Effects of enhanced nitrogen deposition on foliar chemistry and physiological processes of forest trees at the Bear Brook Watershed in Maine

Effects of enhanced nitrogen deposition on nutrient foliar concentrations and net photosynthesis of sugar maple (Acer saccharum Marsh.), American beech (Fagus grandifolia Ehrh) and red spruce (Picea rubens Sarg.) were evaluated at the Bear Brook Watershed in Maine (BBWM). The BBWM is a paired-watershed forest ecosystem study with one watershed treated since 1989 with bimonthly dry ammonium sulfate ((NH₄)₂SO₄) additions at a rate of 25.2 kg N ha⁻¹ year⁻¹, while the other watershed serves as a reference. The (NH₄)₂SO₄ treatment resulted in significant increases in foliar N concentrations for all three species and significant reductions in foliar Ca, Mg and Zn concentrations for American beech and red spruce. Treatment effects on foliar concentrations of other nutrients were not significant in any species. Despite higher N concentrations in all species, only treated sugar maple showed significantly higher photosynthetic rates. The non-response in net photosynthesis to higher foliar N in American beech and red spruce might be attributed to their low foliar Ca and/or Mg concentrations. Higher net photosynthetic rates in sugar maple might be explained by the higher foliar N and by the ability of this species to maintain an adequate Ca and Mg supply. Results suggested that nutrient imbalances due to inadequate supply of Ca and Mg might have counteracted a potential increase in net photosynthesis induced by higher N concentrations in American beech and red spruce.     

Fine-root distribution and morphology in an acidic Norway spruce (Picea abies (L.) Karst.) stand in SW Sweden in relation to granulated wood ash application

Wood ash is recommended as a compensatory fertiliser to counteract the effects of acidic deposition on forest ecosystems. Spatial distribution of biomass, necromass and morphology parameters of the fine roots (diameter classes <1, 1–2, <2 mm) of Norway spruce (Picea abies (L.) Karst.) were analysed in response to fertilisation with granulated wood ash (GWA) in a long-term field experiment in SW Sweden. GWA was applied as a single dose of 3200 kg ha⁻¹ and the fine roots were sampled 9 years later by soil coring. Soil cores were divided into 1-cm strata within the top 0–2.5 cm humus limits, the lower humus below 2.5 cm (with varying thickness) and the mineral soil to 50 cm depth (from ground surface). Total fine-root biomass in the control (C) and GWA treatment, 256 ± 20 and 258 ± 25 g m⁻², respectively, and length 2072 ± 182 and 1800 ± 198 m m⁻², respectively, did not differ statistically from each other. Total fine-root necromass in the 1–2 mm fraction was significantly higher in C than in the GWA treatment, 130 ± 12 and 80 ± 10 g m⁻², respectively. Fine-root biomass in the <1 mm fraction was significantly greater in the lower humus in the GWA treatment, but this did not affect the total biomass in the <1 mm fraction in the whole soil profile. The biomass-to-necromass ratio (1–2 mm) was significantly higher in the GWA treatment in the 0–30 cm soil layer than in the corresponding layer of the control. Specific root length (SRL) was lower in the GWA treatment than in the control in the 0–5 cm soil layer. The lower necromass and SRL were more clearly related to the GWA treatment, whereas the difference in the vertical distribution of biomass may have been related to the thicker humus layer in the GWA plots.     

