Stand and site characteristics in the decline of pinus sylvestris caused by gremmeniella abietina shoot disease in a severely damaged 50‐year‐old plantation in se lapland

The role of stand and site characteristics in a 50‐year‐old, sowed Scots pine (Pinus sylvestris L.) stand on the decline of Scots pine caused by Gremmeniella abietina was studied. Treewise damage of Scots pine was modelled using stand and site variables and stepwise regression analysis. The significant variables included in the model were stand elevation from the main cold air centre and the pH of the humus layer, while the other less significant variables were the number of stems per hectare of Scots pine and Norway spruce, content of Cu and Ni in the humus layer, and the thickness of the humus layer and the A₂ horizon. The studied variables excluded from the model were the number of broad‐leaved trees per hectare, Al and Pb contents in the humus layer, and Al, Cu, Pb and Ni contents in the A₂ horizon. The results strongly suggest that the vicinity of water as well as local depressions act as vulnerable sites for Scots pine and the said vulnerability is enhanced by increased pH of the humus layer.     

Seed cone abortion and full seed production in an indoor seed orchard with potted grafts of Picea abies

Abortion of seed cones and production of full seeds have been recorded in an indoor‐potted Norway spruce (Picea abies (L.) Karst.) seed orchard at Biri nursery in Norway. Seed cone buds aborted their normal development at much higher frequencies (22–45%) inside isolation bags than in an open‐air (4–12%) greenhouse environment. The grafts had been induced to flower the year prior to seed cone development. Treatment with gibberellin A_4/7 in conjunction with heat reduced the proportion of aborted, isolated flowers (13%) compared to treatment with heat alone (22%) in a clonally balanced material. The production of full seeds per cone increased from 3 in 1987 to 50 in 1991. The full seed production is discussed in relation to methods and timing of pollen forcing. It is anticipated that 1 kg of seeds could be produced annually in a 1250 m² greenhouse.     

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.     

Effect of uninucleate Rhizoctonia on Scots pine and Norway spruce seedlings

One‐year‐old container‐grown seedlings were planted in spring on clear cut areas: the Norway spruce (Picea abies) on a moist upland site (Myrtillus‐type) and Scots pine (Pinus sylvestris) on a dryish upland site (Vaccinium‐type). While still in the nursery, half of the seedlings of each species had been inoculated during the previous summer, with a uninucleate Rhizoctonia sp., a root dieback fungus. At outplanting all the seedlings appeared healthy and had a normal apical bud, although the height of the inoculated seedlings was less than that of the uninoculated control seedlings. At the end of the first growing season after planting, the mortality of inoculated Scots pine and Norway spruce seedlings was 25 and 69%, respectively. After two growing seasons the mortality of inoculated seedlings had increased to 38% for Scots pine and 93% for Norway spruce. The mortality of control seedlings after two growing seasons in the forest was 2% for Scots pine and 13% for Norway spruce. After outplanting the annual growth of inoculated seedlings was poor compared with the growth of control seedlings. These results show that, although Rhizoctonia‐affected seedlings are alive and green in the nursery, the disease subsequently affects both their survival and growth in the forest.     

Quantitation of gibberellins A9, A1 and A3 in relation to flower bud differentiation in Picea abies

Endogenous gibberellin A₁ (GA₁), GA₃, GA₄ and GA₉ were quantitated in elongating shoots of Norway spruce [Picea abies (L.) Karst.] grafts with a good or a poor flowering history. The grafts were grown either in a natural environment outdoors, cool and wet (CW) treatment, or in a greenhouse with elevated temperatures and controlled drought stress, hot and dry (HD) treatment. The GAs were quantitated by gas chromatography‐mass spectrometry selected ion monitoring (GC‐MS SIM) using deuterated GA₁, GA₃, GA₄ and GA₉ as internal standards. Terminal shoots from the second whorl of branches were harvested at 76%, 86% and 96% shoot elongation for the CW treated grafts and at 90% and 99% shoot elongation for the HD treated grafts. The content of GAs in the CW grafts was highest during most rapid shoot elongation, GA₉ being the dominant GA. The levels decreased as shoot elongation ceased. This was also noted for GA‐content in shoots of the HD treated grafts. A comparison of the GA‐amounts at ca. 96% of total shoot elongation for the CW treated grafts and ca. 99% of total shoot elongation for the HD treated grafts revealed that shoots of the good flowering clone had a higher content of GA₉ and a lower content of GA, and GA₃. When comparing the HD treated and CW treated grafts, the shoots of HD treated grafts contained higher concentrations of GA₉ but lower concentrations of GA, and GA₃. Calculating the ratios between [GA₉] and [GA₁] resulted in a ratio of 12.5 and 36.6 for the good flowering clone grown outdoors and in greenhouse, respectively. The same ratios were for the poor flowering clone 1.45 and 3.8 when grown outdoors and in greenhouse, respectively. A higher ratio may indicate a higher capacity of synthezise the importance of flowering GA₄ from GA₉ and a lower conversion of GA₄ to GA₁, thereby favouring the diffentiation to reproductive buds.     

