Canopy density in stands of Picea abies and Pinus sylvestris after different thinning methods

Altogether 82 plots (261 estimations) of Picea abies (L.) Karst, and 193 plots (360 estimations) of Pinus sylvestris (L.) stands were estimated by a vertical tube. The “crown free projection”, CFP, of stands thinned in three methods with different thinning grades was measured: unthinned, heavily and very heavily thinned, heavily thinned delayed first thinning, extra heavily thinned and thinned from the top. Basal area (m²ha^?1) density (stems ha^?1) and diameter sum (m ha^?1) were plotted against CFP. Basal area was the best practical measure of stand in this study. Generally Scots pine stands have higher CFP and the curves are steeper than in Norway spruce stands. Depending on the grade of thinning, heavily and very heavily thinned spruce stands, delayed first thinning included, have CFP values of 10–15% and stands thinned from the top, 20–40%, compared with 30–80% and 30–60% respectively in pine stands. Extra heavily thinned stands have the highest CFP, 20–80% in spruce and 50–90% in pine stands. The CFP levels after thinning are too high in pine stands for avoidance of sucker and sprout production of aspen and birch. In dense Norway spruce stands thinned from the top or heavily and very heavily thinned, the CFP values are low enough (≤30%) to diminish the production of suckers.     

Root-associated fungi of healthy-looking Pinus sylvestris and Picea abies seedlings in Swedish forest nurseries

The aim of this study was to assess fungal communities in roots of healthy-looking Pinus sylvestris L. and Picea abies (L.) Karst. seedlings in nine forest nurseries in Sweden using a combination of traditional culturing and direct sequencing of internal transcribed spacer of fungal ribosomal RNA (ITS rRNA) from the roots. Culturing from 1800 surface-sterilised root segments resulted in 2387 fungal cultures representing 42 different taxa. Direct sequencing from 180 root segments resulted in 119 ITS rRNA sequences representing 25 different taxa. In total, 55 different fungal taxa were detected using both methods. Although direct sequencing was more efficient than culturing in detecting different fungal taxa, both methods provided complementary information about fungal communities in roots since each detected rather different groups of fungi. The most dominant taxa detected by culturing were Trichoderma viride Pers. (19.5%), Phoma mucivora Davey & Currah (19.1%), Phialocephala fortinii Wang & Wilcox (17.4%) and Meliniomyces variabilis Hambl. & Sigler (10.2%), while Thelephora terrestris Ehrh. (26.1%), Unidentified sp. NS126 (25.2%) and Heliotales sp. C20 (10.1%) were most commonly detected by direct sequencing. In conclusion, results showed that forest nurseries in Sweden harbour diverse communities of fungi associated with the roots of healthy-looking P. sylvestris and P. abies seedlings. Although fungal communities were often dominated by saprotrophs and endophytes, several facultative pathogens were also detected indicating that under suitable conditions they may be a potential threat to the plants.     

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.     

Intraspecific variation in Heterobasidion annosum for growth in sapwood of Picea abies and Pinus sylvestris

Two greenhouse experiments were conducted to study intraspecific variation in growth of the root rot fungus Heterobasidion annosum in living host sapwood. In experiment 1, Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) seedlings were inoculated with H. annosum isolates, 14 each of the S-and P-intersterility groups, collected from various parts of Sweden. In pine, the P-group isolates were more virulent than the S-group isolates both in terms of infection frequency, induced mortality rate (p < 0.05), and fungal growth in sapwood (p < 0.05). In spruce, the P-group isolates were also more virulent on average, but the difference was not statistically significant. Both S and P isolates had a higher infection frequency and a significantly longer sapwood growth on spruce than on pine. The P-group caused higher mortality on pine than on spruce. The length of the lesion in the inner bark was strongly correlated with fungal growth in spruce, but not in pine where the lesions were short or absent. In experiment 2, ten Norway spruce clones were inoculated with 18 S-isolates, originating from nine live-decayed trees and from nine spore-infected stumps in a single Norway spruce stand. The objective was to test whether any selection for growth rate in sapwood was detectable among individuals of H. annosum originating either from stumps or trees. The results gave no support for such selection since no difference in sapwood growth between the two groups of isolates was found.     

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.     