Effects of experimental acidification,nitrogen addition and liming on ground vegetation in a mature stand of Norway spruce (Picea abies (L.) Karst.) in SE Sweden

During the period 1976–1991, a combined experiment of acidification, liming and nitrogen addition in a mature spruce stand was conducted at Farabol in south-east Sweden. The aim of this study was to investigate the effects of these treatments on the ground vegetation 0, 1, 5 and 15 years after experimental establishment. The treatment regimes were nitrogen (200 kg N ha⁻¹, repeated three times at 4–5-year intervals, totally 600 kg N ha⁻¹), sulphur powder (50 and 100 kg S ha⁻¹ a⁻¹, totally 600 and 1200 kg ha⁻¹), sulphur plus nitrogen (600+600 kg ha⁻¹) and limestone (500 kg ha⁻¹ a⁻¹, i.e. totally 6000 kg ha⁻¹). The results showed that nitrogen addition and liming promoted the abundance of the grass Deschampsia flexuosa, while acidification had a negative effect on D. flexuosa and herbs in the field layer. There was a negative reaction giving immediate damage to the bryophytes in connection with additions of nitrogen, sulphur powder and lime. The magnitude of damage and the capacity to recover varied among species as well as among treatments. The recovery from immediate damage after liming was much faster than after the treatments with sulphur powder and/or nitrogen. A negative interaction between sulphur powder and nitrogen was found for herbs and mosses where the combined effects were stronger than the effects of a single treatment alone. Acidification also had a negative effect on the total number of species. The results of this study showed that acidification and nitrogen deposition could negatively influence forest vegetation by changing the nutrient availability in the soils. Liming led to an improved growth of the forest ground vegetation and the flora changed towards a more nitrophilic species composition.     

Regeneration of Norway spruce in canopy gaps in Sphagnum-Myrtillus old-growth forests

Gap-associated spruce (Picea abies (L.) Karst.) regeneration in Sphagnum-Myrtillus stands of south taiga forests (Central Forest Biosphere reserve, Tver region, Russia) was studied to evaluate the role of different disturbances in spruce dynamics. Sampled gaps (n=70) ranged from 40 m² to 1.7 ha in size, and from 1 to 70 years since disturbance moment. Formation of gaps lead to increase in the number of stems per ha in all gap size classes (small: 40–200 m², medium: 200–3000 m², and large: >3000 m² gaps). Spruce was the most important species in gap refilling, although its role was not the same in different gap classes. The highest values of relative abundance (compared to other species) were recorded in small gaps, and much lower values – in middle and large gaps. However, as refilling of gaps proceeded, spruce showed rather active regeneration in middle and large gaps and partly regained its abundance in middle-age disturbances. In general, all types of gaps studied supported spruce regeneration into the forest canopy. Almost perfect correlation between predicted outcome of spruce dynamics in gaps and its current role in the canopy of Sphagnum-Myrtillus stands suggests a good adaptation of this species to the current disturbance regime and a steady state of the these forests.     

Stand development after different thinnings in two uneven-aged Picea abies forests in Sweden

Two field experiments, located in Central and Northern Sweden, were used to study the influence of standing volume on volume increment and ingrowth in uneven-aged Norway spruce (Picea abies (L.) Karst.) stands subjected to different thinnings. Each experiment had a 3 × 2 factorial block design with two replications. Treatments were thinning grade, removing about 45, 65, and 85% of pre-thinning basal area, and thinning type, removing the larger or the smaller trees, respectively. Each site also had two untreated control plots. Plot size was 0.25 ha. Volume increment was 0.5–6.8 m³ ha⁻¹ year⁻¹ for the plots, and significantly positively (p < 0.01) correlated with standing volume. Within treatment pairs, plots thinned from Above had consistently higher volume increment than plots thinned from Below. Ingrowth ranged from 3 to 33 stems ha⁻¹ year⁻¹, with an average of 14 and 21 stems ha⁻¹ year⁻¹ at the northern and southern site, respectively. At the southern site ingrowth was significantly negatively (p < 0.01) correlated with standing volume, but not at the northern site. Mean annual mortality after thinning was 2 and 7 stems ha⁻¹ year⁻¹at the northern and southern site, respectively.     