Irradiance in young stands of picea abies (L.) karst. and pinus sylvestris L. and the possibilities to prevent suckers of broad‐leaved trees

Incoming shortwave global radiation (Q _g) and photosynthetically active radiation (PAR; Q _pa as a fraction of full daylight, relative irradiance (%Q), were measured at the same time in young stands of Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.). Measurements were made on three levels above ground: 20 cm above ground and 50 and 15 % of stand height. Stands of three heights (75, 150 and 300 cm) were studied during two months. The stands were created by arranging young trees cut from natural stands, in nine quadratic spacings: 0.1, 0.2, 0.3, 0.4, 0.5, 0.7, 1.0, 1.4 and 2.0 m. The leaf area index (L) was estimated. Differences in %Q‐values for Q _pa and Q_g in the same species and at the same stand height and level of light measurements above ground were significant only in 150 and 300 cm stands. In 75 cm high stands of Scots pine, the %Q was <60% at spacings <0.2 m and in 75 cm high stands of Norway spruce, the %Q was <60% at spacings <0.3 m. Only at 0.1x0.1–0.3x0.3 m, %Q was <20% in Norway spruce stands. In 150 cm stands %Q<20% was measured at spacings <0.7 m in Norway spruce and <0.5 in Scots pine. In 300 cm stands of Norway spruce it was measured up to 1.2 and in Scots pine <1.1 m. Light extinction coefficients, K and K_g for spruce and pine stands were 0.17–0.40 and 0.16–0.31 respectively. Some practical implications of the study are presented. Lack of light (%Q<10%) as a single factor of seriously suppressed growth and development of broad‐leaved plants and suckers by competition in young stands of spruce and pine only occur in dense stands 0.3x0.3 m‐1.1xl.l m (8000–100000 stems/hectare). Competition by light on a regenerated area generally occurs in the level of 50% of tree height (150–300 cm) and higher due to the rapid growth of broad‐leaved trees (sprouts) compared with planted conifers but the light intensity at these levels %Q>10%.     

Growth response of coniferous stands to whole‐tree harvesting in early thinnings

A series of 15 field experiments was established to quantify the growth response of first‐thinning stands of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst) to whole‐tree harvesting and to estimate the need for nutrient compensation. The experiments were undertaken in Finland, Norway and Sweden and represent a wide range of site conditions. The site index (H ₁₀₀) of Scots pine stands varied from 19 to 29 m, and that of Norway spruce stands from 28 to 36 m. Total amounts of biomass and nutrients removed were calculated based on data obtained from felled sample trees. During the first 5‐yr period the growth response to the removal of logging residues varied considerably in both pine and spruce stands. Regression analyses did not reveal any functions that explained the variation in results satisfactorily. In cases where whole‐tree harvesting influenced tree growth negatively, this effect was counteracted by compensatory fertilization. It was concluded that to determine the response of remaining trees to harvesting intensity reliably, the post‐harvest period analysed must be longer than 5 yrs.     

Infection experiments with Gremmeniella abietina on seedlings of Norway spruce and Scots pine

Conidia of Gremmeniella abietina infected and caused disease symptoms in annual shoots of both Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) seedlings. In Norway spruce shoots the infection remained largely latent, with only a few seedlings showing symptoms. Mycelial growth inside the shoots was faster in Scots pine than in Norway spruce and was favoured by low temperature in both hosts. The shoots of Norway spruce seedlings had higher endophyte populations than those of Scots pine, and the populations were decreased by low temperatures. Reductions in the normal epiphytic or endophytic flora by acid mist treatments seemed to favour the development of G. abietina.     

