Instantaneous and potential radiation effect on underplanted European beech below Norway spruce canopy

In even-aged, 120-year-old Norway spruce stands with underplanted beech (in 1995) four permanent research plots were established, each 100 × 100 m in size. Twelve subplots were selected on each plot along a light gradient from complete canopy closure to open light conditions. On each plot, photon flux density (PFD) was measured continuously, and during same time interval (August 2008, 2009), color digital hemispherical photographs were taken. Cumulative PFD values for measuring points were compared with potential PFD radiation obtained from hemispherical photograph analysis for different angle of hemisphere and different parameters from hemispherical image analysis (gap fraction, total openness and direct, indirect and total amount of radiation). Cumulative and average daily values for the plots were compared; 120° hemispherical photograph angle, gap fraction and total openness were the variables that explained the largest proportion of variance in light transmittance. Determination coefficients between direct and total light component were highest for the total potential radiation and lowest for the diffuse light component. Comparison between potential and instantaneous light measurements for radial, height increment and SLA of young beech showed that instantaneous radiation measurements explained height increment best; 120° of hemisphere proved to be the best explaining angle.     

Competition modifies effects of enhanced ozone/carbon dioxide concentrations on carbohydrate and biomass accumulation in juvenile Norway spruce and European beech

Elevated concentrations of carbon dioxide ([CO2]) and ozone ([O3]) affect primary metabolism of trees in opposite ways. We studied their potential interactions on carbohydrate concentrations and contents. Two hypotheses currently under debate were tested. (1) Stimulation of primary metabolism by prolonged exposure to elevated [CO2] does not compensate for the adverse effects of O3 on carbohydrate accumulation and biomass partitioning to the root. (2) Growth in a mixed-species planting will repress plant responses to elevated [O3] and [CO2] relative to conditions in a monoculture. To this end, European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst.) saplings grown under conditions of intra- and interspecific competition were pre-acclimated for 1 year to ambient or elevated [CO2]. In the following 2-year phytotron study, trees were exposed to factorial combinations of ambient and elevated [O3] and [CO2]. The total carbohydrate content (sugar and starch) of spruce was greater in plants exposed to elevated [CO2] than in plants exposed to ambient [CO2]. In beech, the opposite response was observed, especially when this species was grown in combination with spruce. Overall, the data did not support Hypothesis 1, because the adverse effects of O3 were counteracted by elevated [CO2]. Support for Hypothesis 2 was species-dependent. In beech saplings, reduction of carbohydrates by elevated [O3] and stimulation by elevated [CO2] were repressed by competitive interaction with spruce. In contrast, in spruce, stimulation of carbohydrates by elevated [CO2] was similar in mono- and mixed cultures. Thus Hypothesis 2 was supported for beech but not spruce. We conclude that, in juvenile beech and spruce, a 3-year exposure to elevated [CO2] counteracts the adverse effects of O3 on carbohydrate concentrations and contents. For beech, sensitivity to elevated [CO2] and [O3] was high in monoculture but was largely repressed by interspecific competition with spruce. In contrast, the response of spruce to perturbations of atmospheric chemistry was not significantly affected by either intra- or interspecific competition.     

Effects of shade on growth and mortality of maple (Acer pseudoplatanus), ash (Fraxinus excelsior) and beech (Fagus sylvatica) saplings

We compared shade tolerance of maple, ash and beech in the saplingstage from two sites with rich soils differing in water supply,growing in dense thickets underneath a beech shelterwood ofvarying canopy densities. Shade tolerance was described by twocomponents: mortality in shade and height growth in high light.At low light, beech showed the least mortality, maple the highestand ash in between on both sites. The decline with increasinglight was steepest in beech and more gradual with ash and maple.At 15 per cent above canopy light, all three species approachedzero mortality. Beech as the most shade-tolerant species hadthe highest survival rate under low light and the least lengthgrowth rate under high light (>17 per cent). Ash had a lowersurvival rate at low light than beech and a highest growth rateat high light. Maple showed a bit weaker trade-off with thelowest survival rate but a growth rate inferior to ash. On thebetter water-supplied site, height growth was significantlysuperior in all three species only under high light. On thebasis of these results, silvicultural conclusions are drawnwith respect to appropriate light levels and cutting types.     