Influence of slope position, stand type and rhododendron (Rhododendron ponticum) on litter decomposition rates of Oriental beech (Fagus orientalis Lipsky.) and spruce [Picea orientalis (L.) Link]

Oriental beech (Fagus orientalis Lipsky.) and Oriental spruce [Picea orientalis (L.) Link] are the two most common tree species in northeast Turkey. Their distribution, stand type and understorey species are known to be influenced by topographical landforms. However, little information is available as to how these changes affect litter decomposition rates of these two species. Here, we investigated the effects of slope positions (top 1,800 m, middle 1,500 m and bottom 1,200 m), stand type (pure and mixed stands) and purple-flowered rhododendron (Rhododendron ponticum) on litter decomposition rates of Oriental beech and spruce for 4 years using the litterbag technique in the field. Among these three factors, stand type had the strongest influence on litter decomposition (P < 0.001, F = 58.8), followed by rhododendron (P < 0.001, F = 46.8) and slope position (P < 0.05, F = 11.6). Litter decomposition was highest under mixed beech/spruce forest, followed by pure beech and spruce forest. Beech and spruce litter decomposed much faster in mixed bags (beech–spruce) than they did separately under each stand type. Purple-flowered rhododendron significantly reduced litter decomposition of Oriental beech and spruce. Beech and spruce litter decomposed much slower at top slope position than at either bottom or middle position. Differential litter decomposition of Oriental beech and spruce was mainly due to adverse conditions in spruce forest and the presence of rhododendron on the ground which was associated with lower soil pH. Higher elevations (top slope position) slowed down litter decomposition by changing environmental conditions, most probably by decreasing temperature as also other factors are different (pH, precipitation) and no detailed investigations were made to differentiate these factors. The adverse conditions for litter decomposition in spruce forest can be effectively counteracted by admixture of beech to spruce monoculture and by using the clear-cutting method for controlling rhododendron.     

Nitrogen budgets for Scots pine and Norway spruce ecosystems 12 and 7 years after the end of long-term fertilisation

The magnitude of nitrogen storage and its temporal change in forest ecosystems are important when analysing global change. For example, the accelerated growth of European forests has been linked to increased nitrogen deposition, but the changes in the N inputs that cause long-term changes in ecosystems have not yet been identified. We used two Swedish forest optimum nutrition experiments with Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) to study the long-term fate of N applied to these forest ecosystems. In the pine experiment, in addition to fertiliser (NPK) application, soil acidity was manipulated by application of lime and dilute sulphuric acid. From the spruce experiment, we selected treatments with similar fertiliser doses as in the pine experiment and with and without lime addition.We quantified various terms in the N budget 12 years (pine) and 7 years (spruce) after the last N addition. In the pine stand the NPK-treatment was the only treatment to produce a significant increase in N in the tree biomass (97% above control), whereas in the spruce stand the N additions increased tree N in all treatment combinations (207% above control). In the pine stand the relative distribution of nitrogen between trees and soil did not vary across treatments, with trees containing around 12% of ecosystem N and humus containing around 44% of soil N. The increases in N stocks in the pine stands were mainly in the soil. In contrast, in the spruce ecosystem trees accumulated most of the added N and the increase in the soil was restricted to the humus layer.In the pine ecosystem, large losses of added N (between 254 and 738 kg ha⁻¹ out of 1040 kg ha⁻¹ added as fertiliser) occurred, whereas in the spruce ecosystem we recovered more N than could be accounted for by inputs (between 250 and 591 kg ha⁻¹). There was no clear pattern in the interaction between acidification/liming and N additions.     

Water and element fluxes during the regeneration of Norway spruce with European beech: Effects of shelterwood-cut and clear-cut

