced insect herbivory due to predation by wood ants. The study was conducted in medium-fertile spruce-dominated stands in eastern Finland. We used stands of four different age classes (5, 30, 60 and 100 years) and selected ten spruces heavily visited and ten spruces lightly visited by ants around five medium-sized ant mounds in each stand age class. The access of ants was blocked on half of the trees in both groups. In the 5-year-old stands, the mean annual height growth of individual heavily visited seedlings was 16.3% greater than in the ones where ant traffic was blocked, but this difference was not significant. In the 30-year-old stands, the mean annual radial growth of the heavily visited spruces was 7.3% smaller than in trees where ant traffic was blocked, and this difference was significant. The mutualism had no significant effect on the radial growth in the 60- and 100-year-old stands. In the 60-year-old stands, however, the spruces that were visited heavily prior to the beginning of the study grew significantly less relative to their past growth than the initially lightly visited trees during the study. This suggests that the ant–aphid mutualism may have long-term effects on tree growth. The ant–aphid mutualism had no significant effect on the growth at the stand level. The results indicate that ant–aphid mutualism can have a significant effect on the growth of individual spruces, but the effect is negligible at ecosystem level.     

Partitioning of soil respiration into its autotrophic and heterotrophic components by means of tree-girdling in old boreal spruce forest

Forests accumulate much less carbon than the amount fixed through photosynthesis because of an almost equally large opposing flux of CO₂ from the ecosystem. Most of the return flux to the atmosphere is through soil respiration, which has two major sources, one heterotrophic (organisms decomposing organic matter) and one autotrophic (roots, mycorrhizal fungi and other root-associated microbes dependent on recent photosynthate). We used tree-girdling to stop the flow of photosynthate to the belowground system, hence, blocking autotrophic soil activity in a 120-yr-old boreal Picea abies forest. We found that at the end of the summer, two months after girdling, the treatment had reduced soil respiration by up to 53%. This figure adds to a growing body of evidence indicating (t-test, d.f. = 7, p < 0.05) that autotrophic respiration may contribute more to total soil respiration in boreal (mean 53 ± 2%) as compared to temperate forests (mean 44 ± 3%). Our data also suggests that there is a seasonal hysteresis in the response of total soil respiration to changes in temperature. We propose that this reflects seasonality in the tree below-ground carbon allocation.