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.     

Overstorey density influence on the height of Picea abies regeneration in northern Sweden

The study aimed specifically at investigating if canopy opennesswas a better predictor of the height growth of Norway spruce(Picea abies (L.) Karst.) advance regeneration than overstoreybasal area or overstorey standing volume. In 1990, a field experimentwith 3 x 2 factorial design and two replications (blocks) wasestablished in an uneven-aged Norway spruce forest. Plots hada net plot area of 30 x 30 m, each with a 10-m-wide treatedbuffer zone. Three overstorey density levels retained approximately15, 40 and 70 per cent of the pre-harvest overstorey standingvolume and were allotted to the plots. Two types of thinningthat harvested smaller trees or harvested larger trees wererandomly allocated to each pair of overstorey density plots.In mid-June 2000, canopy openness was estimated from hemisphericalphotographs taken at five marked points in the centre of eachof the plots at 0.9 m from ground to the top of the ‘fish-eye’camera lens. Regression results showed that canopy opennesswas a better predictor of height increments of spruce seedlings(0.1< height < 0.5 m), saplings (0.5 height < 2.0m), and small trees (height 2.0 m, diameter at 1.3 m height< 5 cm) than with overstorey basal area (m² ha^–1) oroverstorey standing volume (m³ ha^–1). The height incrementof the spruce advance regeneration was not significantly correlatedto stand basal area or to standing volume. Overstorey basalarea in the net plots was significantly negative (P 0.05) withmean canopy openness estimates, and the r² value was 0.40. Resultsindicated that basal area was not linearly related to canopyopenness as it increased, which might explain the lack of predictivepower of retained basal area on spruce regeneration height indense stands in boreal Sweden.     

Yield capacity of planted Picea abies in northern Sweden

Norway spruce (Picea abies (L.) Karst.) is the dominant species in the older forests of central northern Sweden. However, spruce has seldom been planted in the area, partly because existing tools for site classification have indicated a low yield capacity for the species. The aim of this study was to examine the yield capacity of spruce on the basis of existing plantations. In total, 91 operational and experimental plantations in the age interval 27–46 yrs were sampled. Stands were located between 62° and 65° N at altitudes 130–620 m a.s.l. Site index was estimated by height growth and site‐factor equations previously developed from old‐growth stand data. Height developments in the plantations indicate that site index for these stands is on average 4.6 m higher than predicted by site‐factor equations. The differences between the two methods are larger on poor sites than on rich sites. No systematic deviations of top height development from the site index curves could be detected on remeasured sample plots. Existing growth models were applied on measured stand data to predict future growth. Calculated mean annual increments were on average 20% lower when site index was predicted by site factors instead of height and age. The bias means that the yield capacity of planted spruce in northern Sweden has been underestimated by about 35%.     

Windthrow after Shelterwood cutting in Picea abies Peatland forests

The aim of the study was to address the question of whether shelterwood cutting should be discouraged as a forest regeneration method in Norway spruce (Picea abies (L.) Karst.) forests on highly productive peatlands due to a higher risk of windthrow in such forests. The total extent of windthrow was observed during six years after shelterwood cutting in nine spruce forests on fertile peatlands in north, central and south Sweden. Shelterwoods at densities of 140 and 200 stems ha^‐1 were studied. At the end of the six year study period, 43% of all trees in the sparser shelterwoods and 38% of the trees in the denser shelterwoods had blown down. With respect to results from recent studies of the development of naturally regenerated and planted seedlings under the shelter trees, the extent of windthrow was regarded acceptable. Analysis of wind data from national weather stations close to the experiments showed that the acceptable extent of windthrow could not be explained by low frequencies of high winds during the shelterwood period. On the contrary, according to the wind observations made at the weather stations selected in central and south Sweden, the annual number of days with high winds (>21 ms^‐1) during the six years after the shelter‐wood cutting was larger than the average high wind frequency for the last 40 years. Observations of maximum wind speeds show that the mean annual gale force during the shelterwood period was higher than the long term average. Thus, the results from this study do not justify avoidance of shelterwood cutting in spruce peatland forests because of the risk of windthrow. Each of the shelterwoods in the experiment was located close to a large clearcut, and the shelterwood cuttings were carried out as heavy thinnings. For further reduction of windthrow, denser shelterwoods and no clearcutting in adjacent stands are suggested when using shelterwood systems in practical forestry.     