The regeneration of mature Norway spruce with European beech using the shelterwood silvicultural system is a good example of continuous cover forestry. In contrast, the regeneration may also start with clear-cut plots, which often occur after calamities like wind-throw or bark beetle attack. During regeneration the forest ecosystem becomes a highly dynamic system. Nutrient losses from the soil may occur as the element turnover is affected by the reduced nutrient uptake of forest trees as well as the enhanced mineralisation and nitrification due to higher soil temperature and soil moisture. Continuous cover forestry may help to reduce these nutrient losses. In order to test this, we investigated water and element fluxes of two chronosequences. The first investigated regeneration in the shelterwood system, while the second concerned itself with regeneration on clear-cut plots. In a shelterwood-cut about 30% of the mature spruce trees are removed and young beech trees are planted. Some 10 years later a secondary felling is carried out and at age 20 of the beech regeneration the final harvest of the mature trees occurs. Thus, the studied time steps were (a) the first 5 years after the initial cut and planting, (b) 10-year-old beech regeneration after the second shelterwood cut and (c) 20-year-old beech regeneration after the final harvest.Our results indicate that nutrient losses with seepage water – especially nitrogen, calcium and magnesium – occur during the first years after the clear cut and, to a lesser extent, after secondary felling on the selective-cut plot. This may temporarily affect seepage water quality due to elevated nitrate concentrations, which reached values of more than 100 mg l⁻¹. In the time span between planting and an age 20 of the beech regeneration, total losses of nitrogen from the main rooting zone reach 230 kg ha⁻¹ after clear cut. Preliminary estimates of the total nitrogen loss in the shelterwood system range between 150 and 230 kg ha⁻¹ indicating either significantly lower or equal losses of nutrients. In the second case, however, element output is distributed more equally over the 20-year-period than after clear felling where 85% of the nitrate leaching occurs during the first 3 years.     

Wood volume yield and stand structure in Norway spruce understorey depending on birch shelterwood density

Wood volume yield and stand structure were investigated for Norway spruce understorey growing at 1500 trees ha⁻¹ under birch shelters of two different densities, 300 and 600 trees ha⁻¹, and Norway spruce growing without shelter, in a field trial in the boreal coniferous forest, 56 years after the establishment of the stand and 19 years after establishment of the trial.Wood volume yield in sheltered spruce (mean annual increments of 1.87 and 1.78 m³ ha⁻¹ year⁻¹ under the dense and sparse shelterwoods, respectively) was significantly lower than that of unsheltered spruce (mean annual increment 2.43 m³ ha⁻¹ year⁻¹). The loss in wood volume yield for sheltered spruce was more than compensated for by the additional wood volume yield in the shelterwoods (mean annual increments 3.26 and 1.88 m³ ha⁻¹ year⁻¹ for the dense and sparse shelterwood respectively).Shelterwood density did not produce any significant differences in inequality of the understorey stands, measured as skewness and the Gini coefficient for the wood volume distributions. This implies that two-sided competition for nutrients and water was more significant than competition for light.Immediately after trial establishment, trees in the no shelterwood treatment (i.e. where all overstory trees had been removed) showed a marked increase in diameter growth. Over time, the growth rate of unsheltered Norway spruce was reduced to a level comparable to that of sheltered spruce. The difference in average diameter has persisted during the trial period. There was no similar effect on height growth, resulting in an increased slenderness index (h/d) with increased shelterwood density for the understorey trees.     

Spruce beetles and forest ecosystems in south-central Alaska: A review of 30 years of research

From 1920 to 1989, approximately 847,000 ha of Alaska spruce (Picea spp.) forests were infested by spruce beetles (Dendroctonus rufipennis). From 1990 to 2000, an extensive outbreak of spruce beetles caused mortality of spruce across 1.19 million ha of forests in Alaska; approximately 40% more forest area than was infested the previous 70 years. This review presents some of the most important findings from a diversity of research and management projects from 1970 to 2004 to understand the biology, ecology, and control of this important forest insect, and the causes and effects of their outbreaks. We suggest that future research should examine the long-term effects of the spruce beetle outbreaks and climate variability on forest ecosystems in the region. Research into how different management actions facilitate or interrupt natural successional processes would be particularly useful.     