Incidence of Gremmeniella abietina in planted seedlings of Picea abies and Pinus sylvestris in northern Finland

The ascomycete Gremmeniella abietina causes a disease (scleroderris canker) on conifers. The pathogen kills terminal shoots and branches of Pinus sylvestris and Picea abies, sporulates on host shoots and causes leader changes (i.e. dieback). Damage caused by G. abietina was investigated in young P. abies and P. sylvestris plantations in northern Finland. Side branches from seedlings of both species and the main stems of P. sylvestris were collected from three sites in two locations (Poksa 1 and 2, Kivalo) in northern Finland. The number of cankers and leader changes was counted in branches of each age (i.e. year) in a total of ca. 6300 shoots of P. abies and 1200 shoots of P. sylvestris. Cankers were common on both P. abies and P. sylvestris in Poksa 1 (2002–2008) and on P. abies in Poksa 2 (1998–2001). In Kivalo, cankers occurred only sporadically on P. sylvestris. Leader changes were most frequent in 1999–2009 in Poksa 1, in 2001–2009 in Poksa 2 and in 2003 in Kivalo. Cankers and other symptoms of infection were more obscure on P. abies than on P. sylvestris. According to both conidial morphology and molecular analysis, the strain of G. abietina infecting both P. abies and P. sylvestris was small‐tree type (STT). This is the first report of G. abietina, STT or B type, injuring P. abies in plantations.     

Stem Form and Taper Changes After Thinning and Nitrogen Fertilization in Picea abies and Pinus sylvestris Stands

Stem form and taper changes after thinning and thinning combined with N fertilization were studied in 23 Norway spruce [Picea abies (L.) Karst.] and 46 Scots pine (Pinus sylvestris L.) stands in Sweden observed for 5-28 yrs. Average taper was calculated for the stem section 1.3-6.0 m above ground level and as the ratio between the diameter at breast height and total tree height for periods of 5-10 yrs. At the start of the experiment, before the first thinning, the dominant height was 12-15 m. Once 65% of the basal area had been removed in a single thinning from below, the remaining trees showed a strong increase in taper compared with trees in unthinned stands. Stems in stands treated with heavily recurrent thinnings from below also increased in taper, although the increases were not always statistically significant. Trees in thinned, N-fertilized Scots pine stands in middle and northern Sweden developed a more pronounced taper compared with stems in equally thinned, unfertilized stands.     

Mean canopy stomatal conductance responses to water and nutrient availabilities in Picea abies and Pinus taeda

We compared sap-flux-scaled, mean, canopy stomatal conductance (GS) between Picea abies (L.) Karst. in Sweden and Pinus taeda (L.) in North Carolina, both growing on nutritionally poor soils. Stomatal conductance of Picea abies was approximately half that of Pinus taeda and the sensitivity of GS in Picea abies to vapor pressure deficit (D) was lower than in Pinus taeda. Optimal fertilization increased leaf area index (L) two- and threefold in Pinus taeda and Picea abies, respectively, regardless of whether irrigation was increased. Although it increased L, fertilization did not increase GS in Picea abies unless irrigation was also provided. In Pinus taeda growing on coarse, sandy soils, the doubling of L in response to fertilization reduced GS sharply unless irrigation was also provided. The reduction in GS with fertilization in the absence of irrigation resulted from the production of fine roots with low saturated hydraulic conductivity. When Pinus taeda received both fertilization and irrigation, the increase in L was accompanied by a large increase in GS. In Pinus taeda, a reference GS (defined as GS at D = 1 kPa; GSR) decreased in all treatments with decreasing volumetric soil water content (theta). In Picea abies, theta varied little within a treatment, but overall, GSR declined with theta, reaching lowest values when drought was imposed by the interception of precipitation. Despite the large difference in GS both between Picea abies and Pinus taeda and among treatments, stem growth was related to absorbed radiation, and stem growth response to treatment reflected mostly the changes in L.     

Responses of Pinus sylvestris and Picea abies Seedlings to Limited Phosphorus Fertilization and Treatment with Elevated Ozone Concentrations