Silvicultural alternatives in an uneven-sized forest dominated by Picea abies

In this article, we report on the increments in basal area and tree diameter as well as the structural development observed in variously thinned plots that underwent either uneven- or even-sized treatment. The experimental forest was originally an uneven-sized mixed stand dominated by Picea abies. Twenty-eight randomized sample plots underwent each treatment, and the trees were monitored for 15 growing seasons after thinning. The uneven-sized plots retained a reverse J-shaped diameter distribution, but this was changed into a bell shape by low thinning in the even-sized plots. Absolute basal area increment was positively correlated with basal area in the even-sized treatment but not in the uneven-sized treatment. In the latter, all of the plots grew almost equally well, and only the basal area of broadleaves explained slightly positively the increment variation. Relative basal area increment was negatively correlated with basal area in both treatments. Additionally, the basal area of Scots pine was a positive explanatory variable in the relative increment variation in the even-sized treatment. For the dominant Norway spruce trees, diameter increment was negatively correlated with basal area in both treatments and, conversely, heavy removal increased the diameter increment. Relative basal area increment averaged 5% annually in uneven-sized plots representing the “target selection.” This was more than double the increment observed for the even-sized plots that represented the “prevailing practice.” Likewise, the diameter increment of Norway spruce trees was 48% greater in the uneven-sized compared to the even-sized plots.     

Diameter Increment in Picea abies Shelterwood Stands in Northern Sweden

In this study, the effect of shelterwood density on radial increment and stem form changes was evaluated after 9 yrs for a trial in a forest in northern Sweden dominated by Norway spruce [Picea abies (L.) Karst.]. Radial increment increased after the shelterwood cut, compared with control stands. The effect was of the same magnitude for sparse and dense shelterwoods. The response started in the third growing season and increased until the seventh season after release. The relative increment response was highest in the lower parts of the tree trunk, which indicates a reallocation of increment during the shelterwood period, causing a slight taper deterioration.     

Forecasting development of understory American beech after partial cutting in uneven-aged northern hardwood stands

American beech (Fagus grandifolia Ehrh.) in the understory of northern hardwood stands can interfere with the development of more desirable species such as sugar maple (Acer saccharum Marsh.). Yet, managers have had only limited methods for identifying stands where beech might dominate a new cohort. We examined five uneven-aged stands in New York State to determine the degree of influence that understory beech has on species development after a cutting treatment. No significant differences between stands were found, allowing us to develop a single model for evaluation. Understory beech was assessed using a species index value (SIV), which measures the proportion of stems, weighted by height, for a species on a milacre plot. We used a 5×5 transition matrix to compare pre- and post-cut beech importance levels. Stand-level trends showed an overall increase in the proportion of sample plots with higher levels of understory beech after cutting, with the likelihood of change on individual plots dependent on their pre-cut status. Plots with either no beech or very high levels of beech changed little after cutting, while beech was more likely to reach critical levels of dominance when pre-cut SIV was medium or higher. The transition matrix gives forest managers a means to forecast the probable level of future beech importance following cutting, based upon precursor conditions.     

Fungi in bark peeling wounds of Picea abies in central Sweden

A total of 210 Norway spruce trees with stem wounds resulting from bark peeling by moose (Alces alces) were examined in three 45–50-year-old stands that contained 20-30% of damaged trees. Injured stems were between 8 and 40 cm diameter at breast height (d.b.h.) and showed 1 to 23-year-old wounds. Wounds varied in size from 2 to 4815 cm². The size of injury correlated negatively with the age of the injury (r=-0.24; p=0.004), and positive correlation was established between the age of the injury and d.b.h. of the wounded tree (r=0.50; p=0.0001). Each wound was sampled once using an increment borer and fungi were isolated. Among basidiomycetes, Stereum sanguinolentum was the most common (isolated from 26.7% of the damaged stems). Other common species were Cylindrobasidium evolvens (23.8%), Amylostereum areolatum (5.2%), A. chailletii (0.5%), Heterobasidion annosum (5.2%), Peni-ophorapithya (1.4%), Sistotrema brinkmannii (1.0%). The ascomycete Nectria fuckeliana was the most common among all fungi (present in 35.7% of bark peeling wounds). The frequency of S. sanguinolentum infection correlated positively with the age of the injury (r=0.27; p=0.001) and the opposite relationship was revealed for C. evolvens (r=?0.30; p=0.0001). Furthermore, C. evolvens infection correlated positively with the wound size (r=0.30; p=0.0001) and negatively with the tree d.b.h. (r=?0.20; p=0.004). A positive correlation was found between tree d. b. h. and the occurrence in stems of H. annosum (r=0.23; p=0.001 and N. fuckeliana (r=0.23; p=0.0006). The spruce bark beetle Dendroctonus micans attacked 14.8% of wounded trees. Presence in stems of N. fuckeliana was associated significantly with the D. micans attack (r=0.190; p =0.006; χ² test: p=0.01). Except for a negative correlation between infections of S. sanguinolentum and H. annosum (r=-0.140; p=0.04), no significant relationship between fungal species was found.     