Estrogenic and anti-estrogenic activities of hispolon from Phellinus lonicerinus (Bond.) Bond. et sing

Hispolon was the main antitumor active ingredient in Phellinus sensu lato species. In order to confirm the dual regulating estrogenic ingredient and obtain more effective natural estrogen replacement drugs, hispolon was separated from Phellinus lonicerinus (Bond.) Bond. et sing. Hispolon exhibited significant anti-proliferative effect against estrogen-sensitive ER (+) MCF-7 cells in the absence of estrogen, and exhibits antagonistic effects on 17β-estradiol (E₂)-induced MCF-7 cell proliferation when E₂ and the different concentrations of hispolon were treated simultaneously. Hispolon also inhibited the proliferation of estrogen-negative ER (−) MDA-MB-231 cells at the concentration of 5.00 × 10^− 5 M. The yeast two-hybrid experiments showed that hispolon had strong and non-selective effects on the estrogen receptor (ER) α and ERβ at a concentration of 1.00 × 10^− 6 M. The ERβ-binding ability of hispolon was larger than ERα in the concentration range of 1.00 × 10^− 9 M and 1.00 × 10^− 7 M. Hispolon could increase the body weight coefficient, serum E₂ and progesterone contents in immature female mice at dose of 9.10 × 10^− 6 mol/kg, and increase coefficient of thymus and spleen in mice. The Gscores of hispolon-ERα and hispolon-ERβ docked complexes were − 7.93 kcal/mol and − 7.79 kcal/mol in docking simulations. Hispolon presented dual regulating estrogenic activities, which showed estrogenic agonist activity at low concentration or lack of endogenous estrogen, and the estrogenic antagonistic effect was stimulated at high concentrations or too much endogenous estrogen. Hispolon could be used for treating the estrogen deficiency-related disease with the benefit of non-toxic to normal cells, good antitumor effects and estrogenic activity.     

Production,allocation, and stemwood growth efficiency of Pinus taeda L. stands in response to 6 years of intensive management

Loblolly pine (Pinus taeda L.) is a highly plastic species with respect to growth responses to forest management. Loblolly pine is the most planted species across the southern United States, a region with the most expansive and intensively managed forest plantations in the world. Management intensity, using tools such as site preparation and fertilization, is increasing greatly in scope over time. To better define to the productive potential of loblolly pine under intensive management, the influence of 6 years of management with weed control (W), weed control plus irrigation (WI), weed control plus irrigation and fertigation (irrigation with a fertilizer solution) (WIF), or weed control plus irrigation, fertigation, and pest control (WIFP) since plantation establishment on stand productivity in loblolly pine was examined. The site is located near Bainbridge, GA (30°48′N latitude and 84°39′W longitude) and is of medium quality (site index=18 m, base age 25). Increasing management intensity greatly accelerated stand development and biomass accumulation. At age 6 total production (above plus belowground) was nearly doubled from 50 to 93 Mg ha⁻¹ in WIFP stands compared to W stands, and standing stem biomass increased from 24 Mg ha⁻¹ in W stands to 48 Mg ha⁻¹ in response to WIFP treatment. Stem current annual increment (CAI) peaked at age 5 in the WIF and WIFP stands at 17–18 Mg ha⁻¹ per year at a basal area between 18 and 21 m² ha⁻¹. Year to year variation in CAI was better explained by previous-year leaf area index (LAI) than current-year LAI. Maximum stemwood production in loblolly pine was achieved through large increases in LAI and small decreases in allocation to woody roots (tap+coarse roots) versus woody shoots (stem+branches) associated with intensive treatments.     