Scots pine ( Pinus sylvestris L.) and Norway spruce [ Picea abies (L.) Karst.] seedlings were exposed to high phosphorus (HP) or low phosphorus (LP) availability for one growing season in the open field, and to combined P availability and elevated ozone (O 3 ) concentrations (0, 55, 110 and 210 ppb for Scots pine and 0, 40, 75 and 150 ppb for Norway spruce, respectively) for 28 days in controlled laboratory chambers. Compared with HP, the LP treatment reduced Scots pine current-year (C) shoot and root dry masses and Norway spruce total dry mass, whereas the highest O 3 concentrations increased the magnesium concentration of Scots pine C needles and P concentrations of the C needles of both tree species. Chlorophyll a, a+b and carotenoid concentrations of Scots pine C needles were significantly higher in the LP treatment compared with HP under the highest O 3 concentration (210 ppb). In the mesophyll tissue of C needles of both tree species, LP treatment increased the size of mitochondria and elevated O 3 -induced granulation of chloroplast stroma and disintegration of cytoplasm. Exposure to elevated O 3 concentrations increased swelling of chloroplast thylakoids and reduced the amount of vacuolar tannin in the LP Scots pine C needles. The results suggest disturbances in needle photosynthetic machinery due to acute exposure to the combination of elevated O 3 and low P availability. However, clear additive effects were found only in needle P concentrations < 1 mg g -1 in short-term O 3 exposure.     

Variation in sapwood thickness of Picea abies in Estonia depending on the tree age

Variation of sapwood thickness in terms of a linear measurement and a growth ring count with reference to the age was studied in dominant and suppressed Norway spruce trees. In the trees of both dominance classes the sapwood thickness increased in absolute terms, while its relative portion decreased with the age of the trees growing. Great differences were found in sapwood thickness between dominant and suppressed trees in linear measurements but not in the number of sapwood rings or the rate of the heartwood formation. The number of sapwood growth rings reached 40 in old trees. The index of vigour differed considerably in dominant and suppressed trees.     

Growth rates of Heterobasidion annosum s.s. and H. parviporum in functional sapwood of Pinus sylvestris and Picea abies

Growth rates of H. annosum s.s. and H. parviporum were investigated in the functional sapwood of young Pinus sylvestris and Picea abies plants as an indicator of the relative susceptibilities of the hosts to these pathogens. The stems of 520 five‐year‐old P. abies and 321 four‐year‐old P. sylvestris plants were inoculated and the extent of infection determined 16 weeks later. H. annosum s.s. grew further than H. parviporum in P. sylvestris sapwood, while in P. abies, no differences between the two Heterobasidion spp. were found. Both H. annosum s.s. and H. parviporum spread faster in the sapwood of P. abies than in P. sylvestris. There was high within‐host species variation in growth rates for both P. sylvestris and P. abies suggesting it may be possible to identify tree genotypes with lower susceptibility.     

Age-related changes in foliar morphology and physiology in red spruce and their influence on declining photosynthetic rates and productivity with tree age

The contribution of changes in meristem behavior to age-related decline in forest productivity is poorly understood. We studied age-related trends in needle morphology and gas exchange in a population of red spruce (Picea rubens Sarg.) growing in a multi-cohort stand where trees ranged from first-year germinants to trees over 150 years old, as well as in grafted scions from these trees. In the field study, age-related trends in foliar morphology were determined in six cohorts ranging in age from 2 to 120 years, and differences in gas exchange characteristics were compared between 60- and 120-year age classes. In a common-rootstock study, scions from trees representing 20-, 60-, and 120-year cohorts were grafted onto juvenile rootstock and maintained for three growing seasons, after which morphological and physiological foliar attributes were evaluated. The field study revealed significant age-related trends in foliar morphology, including decreasing specific leaf area, and increasing needle width, projected area, and width/length ratio. Similar trends were apparent in foliage from the grafted scions. Both in situ foliage and shoots of grafted scions from the oldest cohort showed significantly lower photosynthetic rates than their counterparts from younger trees; however, differences in stomatal conductance and internal CO(2) concentrations were not significant. These results suggest that: (1) foliage of red spruce exhibits age-related trends in both morphology and physiology; (2) age-related decreases in photosynthetic rates contribute to declining productivity in old red spruce; (3) declines in photosynthetic rates result from nonstomatal limitations; and (4) age-related changes in morphology and physiology are inherent in meristems and persist for at least 3 years in scions grafted to juvenile rootstock.     

Application of growth regulators in aqueous media and organic solvents to seeds of Picea abies and Pinus sylvestris

Incubation with acetone and dichloromethane had little effect on the germination of different lots of Norway spruce seeds. This was not the case with Scots pine where almost half the seed lots tested were adversely affected by acetone and dichloromethane. The auxins indole‐3‐acetic acid (IAA) and indole‐3‐butyric acid (IBA) were applied to Scots pine seeds in acetone, dichloromethane and phosphate‐citrate buffer. The uptake of the two compounds was most rapid from acetone. Although the uptake from dichloromethane was slower than from acetone it reached the same level after 6 h. Uptake of the growth regulators from buffer was significantly lower than from the organic solvents. The accumulation of IAA and IBA into the seeds from both acetone and dichloromethane was linear between 0.5 and 10 mM, and between 0.5 and 15 mM from buffer. IAA and IBA levels in infused seeds declined over a 5‐day germination period, the former more rapidly than the latter. The metabolic activity of the IAA degradation system was as effective in seeds after immersion in the organic solvents as seeds immersed in buffer.     