A selection experiment for height growth with cuttings of Picea abies (L.) Karst

Twenty sets, each consisting of a tall, a medium, and a small plant, were selected from a 2 by 2‐year‐old nursery crop. Selection differential was measured. Cutting propagation was undertaken and the subsequent clones were tested in a field experiment. Differences between groups at age 1, 2, 3, 6, 8, and 10 were calculated, and ortet/ramet regressions and age‐to‐age correlations were conducted. Selection of the tall plants gave a response of approximately 17% at age 10. Accumulated genetic gain at this age by a specified stepwise selection procedure is estimated to 36%. Step one is the early selection in the nursery, step two is a later selection amongst the early selected clones and performed after 7 years’ field testing. Ageing of clones, selection for other traits and application of the clones over a variety of sites will reduce these gains.     

Effects of Stump-treatment Substances for Root-rot Control on Ground Vegetation and Soil Properties in a Picea abies forest in Sweden

There is increasing commercial interest in treating stumps to restrict the spread of root rot [Heterobasidion annosum (Fr.) Bref.] when thinning Swedish coniferous forests during the summer. Both chemical and biological substances are used for this purpose. During the treatment, however, a large proportion of the substance applied is spilled beside the stumps. A field study was conducted on the effects of stump-treatment substances on various ground-vegetation species in a Picea abies (L.) Karst. forest in Sweden. Three different substances commonly used in forestry were studied: urea solution (0.23 kg N m^-2), borate solution (10 g B m^-2) and a fungal preparation of Phlebiopsis gigantea (Fr.) Jül. spores (1 g spores m^-2). The principal objectives were to assess whether any of the substances were harmful to plants and whether plant species differed in their sensitivity. Both borate and urea solution caused severe damage to most ground-vegetation species tested. Bryophytes were affected more strongly than vascular plants and urea was slightly more toxic than borate. Treatments with P. gigantea caused no obvious damage. The size and persistence of chemical changes in the soil induced by the treatments were also analysed. Transient changes were apparent in topsoil where borate or urea had been added. Very high concentrations of B were initially observed where borate had been applied, and even after 1 yr they were slightly higher than the threshold concentration at which plant injuries are expected. Urea treatment initially resulted in a pH increase of 2 units and a substantial increase in soil ammonium content. After 1 yr these effects had largely disappeared, although some increase in ammonium was still detectable.     

Environmental influences on juvenile shoot growth in Picea abies

Under certain environmental conditions, juvenile shoot growth characters might give early indications of adaptability and growth potential. Therefore, the reaction patterns of predetermined and free shoot growth were studied on Picea abies (L.) Karst, plants, which at different times during the growth period were treated by (1) application of nutrients, (2) defoliation, and (3) short days. Fertilization caused an increase in both predetermined and free growth, while defoliation and short day treatment caused a decrease in both forms of shoot growth and short days did not allow any free growth to occur. In the growth period following the treatments, fertilization caused more predetermined and less free growth to occur, while defoliation caused less predetermined and more free growth. Predetermined growth is explainable by the finite number of predetermined needle primordia. Free growth is initiated thereafter. It is influenced strongly by the environmental conditions and seems not to be influenced by the amount of preceding predetermined growth. Free growth enables young plants to utilize favourable growth conditions in summer for height growth precociously, which might increase their adaptability and competitive value. However, predetermined growth is the preferable mode of shoot growth because favourable conditions in successive growth periods induce more predetermined than free growth. Predetermined and free growth are well integrated forms of shoot growth giving no supporting evidence to the hypothesis that free and predetermined growth are inherited independently of each other. Rank changes in provenance development are probably not explainable by a loss in height growth when free growth occurs no more due to age, but by other causes.     

Biomass,decomposition and nutrient release of Vaccinium myrtillus leaf litter in four forest stands

Biomass and nutrient transfer (N, P, K, Ca, Mg) of bilberry (Vaccinium myrtillus L.) leaf litter fall, as well as decomposition and nutrient release, were studied in four mature forest stands situated in Central and South Sweden. Bilberry leaf litter fall amounted to between 33 and 55 kg ha^‐1 yr^‐1 in the four stands. Only minor differences between sites were noted for litter concentrations of N, P and Ca, whereas K and Mg showed somewhat larger variability. Relative amounts of the five nutrient elements in the litter fall were generally in the order N > Ca > K > Mg > P. The amounts of nutrients returned to the forest floor by the annual leaf litter fall in the stands ranged from 0.4 to 0.8 kg ha^‐1 for N, 0.4 to 0.6 kg ha^‐1 for Ca, 0.2 to 0.7 kg ha^‐1 for K, 0.1 to 0.2 kg ha^‐1 for Mg and 0.04 to 0.08 kg ha^‐1 for P.