Effects of soil temperature on shoot and root growth and nutrient uptake of 5-year-old Norway spruce seedlings

Soil temperature is a main factor limiting root growth in the boreal forest. To simulate the possible soil-warming effect of future climate change, 5-year-old Norway spruce (Picea abies (L.) Karst.) seedlings were subjected to three simulated growing seasons in controlled environment rooms. The seedlings were acclimated to a soil temperature of 16 degrees C during the first (GS I) and third growing seasons (GS III), but were assigned to random soil-temperature treatments of 9, 13, 18 and 21 degrees C during the second growing season (GS II). In GS II, shoot diameter growth was lowest in the 21 degrees C treatment and root growth was lowest in the 9 degrees C treatment. In GS III, shoot height and root length growth improved in seedlings that had been kept at 9 degrees C during GS II, indicating compensatory growth in response to increased soil temperature. The temporary decrease in soil temperature had no long-lasting significant effect on seedling biomass or total nutrient uptake. At the end of GS III, fine roots of seedlings exposed to a soil temperature of 21 degrees C in GS II were distributed more evenly between the organic and mineral soil layers than roots of seedlings in the other treatments. During GS II and GS III, root growth started earlier than shoot growth, decreased during the rapid shoot elongation phase and increased again as shoot growth decreased.     

Effects of simulated acid deposition on the surface structure of Norway spruce and silver fir needles

Treatments with acidified solutions of pH 3.5 cause alterations in the wax structure of the needles of silver fir and Norway spruce. The wax structures, especially those of the stomata, once they have been damaged, are not capable of complete reorganization even if the needle is repeatedly treated with distilled water.     

Storm-induced tree resistance and chemical differences in Norway spruce (Picea abies)

Introduction   

Windstorm is one of the most destructive environmental disturbance factors on forests, but its influence on conifer defense chemistry and susceptibility to insects and diseases is not well understood.     

Impact of bark wounds on sapwood in Norway spruce and silver fir

Bark wounds by damage during harvesting are a serious problem in forestry due to fungi infection and wood deterioration. This paper presents results of an investigation about the influence of the wounds on the internal structure of such injured stems. In an experiment, bark wounds were artificially created at the stem base of Norway spruce [Picea abies (L.) Karst.] and silver fir (Abies alba Mill.). 2 years later, the injured stems along with undamaged controls were cut and the trunk portion below breast height subjected to computer tomographic (CT) analysis. Analysis of the CT-images revealed a substantial impact of wounding on sapwood properties in spruce: directly adjacent to the wound surface in all examined trees, a large disturbance zone was detected affecting on average 17% of the potential sapwood area. With increasing distance from the wound, the size of this disturbance zone diminished, but was still detectable in all trees at breast height ca. 1 m above the bark wound.     

Pine weevil (Hylobius abietis) damage to cuttings and seedlings of Norway spruce

Damage caused by pine weevil (Hylobius abietus L.) to planted seedlings and cuttings of Norway spruce (Picea abies (L.) Karst.) was studied at five clearcut sites in south-eastern Sweden. The main objective was to compare the two types of stock in terms of attack frequency and mortality due to pine weevil feeding. Cuttings and seedlings with the same initial stem-base diameter (4 mm) were compared. Two sites were harvested and scarified shortly before planting, two were harvested shortly before planting, but were not scarified, and one was harvested 2 years before and scarified the autumn before planting. The total mortality 5 years after planting was highest, greater than 90%, at the new, non-scarified sites, and lowest, 23%, at the old, scarified site. More than 90% of the mortality was caused by pine weevil feeding. Attack frequency and pine weevil induced mortality were significantly higher among seedlings than among cuttings. Mortality due to pine weevil damage was 4–43% higher in seedlings than in cuttings after the fifth year. Of the cuttings and seedlings that were attacked in the first year, a significantly higher frequency of the seedlings were girdled. The higher resistance of cuttings to pine weevil damage may partly explain the more rapid growth of cuttings reported in other studies. However, the causes of their higher resistance need to be further investigated. The thicker bark and needles on the stem base of the cuttings could be important in this respect.     