Freezing temperatures in the root zone—effects on growth of containerized Pinus sylvestris and Picea abies seedlings

Roots of 1‐year‐old containerized seedlings of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) were experimentally frozen in December. The seedlings were then grown for 3 weeks in a growth chamber and evaluated with regard to root growth capacity (RGC) and shoot elongation. The subsequent RGC of Scots pine declined as root zone temperatures were lowered from ‐6°C to ‐11°C and from ‐11°C to ‐16°C. Almost no root growth was observed after exposure to ‐20°C. Shoot growth was also negatively affected by low root temperatures but less than root growth. Low root temperatures did not affect Norway spruce as much as Scots pine, although root and shoot growth of Norway spruce were reduced after exposure to the lowest test temperatures (‐16°C and ‐20°C). The length of exposure, ranging between 1 and 8 hours had no effect on subsequent growth.     

Effect of planting Scots pine around Norway spruce stumps on the spread of Heterobasidion coll.

Management of a Norway spruce stand planted on a site infected by Heterobasidion coll. is problematic because the fungus spreads vegetatively from the colonized stumps of the previous generation to the new seedlings. Growing of mixed stand with more resistant tree species has been suggested to decrease the economic losses caused by butt rot in Norway spruce trees. The mechanistic simulation model Rotstand describing the spread of Heterobasidion coll. in coniferous stands of southern Finland was used to study the effect of planting Scots pines around colonized clear‐felling stumps of Norway spruce of the previous generation. Planting of Scots pines in clusters around colonized stumps markedly decreased the butt rot of Norway spruce trees at the age of 20 years and at clear felling. If the same number of Scots pines were planted randomly, the effect was weak. When the average diameter of colonized clear‐felling stumps was 30 cm, a Scots pine circle with a radius of 3 m resulted in the highest soil expectation value (SEV) at 2% discounting rate, whereas with 40‐cm stump diameter, a 4‐m radius produced the highest SEV. When the proportion of Heterobasidion parviporum in the old colonized stumps was 50% instead of 95%, planting pines around colonized stumps still clearly decreased the butt rot at the age of 20 years and in final felling.     

Variations in seed germination of Hippophae salicifolia with different presoaking treatments

Mature seeds of H. salicifolia, collected from five provenances (i.e. Hanuman Chatti, Helang, Lata, Rambara and Janggal Chatti) in west Himalaya, India, were treated with stratification (at 4°C for 15, 30 and 60 days) and in different concentrations of GA₃ (5, 10, 20 mM), KNO₃ (50, 100, 200 mM) and Thiourea (50, 100, 200 mM) solution to determine the variations in seed germination. Results reveal that the germination rates of seeds from different provenances under different pre-sowing treatments are significantly increased compared to those in control (24%–30%). The seeds treated with Thiourea (100 mM) have highest germination rate (76%–83% for different seed sources), followed by those (63%–71% for different seed sources) pretreated with stratification (4°C, 30 days). GA3 treatment significantly shortens the mean germination time (MGT) and improves seed germination percentage. Considering the practical applicability and cost effectiveness, thiourea (100 mM) and stratification (at 4°C) treatments for seed germination are recommended for mass multiplication through seeds of H. salicifolia in village/forest nurseries of the west Himalaya, India.     

Long‐term effects of liming on the fine‐root standing crop of Picea abies and Pinus sylvestris in relation to chemical changes in the soil

The long‐term effects of lime application on fine roots of Norway spruce, Picea abies (L.) Karst, and Scots pine, Pinus sylvestris (L.), have been studied in five experimental forest stands subjected to different lime applications 5 to 18 years before the present study was undertaken. The effects of liming does not seem to significantly influence fine‐root development in forest stands in the long term. The only response to liming in measured root variables was a tendency to increased specific root length (SRL = fine‐root length/fine‐root dry weight, m/g). A correlation between increased SRL, decreased root biomass and increased stem volume growth was indicated. Changes in water extractable amounts of mineral elements—P, K, Ca, Mg, Mn, S, Al and Fe‐in bulk soil and rhizosphere soil from the mineral soil layers were studied in a control area and an area treated with 3830 kg CaCO₃ ha^‐1. Few significant differences were found between treatments, and then mainly in the case of Ca.     