Estimating foliage biomass in Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) plots

Dynamic decomposition models are needed to estimate changes in the carbon stock of boreal soil because these changes are difficult to measure directly. An important aboveground carbon flux to the soil is foliage litterfall. To estimate this flux, both the amount and the turnover rate of the foliage biomass component must be known. Several methods for estimating foliage biomass of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.), including biomass equations and biomass expansion factors (BEFs), were compared with predicted foliage biomass based on forest inventory plot-level measurements. Measured foliage biomass was up-scaled from the branch-level to the plot-level by combining forest inventory variables (diameter, height, height at the crown base and crown base diameter) based on the assumptions of pipe model theory. Combining the foliage biomass: cross-sectional area ratio with the forest inventory variables provided accurate estimates of foliage biomass at the plot-level for plots in southern Finland. The results emphasize the need to test biomass equations with independent data, especially when the equations applied are based on neighboring regions.     

The effect of tree architecture on conduit diameter and frequency from small distal roots to branch tips in Betula pendula, Picea abies and Pinus sylvestris

We studied the effect of tree architecture on xylem anatomy in three Betula pendula Roth., three Picea abies (L.) H. Karst. and three Pinus sylvestris (L.) trees (mean age 35 years). First, the analysis of conduit anatomy in different tree parts showed that conduits tapered and their frequency increased from roots (≥ 2 mm) to stem, from stem to branches and further to leaf petioles in B. pendula. Conduit anatomy in lateral and main roots, as well as lateral and main branches, significantly differed from each other in all the studied species. The increase in conduit diameter and decrease in frequency from the pith to the bark were clear aboveground, but variable patterns were observed belowground. In the leaf petioles of B. pendula, conduit diameter increased and conduit frequency decreased with increasing individual leaf area. Second, the results concerning the scaling of conduit diameter were compared with the predictions of the general vascular scaling model (WBE model) and Murray's law. The scaling parameter values at the tree level corresponded with the predictions of the WBE model in all the studied trees except for one tree of both conifer species. However, the scaling parameter values changed from one tree compartment to another rather than remaining uniform inside a tree, as assumed by the WBE model. The assumptions of the WBE model of a constant conductivity ratio, constant tapering and an unchanged total number of conduits were not fulfilled. When the conductivity ratio and relative tapering were plotted together, the results aboveground corresponded quite well with Murray's law: the conductivity ratio increased when relative tapering decreased. Our results support the theory that trees adjust both their macro- and microstructure to maximize their water transport efficiency, but also to prevent embolism and ensure mechanical safety.     

Aluminium concentration versus the base cation to aluminium ratio as predictors for aluminium toxicity in Pinus sylvestris and Picea abies seedlings

Aluminium (Al) toxicity is considered an important factor in forest deterioration caused by soil acidification. A ratio of base cations (BC) to Al in the soil solution lower than 1 is widely used as an indicator for potentially adverse effects on tree health. In our view, the validity of the assumptions underlying the use of the BC:Al ratio as an indicator for Al toxicity in trees has never been evaluated properly.Here, we evaluate the importance of the base cations Ca and Mg in counteracting Al toxicity. Pinus sylvestris and Picea abies seedlings were grown on nutrient solution with a range of Al (0–0.25–0.5–1–2 mM) and base cation (0.25–0.5–2 mM) concentrations, giving BC:Al ratios of 1 at different levels of Al. Increasing concentrations of Al in solution caused growth reductions, which could not be counteracted by increasing concentrations of BC in solution with P. sylvestris and only partly counteracted with P. abies. Increased concentrations of Al in solution decreased the concentrations in shoot and root of both Ca and Mg, while increased concentrations of BC in solution increased tissue concentrations of BC. Growth reductions were, however, not a result of BC deficiencies, as growth reduction already occurred in tree seedlings that maintained adequate concentrations of Ca and Mg.All growth and uptake variables measured showed a higher or equal correlation with the absolute concentrations of Al or Al+BC in solution than with the BC:Al ratio. We conclude that Al toxicity is determined solely by the concentration of Al in solution. Shoot growth decreased significantly as dissolved Al increased at a constant BC:Al ratio of 1. In P. abies, but not in P. sylvestris, dissolved BC can positively affect uptake of BC and growth, which might partly alleviate the toxic effects of Al. Our results show that the mechanistic explanation for the effect of the BC:Al ratio is insufficient to describe Al toxicity. Care should be taken when using models based on the BC:Al ratio to predict the effect of Al on tree growth.     