Dead trees and protected polypores in unmanaged north-temperate forest stands of Lithuania

The availability of coarse woody debris (CWD) and distribution of dead trees into categories of mortality (dead standing, broken and uprooted) were investigated in north-temperate forests of central Europe (Lithuania). The studied area comprised 188.7 ha and included 18 different stands 40–130 years of age with a variety of tree species (spruce (Picea abies (L.) Karst.), pine (Pinus sylvestris L.), alder (Alnus glutinosa (L.) Gaertn.), birch (Betula pendula Roth and B. pubescens Ehrh.), aspen (Populus tremula L.), oak (Quercus robur L.), forest types (caricus-sphagnum, vaccinium-myrtillus, oxalis, myrtillus-oxalis, caricus-calamagrostis) and edaphic conditions (peaty, sandy, loamy soils of different moisture). The stands were excluded from wood harvesting for at least 30 years. A total of 11 365 dead trees (over 10 cm in DBH) or 6160.7 m³ of dead wood was found (60.2 trees/ha and 32.6 m³/ha). The volume of CWD per hectare was larger in older stands (r_S=0.78, P<0.01). Tree mortality during the last 2 years consisted of 482 trees and 381 m³, or 1.28 trees/ha×year and 1.01 m³/ha×year. In 25–33% of cases it was wind-related. Uprooted and broken trees were of larger DBH than dead standing. The distribution into the categories of mortality was strongly dependent on tree species (chi-square test, d.f.=10,P=0). Dead standing dominated in CWD of pine and alder. Broken trees comprised almost a half in CWD of aspen, and about one-third in birch, alder and oak. Uprooting most often occurred in spruce, aspen and birch. Edaphic conditions and stand age had a pronounced impact on distribution into mortality categories for spruce (chi-square test, d.f.=20, P<0.00001) and pine (d.f.=8, P≤0.0003). On peat soil, only a minority of trees of both pine and spruce was uprooted, and standing dead prevailed. In CWD of spruce and pine, the proportions of both dead standing and broken decreased and that of uprooted trees increased on mineral soils of higher moisture and bulk density in older stands. By contrast, uprooting in birch and alder occurred less often on more wet sites, where the proportions of standing snags were higher. A total of 41 species of wood-decomposing polypores were found in the study area. Among those, 10 (24%) were of conservation value.     

Stand and landscape level effects of a major outbreak of spruce beetles on forest vegetation in the Copper River Basin,Alaska

From 1989 to 2003, a widespread outbreak of spruce beetles (Dendroctonus rufipennis) in the Copper River Basin, Alaska, infested over 275,000 ha of forests in the region. During 1997 and 1998, we measured forest vegetation structure and composition on one hundred and thirty-six 20-m × 20-m plots to assess both the immediate stand and landscape level effects of the spruce beetle infestation. A photo-interpreted vegetation and infestation map was produced using color-infrared aerial photography at a scale of 1:40,000. We used linear regression to quantify the effects of the outbreak on forest structure and composition. White spruce (Picea glauca) canopy cover and basal area of medium-to-large trees [≥15 cm diameter-at-breast height (1.3 m, dbh)] were reduced linearly as the number of trees attacked by spruce beetles increased. Black spruce (Picea mariana) and small diameter white spruce (<15 cm dbh) were infrequently attacked and killed by spruce beetles. This selective attack of mature white spruce reduced structural complexity of stands to earlier stages of succession and caused mixed tree species stands to lose their white spruce and become more homogeneous in overstory composition. Using the resulting regressions, we developed a transition matrix to describe changes in vegetation types under varying levels of spruce beetle infestations, and applied the model to the vegetation map. Prior to the outbreak, our study area was composed primarily of stands of mixed white and black spruce (29% of area) and pure white spruce (25%). However, the selective attack on white spruce caused many of these stands to transition to black spruce dominated stands (73% increase in area) or shrublands (26% increase in area). The post-infestation landscape was thereby composed of more even distributions of shrubland and white, black, and mixed spruce communities (17–22% of study area). Changes in the cover and composition of understory vegetation were less evident in this study. However, stands with the highest mortality due to spruce beetles had the lowest densities of white spruce seedlings suggesting a longer forest regeneration time without an increase in seedling germination, growth, or survival.     