Nutrient cycling and soil leaching in eighteen pure and mixed stands of beech (Fagus sylvatica) and spruce (Picea abies)

Studies on the combined effects of beech–spruce mixtures are very rare. Hence, forest nutrition (soil, foliage) and nutrient fluxes via throughfall and soil solution were measured in adjacent stands of pure spruce, mixed spruce–beech and pure beech on three nutrient rich sites (Flysch) and three nutrient poor sites (Molasse) over a 2-year period. At low deposition rates (highest throughfall fluxes: 17 kg N ha⁻¹ year⁻¹ and 5 kg S ha⁻¹ year⁻¹) there was hardly any linkage between nutrient inputs and outputs. Element outputs were rather driven by internal N (mineralization, nitrification) and S (net mineralization of organic S compounds, desorption of historically deposited S) sources. Nitrate and sulfate seepage losses of spruce–beech mixtures were higher than expected from the corresponding single-species stands due to an unfavorable combination of spruce-similar soil solution concentrations coupled with beech-similar water fluxes on Flysch, while most processes on Molasse showed linear responses. Our data show that nutrient leaching through the soil is not simply a “wash through” but is mediated by a complex set of reactions within the plant–soil system.     

Effects of experimental drought on the fine root system of mature Norway spruce

Norway spruce (Picea abies (L.) Karst.) is an economically important, but relatively drought-sensitive tree species that might suffer from increasing drought intensities and frequencies, which are predicted to occur in parts of central Europe under future climatic change. In a throughfall exclusion experiment using sub-canopy roofs, we tested the hypothesis that enhanced drought leads to an increase in fine root mortality, and also to a higher, subsequent fine root growth. Fine root production and mortality were assessed using two independent approaches, sequential soil coring (organic layer) and direct observations in minirhizotrons (organic layer plus upper mineral soil). Six weeks of throughfall exclusion resulted in mild drought stress, i.e. a reduction in average soil moisture from 20 to 12 vol.% during the treatment. Based on the sequential coring data, experimental drought did not result in significant changes in fine root biomass during the 6-week treatment period, but caused an increase in fine root mortality by 61% in the 6 weeks following the drought treatment. Remarkably, fine root production showed a synchronous increase in this period, which more than compensated for the loss due to increased mortality. The minirhizotron data confirmed that the drought treatment increased fine root loss in the organic layer. Based on this method, however, root loss occurred during the drought period and was not compensated by increased root production. The mild drought stress was mainly restricted to the organic layer and did not significantly influence fine root dynamics in the mineral soil. We calculated that the drought event resulted in an extra input of about 28 g C m⁻² and 1.1 g N m⁻² to the soil due to increased fine root mortality. We conclude that even periods of mild drought significantly increase fine root mortality and the associated input of root-derived C to the soil organic matter pool in temperate Norway spruce forests.     

Response of mature stands of Norway spruce (Picea abies) to boron fertilization

The effects of boron (B) fertilizer applied 10 growing seasons earlier were studied in mature Norway spruce (Picea abies (L.) Karst.) trees in long-term factorial fertilization experiments at two field sites. Needle nutrient status, above-ground and below-ground growth and δ¹³C and carbon concentrations in the annual rings were measured. Needle B concentrations varied between 4 and 19 mg kg⁻¹ on the plots that had not received B fertilization. On the B-fertilized plots they varied between 15 and 39 mg kg⁻¹. The lowest B concentrations were on the plots that had received N or NCa fertilization. Needle Mn and Zn concentrations were lower on the B plots than on the plots that had not received B fertilization, although not significantly. Mean annual volume growth was slightly higher on the B plots at the more fertile site, but not at the less fertile one. The living:dead fine root mass ratio and living fine root length were also higher on the B-fertilized plots than on the unfertilized plots, but δ¹³C was not significantly affected, suggesting that the water status of the trees was not markedly altered by the increase in root growth. The carbon concentration in the annual rings was higher in the B-fertilized trees than in the unfertilized ones, suggesting the importance of B for wood formation.     