La culture in vitro de conifères alpins

The in vitro culture of alpine conifers . Seeds of European larch, Scots pine, Mugo pine and Norway spruce were sterilized for 30 min in a bath of 30% hydrogen peroxide. Similarly, seeds of Swiss stone pine, of which the tegument had been removed with a little vice, were sterilized for 15 min. Germination and a sterility test were undertaken on malt-agar. Finally, the seeds were transplanted in vitro on two media for mycorrhizal synthesis.     

Interference of bracken (Pteridium aquilinum L. Kuhn) with Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies L. Karst.) seedling establishment

The nature of interference of bracken with Scots pine and Norway spruce seedling establishment was considered in three field experiments. In a seeding experiment, it was found that Scots pine germination was highest on exposed mineral soil and lowest when intact bracken litter and humus were present, suggesting adverse effects of litter and humus on pine regeneration probably due to phytotoxicity. In a second experiment, smothering by bracken caused high mortality of Scots pine seedlings while Norway spruce seedlings were relatively unaffected. Mortality for both Scots pine and Norway spruce seedlings was low when planted in a adjacent Scots pine-bilberry stand with no bracken. Annual shoot growth of Norway spruce was higher in bracken than in Scots pine-bilberry vegetation while no differences in shoot growth between these two vegetation types occurred for Scots pine. In a third experiment, activated carbon was added to the ground under Norway spruce seedlings planted in bracken to adsorb possible phytotoxic compounds released by bracken. The addition of carbon had no effect on seedling mortality or growth rate, indicating that the seedlings were not susceptible to allelochemicals released by bracken. Since large Norway spruce seedlings were relatively unaffected by bracken interference in this study, artificial regeneration with containerized Norway spruce seedlings is suggested to achieve an acceptable conifer tree establishment on clear-cuts invaded by bracken.     

Geological controls on conifer distributions and their implications for forest management in Finnish Lapland

Abstract

Soil water and nutrient regimes of naturally established old-growth conifer stands and those of intensively managed Norway spruce [Picea abies (L.) Karst.] sites were assessed over a range of lithological provinces in Finnish Lapland. Soil dielectric permittivity (ε), as a measure of soil water content (θ _v) and soil electrical conductivity (σ _a), as a measure of soil solute content, were species specific, such that high soil θ _v>0.27 cm³cm^?3 (ε>15) constitutes an edaphic constraint for Scots pine (Pinus sylvestris L.) and low soil solute content (σ _a<0.5 mS m^?1) is constraining for Norway spruce. The spatial pattern of the soil θ _v was temporally stabile, such that intraseasonal and interseasonal soil θ _v was significantly higher in silty tills of spruce stands compared to sandy tills of pine stands. Scots pine was the only conifer on tills derived from felsic rocks of Hetta granite (HG) and Lapland granulite (LG). Norway spruce dominated on tills derived from the mafic rocks of Lapland greenstone belt (LGB), but tills of LG and HG constitute a dispersal barrier for spruce. Mechanical site preparation (MSP) with ploughing (Marttiini) was not able to amend soil θ _v to meet site requirements of Scots pine at former spruce sites. MSP resulted in significant reduction in soil nutrient content such that untreated control σ _a>tilt/shoulder σ _a>trench σ _a. The results imply that MSP treatments through which cross-contour tracks are created pose a risk to the sustainability of soil quality in Lapland.     

Damage by pine weevil Hylobius abietis to conifer seedlings after shelterwood removal

Abstract

Pine weevil (Hylobius abietis L.) damage to seedlings after overstorey removal was investigated in a survey study in six shelterwoods in the south–central part of Sweden. The shelterwoods predominantly consisted of Scots pine, except at one site where the shelter trees mainly consisted of Norway spruce. Before final cutting, 10 plots were laid out at each site and measurements of shelter trees and marked seedlings were taken. The seedlings were examined during the 2 years after final cutting. The study showed that removal of shelter trees increases the risk of severe damage by pine weevil and the variable that was most strongly correlated with the risk was the seedling root collar diameter. Both Scots pine and Norway spruce seedlings were severely damaged by pine weevil, and most of the feeding occurred during the first year after cutting. The amount of debarked area was significantly larger for Scots pine than for Norway spruce seedlings. Vitality (growth of the leading shoot before final cutting) of the seedlings also affected the probability of damage. Seedlings with high vitality were less damaged by pine weevil than seedlings with low vitality. For Scots pine the shelterwood density before final cutting was correlated to the intensity of pine weevil feeding after cutting. In conclusion, after the final cutting of a pine or spruce shelterwood, pine weevils will probably invade the area. To avoid serious damage, Norway spruce and Scots pine seedlings should have reached a diameter of at least 10–12 mm.     