The impact of six European tree species on the chemistry of mineral topsoil in forest plantations on former agricultural land

Influences on mineral topsoils of common European tree species (oak-Quercus robur L., lime-Tilia cordata Mill., ash-Fraxinus excelsior L., birch-Betula pendula Roth., beech-Fagus sylvatica L. and spruce-Picea abies (L.) Karst.) were studied in 30 to 40-year-old stands planted in adjacent plots on former arable land. Mineral soil samples from two depth layers (0–10 and 20–30 cm) under the different species were compared in terms of pH, base saturation, pools and concentrations of exchangeable macro- and micronutrients, total nitrogen and carbon. With the exception of pH (H₂O) and extractable Al and Fe, no significant differences between species were detected in the lower layer. The upper (0–10 cm) layer was, however, affected differently depending on tree species: significant differences in pH, base saturation, exchangeable base cations and other nutrients were observed. The most prominent differences were between lime and spruce. Lime had considerably higher pH, base saturation, base cation and boron pools compared to spruce, which had the most acidifying effect on the mineral topsoils. Among the deciduous species, beech had the most similar effect to spruce on the upper layer of mineral topsoils. Soil C, N and C/N ratios did not differ significantly among species.     

Carbon sources and sinks in high-elevation spruce–fir forests of the Southeastern US

This paper examines carbon (C) pools, fluxes, and net ecosystem balance for a high-elevation red spruce–Fraser fir forest [Picea rubens Sarg./Abies fraseri (Pursh.) Poir.] in the Great Smoky Mountains National Park (GSMNP), based on measurements in fifty-four 20 m × 20 m permanent plots located between 1525 and 1970 m elevation. Forest floor and mineral soil C was determined from destructive sampling of the O horizon and incremental soil cores (to a depth of 50 cm) in each plot. Overstory C pools and net C sequestration in live trees was estimated from periodic inventories between 1993 and 2003. The CO₂ release from standing and downed wood was based on biomass and C concentration estimates and published decomposition constants by decay class and species. Soil respiration was measured in situ between 2002 and 2004 in a subset of eight plots along an elevation gradient. Litterfall was collected from a total of 16 plots over a 2–5-year period.The forest contained on average 403 Mg C ha⁻¹, almost half of which stored belowground. Live trees, predominantly spruce, represented a large but highly variable C pool (mean: 126 Mg C ha⁻¹, CV = 39%); while dead wood (61 Mg C ha⁻¹), mostly fir, accounted for as much as 15% of total ecosystem C. The 10-year mean C sequestration in living trees was 2700 kg C ha⁻¹ year⁻¹, but increased from 2180 kg C ha⁻¹ year⁻¹ in 1993–1998 to 3110 kg C ha⁻¹ year⁻¹ in 1998–2003, especially at higher elevations. Dead wood also increased during that period, releasing on average 1600 kg C ha⁻¹ year⁻¹. Estimated net soil C efflux ranged between 1000 and 1450 kg C ha⁻¹ year⁻¹, depending on the calculation of total belowground C allocation. Based on current flux estimates, this old-growth system was close to C neutral.     

Significant effects of temperature on the reproductive output of the forest herb Anemone nemorosa L.

Climate warming is already influencing plant migration in different parts of the world. Numerous models have been developed to forecast future plant distributions. Few studies, however, have investigated the potential effect of warming on the reproductive output of plants. Understorey forest herbs in particular, have received little attention in the debate on climate change impacts.This study focuses on the effect of temperature on sexual reproductive output (number of seeds, seed mass, germination percentage and seedling mass) of Anemone nemorosa L., a model species for slow colonizing herbaceous forest plants. We sampled seeds of A. nemorosa in populations along a 2400 km latitudinal gradient from northern France to northern Sweden during three growing seasons (2005, 2006 and 2008). This study design allowed us to isolate the effects of accumulated temperature (Growing Degree Hours; GDH) from latitude and the local abiotic and biotic environment. Germination and seed sowing trials were performed in incubators, a greenhouse and under field conditions in a forest. Finally, we disentangled correlations between the different reproductive traits of A. nemorosa along the latitudinal gradient.We found a clear positive relationship between accumulated temperature and seed and seedling traits: reproductive output of A. nemorosa improved with increasing GDH along the latitudinal gradient. Seed mass and seedling mass, for instance, increased by 9.7% and 10.4%, respectively, for every 1000 °C h increase in GDH. We also derived strong correlations between several seed and seedling traits both under field conditions and in incubators. Our results indicate that seed mass, incubator-based germination percentage (Germ%_Inc) and the output of germinable seeds (product of number of seeds and Germ%_Inc divided by 100) from plants grown along a latitudinal gradient (i.e. at different temperature regimes) provide valuable proxies to parameterize key population processes in models.We conclude that (1) climate warming may have a pronounced positive impact on sexual reproduction of A. nemorosa and (2) climate models forecasting plant distributions would benefit from including the temperature sensitivity of key seed traits and population processes.     