The decomposition of the local bilberry leaf litter was followed by means of litterbags during three years. At all sites there was an extremely rapid mass loss from the litter (between 45% and 54%) during the first four to five months of decomposition. After this initial phase, the decomposition rates decreased markedly and after three years the accumulated mass losses of the litters varied between 64% and 78% at the studied sites. After two and three years of decomposition, three of the sites exhibited almost similar litter mass losses whereas at the fourth site the litter was decomposed to a significantly lower degree. The pattern of nutrient release from the decomposing bilberry leaf litter differed somewhat from site to site. Minor differences were, however, noted for P, Ca and Mg while N and K were more strongly retained in the litter at one of the sites.     

Models for Predicting Wood Properties in Stems of Picea abies and Pinus sylvestris in Sweden

To study and model the variation of wood properties, sample trees were selected from 42 Norway spruce and 20 Scots pine stands covering a wide variation in climatic and site conditions, stand maturation and tree sizes. Plot and tree measurements were followed by sampling wood from different heights in each sample tree and laboratory measurements of wood properties. Mixed linear and non-linear prediction models were developed using diameters, number of annual rings and climatic indices as explanatory variables. The variation in spruce properties explained by these variables was: basic density 50%, latewood content 52%, juvenile wood diameter 85%, heartwood diameter 94% and bark thickness 76%. The corresponding values for pine were 59, 54, 79, 92 and 85%. Random among-tree variance was an important contributor to the remaining variation for density and latewood. In general, only a minor part of the random variation was related to variance between stands. Predictions derived from the models for density and juvenile wood in both species, and heartwood in pine showed good agreement when validated with data sets from two other studies. The resulting models may be used for predicting wood properties in forest planning and in bucking computers in harvesters, provided that the essential information is available.     

Ultrasound emission after cycles of water stress in Picea abies

The relationships among rate of ultrasound acoustic emission (AE), xylem water potential and transpiration rate were investigated in 5-year-old potted saplings of Picea abies Karst. after cycles of water stress. Water-stressed plants displayed minimum xylem water potentials of -3.9 MPa, near-zero transpiration rates and up to 45 AE counts per minute. After rewatering, water-stressed plants no longer produced AEs. Well-watered control plants produced only a small number of ultrasonic AEs. After three cycles of water stress (lasting 24 days in total), it was estimated that about two-thirds of the functional tracheids were embolized. The concomitant reduction in hydraulic conductance was about 70%.     

Genetic parameters of growth and wood quality traits in Picea abies

Genetic parameters were estimated for wood and growth traits in two 19-yr-old clonal trials and a 40-yr-old full-sib progeny trial of Norway spruce [Picea abies (L.) Karst.]. In the clonal trials high (>0.4) broad-sense heritabilities were found for wood density traits, lignin content, number of internal cracks, growth traits, spiral grain and number of resin canals. Moderate (0.2–0.4) heritabilities were found for tracheid lumen diameter and cell wall thickness, microfibril angle and tracheid length, while low heritabilities (<0.2) were found for pulp yield, fibre strength, wood stiffness and wood colour. Lignin content and pulp yield showed low genetic variation, whereas the genotypic coefficient of variation for most other traits ranged between 5 and 15%. Most traits showed low levels of genotype by environment interaction. Among the wood properties, latewood proportion, earlywood density and ring density showed significant, adverse correlations with volume in both clonal trials.     

The impact of Sirococcus shoot blight on radial and height growth of Norway spruce (Picea abies) in young plantations

The impact of Sirococcus shoot blight on the radial and height growth of Norway spruce in a young plantation (approximately 20 years old) was investigated by examining the increment losses for four infection intensities (classes). The average diameter at breast height of trees in the lightly damaged class was 72% when compared with the average diameter of the healthy trees, whereas moderately and severely damaged tree classes were 67 and 57%, respectively. Using tree ring analysis, the development of radial growth over time due to intensity of infection was studied. Height growth of affected trees was also significantly reduced (up to 43%) compared with the healthy trees, thus indicating a dramatic impact of Sirococcus conigenus on the growth of young Norway spruce.     

Long-term Growth Effects Following Forest Nitrogen Fertilization in Pinus sylvestris and Picea abies Stands in Sweden

Forest nitrogen (N) fertilization induces changes in the soil and soil microorganisms that could hypothetically affect the long-term productivity of the soil. The tree growth response following a normal (i.e. 150 kg N ha^?1) single shoot N fertilization has a duration of 7–10 yrs. The aim of this study was to investigate whether any residual effects on tree growth persist, which could be attributed to previous N fertilization. The study included six Pinus sylvestris L. and three Picea abies (L.) Karst. experimental sites, sampled for growth parameters 14–28 yrs after the last fertilization. Residual growth effects were on average small, and not statistically significant. Negative residual growth effects of varying duration could be discerned, especially at low-fertility P. sylvestris sites. However, there was an overall tendency for growth to increase in the long term after N fertilization. The main conclusion is that operational forest N fertilization with a normal N dose should not be regarded as a threat to long-term forest production.