S and P intersterility groups in Heterobasidion annosum; Infection frequencies through bark of Picea abies and Pinus sylvestris seedlings and in vitro growth rates at different oxygen levels

Seedlings of Norway Spruce and Scots Pine were inoculated with S and P strains of the root rot fungus Heterobasidion annosum. Infection frequencies through the living bark layers were not related to subgroup affiliation in neither spruces nor pines. Growth rates in vitro were not related to growth rates in vivo. Oxygen concentration had a stronger effect on the growth rates of P than on S strains.     

Snow-packing as a potential harmful factor on Picea abies,Pinus sylvestris and Betula pubescens at high altitude in northern Finland

Snow-packing, a combination of ice, hoarfrost and snow on trees, and the subsequent tree damage by snow, were estimated on Norway spruce (Picea abies), Scots pine (Pinus sylvestris), and pubescent birch (Betula pubescens) in the winter of 1993-94 in southern Lapland, northern Finland, near the local alpine timberline around 400 m a.s.l. Snow-packing on each fully sized tree increased from a few hundred kilograms at 150–250 m a.s.l. to a maximum of 3290 kg at 300–350 m a.s.l. At 300 m a.s.l., snow-packing per metre of stem increased from 30 to 50 kg on trees < 5 m in height up to 180–200 kg on 20-m trees. There was 300 000-480 000 kg/ha of snow accumulated on tree crowns. No stem breakage by snow-packing occurred at or below 250 m a.s.l., whereas at 290–350 m a.s.l., 0-46%, 39-100%, and 0–33% of the spruce, pine and birch trees, respectively, had broken tops. Birch appeared to be the most resistant and pine the most susceptible to snow breakage.     

Morphological and Ecological Variation of Gremmeniella abietina var. abietina in Pinus sylvestris,Pinus contorta and Picea abies Sapling Stands in Northern Finland and the Kola Peninsula

The morphological and ecological variation of two types of Gremmeniella abietina var. abietina causing scleroderris canker on conifers was investigated in Pinus spp. and Picea sp. sapling stands in northern Finland and the Kola Peninsula. Small - tree type (STT or B type) of G. abietina was detected alone in 13 Scots pine, three lodgepole pine and two Norway spruce sapling stands out of 26 stands investigated, both STT and large - tree type (LTT or A type) were observed in six Scots pine stands, and LTT was detected alone in two Scots pine stands. For the first time, G. abietina was found to injure Norway spruce saplings in a respective plantation in northern Fennoscandia. STT isolates produced statistically significantly more conidia in vitro than LTT isolates. Morphological variation in conidia septation revealed that STT produced conidia with more than five septa more frequently than did LTT. There was a greater range in variation in septation in STT than in LTT, with overlapping between the types. Isolates of both types were equally associated with cankers, coloured wood, pycnidia or apothecia in the infected saplings.     

Functions for Biomass Estimation of Young Pinus sylvestris,Picea abies and Betula spp. from Stands in Northern Sweden with High Stand Densities

New silvicultural regimes with high within-stand competition require new functions for estimation of standing stock and growth of biomass components, since the allometry of trees is changed by light competition. This paper presents functions for estimation of the aboveground biomass dry weights for stem wood, stem bark, branches and leaves of young (diameter at breast height <10 cm) Scots pine (Pinus sylvestris L.), Norway spruce [Picea abies (L.) Karst.] and birch (Betula pendula Roth. and Betula pubescens Ehrh.) trees growing in dense mixed stands. The functions were derived from a sample consisting of 84 Scots pine, 43 Norway spruce and 66 birch trees from six stands in northern Sweden with high stand densities (>10000 st ha^-1). The logarithmically transformed power function displayed a good ability to stabilize the variance of dry weights and showed a good fit to the material (0.37< R ² <0.99). A comparison with the most commonly used biomass functions in Sweden today showed that they overestimated the weight of stem wood and branches, while the weight of foliage was underestimated. The nature of these discrepancies suggested that the precision of biomass estimations might also be improved for young trees at wider spacing.