Effects of pre- and post-planting shading on growth of container Norway spruce seedlings

Norway spruce (Picea abies (L.) Karst.) is shade-tolerant and sensitive to high irradiance, summer frosts and winter desiccation, which can impair its reforestation success. In this study, artificial pre- and post-planting shading was examined to determine their effects on post-planting shoot and root growth as well as the vigor of one- and two-year-old Norway spruce seedlings. Three planting experiments were carried out on open nursery fields (Exp. 1, 2) and on a mounded forest clearcut in central Finland (Exp. 3). Before planting, the seedlings were stored over winter either in a freezer or on open fields under snow cover. For two weeks prior to planting, half of the seedlings were placed in the open and the other half under a horizontal shade netting (light transmittance 56 %) (Exp. 1, 2). All seedlings were planted with or without a vertical post-planting shade, which was located on the southern side. Post-planting shading enhanced shoot growth and reduced damage (better visual vigor and needle color and less pine-weevil damage) on Norway spruce seedlings for at least two years after planting (Exp. 2, 3). Those seedlings, that had been stored over winter in the open and kept in shade prior to planting seemed to benefit most from post-planting shading (Exp. 2). However, post-planting shading may give variable results, depending on the seedling quality and weather conditions after planting, and may even reduce shoot growth if no pre-planting shading is used (Exp. 1). Shoot growth may also be improved at the expense of root growth (Exp. 3). The costs of manufacturing and installing post-planting shades may limit their use in practice, for example, to selected regeneration sites where there is high risk of frost damage but where no alternative silvicultural procedure (shelterwood or nurse crop) has been used.     

The effect of nitrogen fertilization on fungistatic phenolic compounds in roots of beech (Fagus sylvatica) and Norway spruce (Picea abies)

The effect of nitrogen fertilization on fungistatic phenolic compounds in fine roots of beech and Norway spruce growing in afforestation plots was analysed. The plots were situated at two sites in Switzerland on acidic soil with low base saturation. For 9 years, the trees have been treated with dry ammonium nitrate to give 0, 10, 20, 40, 80, 160 kg N ha^?1 year^?1, respectively. The phenolic compounds responded differently to fertilization. Fine roots of beech showed a significant decrease of (?)‐epicatechin and piceatannol with increasing nitrogen fertilization. The concentration of protocatechuic acid was increased with fertilization. Roots of fertilized Norway spruce showed significantly decreased concentrations of 4‐hydroxyacetophenone and piceatannol. The mycelial growth of three isolates each of Heterobasidion annosum s.l. and Cylindrocarpon destructans was tested on agar media containing various phenolic compounds in concentrations found in fine roots of Norway spruce (Picea abies) and beech (Fagus sylvatica). All three H. annosum isolates were inhibited by p‐coumaric acid and (?)‐epicatechin. Two isolates were inhibited by another four phenolic compounds (p‐hydroxybenzoic acid, 4‐hydroxyacetophenone, piceatannol and protocatechuic acid), one by (+)‐catechin. Two of three C. destructans isolates were inhibited by all phenolic compounds except for (+)‐catechin which affected only one isolate, one isolate did not respond at all.     

Effects of sika deer (Cervus nippon yakushimae) population growth on saplings in an evergreen broad-leaved forest

An increase in size of sika deer populations has caused severe damage to forest vegetation in many regions of Japan. In a Natural World Heritage area on the western part of Yakushima Island, existing high densities of deer are thought to be increasing, but the impact of deer on forest vegetation has not been estimated. Roadside censuses of deer between 1988 and 2006 showed that encounter rates are continuing to increase. Long-term monitoring of individual tree saplings in a 0.13 ha area of the forest between 1992 and 2005 showed that many saplings still remain despite increases in sika deer density. However, differences in growth and mortality rates of preferred, neutral (neither preferred nor avoided) and avoided sika deer food species varied over time. As a group, preferred species continuously had a low growth rate, while their mortality rate increased through time. The growth rates of the neutral and avoided groups decreased especially in the second half of our study period at the same time that their mortality rates increased. These data suggest that in spite of food species preference, direct feeding pressures as well as indirect physical effects by increasing sika deer density are affecting growth and mortality of tree saplings. Clearly, it is difficult to predict how plant communities will respond to possible continued sika deer population growth, and it is therefore important to continue monitoring these forest communities in the interest of preserving the unique ecosystems on Yakushima Island.     