Long-term trends in loblolly pine productivity and stand characteristics in response to thinning and fertilization in the West Gulf region

Two levels each of thinning and fertilization were applied to a 7-year-old loblolly pine (Pinus taeda L.) plantation on a nitrogen- and phosphorus-deficient West Gulf Coastal Plain site in Louisiana. Levels of thinning were no thinning, or thinning applied 7 and 14 years after stand initiation. Levels of fertilization were no fertilization or broadcast fertilization with diammonium phosphate at age 7 years plus refertilization with urea, monocalcium phosphate, and potash at age 14 years. Long-term measurements of climate, stand development and productivity, projected leaf area index, and foliar nutrition were initiated at age 11 years. We found that by age 17 years, thinning increased mean live-crown length from 4.2 to 7.8 m, and mean tree diameter from 15.0 to 21.8 cm compared to the unthinned treatment. After rethinning at age 14 years, stand basal area increased 1.2 and 19.2% between ages 15 and 17 years on the unthinned and thinned plots, respectively. Refertilization at age 14 years reestablished foliar N, P and K sufficiency, which increased leaf area index from 4.2 to 6.0 m² m⁻² on the unthinned plots and from 3.2 to 3.8 m² m⁻² on the thinned plots, and subsequently, increased gross stand biomass from 114 to 141 Mg ha⁻¹ on the unthinned plots and from 78 to 95 Mg ha⁻¹ on the thinned plots by age 17 years. Leaf area was an important factor controlling loblolly pine productivity. At our study site, however, competition for light and water and nutrition-limited foliage growth influenced the variability and scope of this relationship. Our results suggest that a positive and linear relationship between leaf area and loblolly pine productivity does not universally occur on loblolly pine sites.     

Productivity of mixed versus pure stands of oak (Quercus petraea (Matt.) Liebl. and Quercus robur L.) and European beech (Fagus sylvatica L.) along an ecological gradient

The mixture of beech (Fagus sylvatica L.) and oak (sessile oak, Quercus petraea (Matt.) Liebl., and pedunculate oak, Q. robur L.) is of considerable importance in Europe and will probably become even more important under climate change. Therefore, the performance of oak and beech in mixture was compared with the species’ growth in pure stands. Data from 37 long-term mixing experiments in Poland, Germany and Switzerland were pooled for analysis of mixing effects on stand productivity and possible interrelationships with mixing portions or site conditions. We found that on average, mixed stands of oak and beech exceeded biomass productivity in pure stands by 30 % or 1.7 t ha^?1 year^?1, as the growth of both species was benefitted by the mixture. However, that the interaction actually ranged from facilitation and overyielding on poor sites to underyielding on fertile sites triggered by competition. An empirically derived interaction model showed volume and dry mass growth changing in mixed stands from gains of 50 % to losses of 10 % depending on site conditions. It is concluded that the analysed mixture grows in accordance with the stress-gradient hypothesis and that our results suggest a site-specific relationship between species mixture and biomass productivity. As a consequence, an adequate species mix should result in increased productivity under steady state as well as climate change.