Influence of light availability on growth, leaf morphology and plant architecture of beech ( Fagus sylvatica L.), maple ( Acer pseudoplatanus L.) and ash ( Fraxinus excelsior L.) saplings

In a field study, we measured saplings of beech, ash and maple growing in a fairly even-aged mixed-species thicket established by natural regeneration beneath a patchy shelterwood canopy with 3–60% of above canopy radiation reaching the saplings. Under low light conditions, maple and ash showed a slight lead in recent annual length increment compared with beech. With increasing light, ash and maple constantly gained superiority in length increment, whereas beech approached an asymptotic value above 35% light. A suite of architectural and leaf morphological attributes indicated a more pronounced ability of beech to adapt to shade than ash and maple. Beech displayed its leaves along the entire tree height (with a concentration in the middle crown), yielding a higher live crown ratio than ash and maple. It allocated biomass preferentially to radial growth which resulted in low height to diameter ratios, and expressed marked plagiotropic growth in shade indicating a horizontal light-foraging strategy. In addition, beech exhibited the highest specific leaf area, a greater total leaf area per unit tree height, a slightly greater leaf area index, and a greater plasticity to light in total leaf area. Ash and maple presented a “gap species” growth strategy, characterized by a marked and constant response in growth rates to increasing light and an inability to strongly reduce their growth rates in deep shade. In shade, they showed some plasticity in displaying most of their leaf area at the top of the crown to minimize self-shading and to enhance light interception. Through this, particularly, maple developed an “umbrella” like crown. These species-specific responses may be used for controlling the development of mixed-species regeneration in shelterwood systems.     

Effects of slash retention and wood ash addition on fine root biomass and production and fungal mycelium in a Norway spruce stand in SW Sweden

In the study reported here we examined the short-term effects (1–3 years) of slash retention (SR) and the long-term effects (13–15 years) of wood-ash application (A) on fine roots and mycorrhizae in a 40-year-old Norway spruce forest in southwest Sweden. Soil cores were used to obtain estimates of the biomass (g m⁻²) of roots in three diameter classes (<0.5, 0.5–1 and 1–2 mm), root length density (RLD), specific root length (SRL) and mycorrhizal root tip density (RTD). Fine root (<1 mm) length production and mortality, and mycelium production, were estimated using minirhizotron and mesh bag techniques, respectively. Compared with the control plots (C), the biomass of fine roots in diameter classes <0.5 mm and 0.5–1 mm was significantly higher in A plots, but lower in SR plots. In addition, RLD was significantly lower in the humus layer of SR plots than in the humus layers of C and A plots, but not in the other layers. None of the treatments affected the SRL. In all soil layers, the SR treatment resulted in significant reductions in the number of ectomycorrhizal root tips, and the mycelia production of fungi in mesh bags, relative to the C treatment, but the C and A treatments induced no significant changes in these variables. Fine root length production in the C, A and SR plots amounted to 94, 87 and 70 mm tube⁻¹ during the 2003 growing season, respectively. Fine root mortality in treated plots did not change over the course of the study. We suggest that leaving logging residues on fertile sites may result in nitrogen mineralisation, which may in turn induce reductions in root biomass, and both root and mycelium production, and consequently affect nutrient uptake and the accumulation of organic carbon in soil derived from roots and mycorrhizae.     

Compared effects of competition by grasses (Graminoids) and broom (Cytisus scoparius) on growth and functional traits of beech saplings (Fagus sylvatica)

? The effect of two weeds (grasses and broom, Cytisus scoparius) competition on the growth and functional traits of European beech saplings (Fagus sylvatica) was investigated in an experimental plantation in the French Massif central.

? We hypothesized that grasses would have a much more harmful effect than broom on beech growth through strong competition for soil water and nitrogen.

? A randomized block design was used with three separate blocks, each possessing three types of vegetation; grasses (mainly Agrostis capillaris, Holcus mollis and H. lanatus), broom, and bare soil as a control.

? Two years after plantation beech sapling growth was significantly greater in bare soil than with competition from grasses, and was intermediate on broom. Growth was positively correlated with beech leaf nitrogen content (N_m) and xylem water potential (Ψ_x). N and Ψ_x were dependent, with nitrogen uptake probably decreasing with increasing drought. There was no clear correlation between growth and light transmitted 10 cm below the apex of the saplings.

? Beech leaf mass on an area basis (LMA) was correlated with the three resource variables (water, nitrogen and light) and with growth, confirming that LMA is a sensitive functional trait integrating the degree of stress experienced by the saplings.

? The results suggest that broom is less competitive than grasses and, if properly managed on a sufficiently well-watered site, can generate satisfactory conditions for beech establishment and growth.

     

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.