Effects of seed water content and storage temperature on the germination parameters of white spruce, black spruce and lodgepole pine seed

The effect of seed water content (WC) (2–3, 5–6 and 22–25%, on a fresh weight basis), storage temperature (+4, −20, −80 and −196°C) and storage duration (6, 12, 24, 48 and 60 months) on the germination of white spruce (Picea glauca (Moench) Voss), black spruce (Picea mariana (Mill.) B.S.P.) and lodgepole pine (Pinus contorta Dougl. ex Loud. var. latifolia Engelm.) seed was investigated. Germination of white spruce control (untreated) seeds and seeds adjusted to 2–3% and 5–6% WC declined after 48 months of storage at −80 and −196°C, with a further decline at 60 months at −20, −80, −196°C. Germination remained high when control white spruce seeds and seeds with 2–3, 5–6% WC were stored at +4°C, over all storage durations. Generally, black spruce and lodgepole pine exhibited high germination at all storage temperatures at 2–3% and 5–6% WC as well as the control (untreated) seed, for up to 60 months in storage. Germination declined for all three species when seed was conditioned to 22–25% WC. This loss in germination was partially recovered in white spruce seed stored at +4, −20 and −80°C after storage durations of 24, 12 and 48 months, respectively, and in black spruce seeds stored at −20 and −196°C after storage durations of 24 months. Mean germination time (MGT) was relatively constant for all species, under all conditions, except for seed conditioned to 22–25% WC, where MGT increased for white spruce seed stored 48 months at −80 and −196°C, and for black spruce seed stored 24 months at +4 and −80°C and 60 months at −196°C. These results show that the optimal storage temperatures are 4°C for white spruce, and 4, −20, −80, and −196°C for black spruce and lodgepole pine, and 2–6% water content is optimal for all 3 species at these temperatures.     

Persistence of a biocontrol strain of Phlebiopsis gigantea in conifer stumps and its effects on within‐species genetic diversity

Fungal isolations and genetic fingerprinting were used to determine whether Phlebiopsis gigantea stump treatment against Heterobasidion annosum sl. using a single genotype (Rotstop) would affect the genetic diversity of P. gigantea populations. The survival time of P. gigantea was longer in Norway spruce (Picea abies) stumps compared to Scots pine (Pinus sylvestris) as no isolates were obtained from pine stumps 6 years after treatment, whereas in about half of the spruce stumps the fungus was still present. The usage of Rotstop did not seem to increase the occurrence of the fungus 5 years after the treatment in fresh (1‐year‐old) untreated stumps within the same forest stands. All the isolates from the 6‐year‐old treated spruce stumps were identical in genotype with the Rotstop‐strain, whereas all isolates from the fresh untreated spruce and pine stumps differed from it. Within the treated pine stand, the biocontrol usage seemed to have caused a slight reduction in genetic markers not related to Rotstop, but there were no statistically significant differences between the marker frequencies and the local natural population. Thus, Rotstop is not likely to cause any immediate threat to the genetic diversity of P. gigantea.     

Equations for estimating above- and belowground biomass of Norway spruce,Scots pine,birch spp. and European aspen in Latvia

This study aims to derive allometric functions to estimate the above- and belowground biomass components of the most important tree species in Latvia. The study material included a total of 81 Norway spruce (Picea abies [L.] Karst), 102 Scots pine (Pinus sylvestris L.), 105 birch spp. (mainly silver birch (Betula pendula Roth)) and 84 European aspen (Populus tremula L.) trees sampled in 124 forest stands. The suitability of three mathematical models for the prediction of total aboveground biomass, stem biomass, total live and dead branch biomass, belowground biomass and small root biomass was evaluated. Our analysis revealed that the use of the Intergovernmental Panel on Climate Change mean default values for the root-to-shoot ratio recommended for temperate and boreal ecological zones leads to the overestimation of root biomass of young trees, especially Scots pine and Norway spruce. Our findings indicate that biomass functions recommended for other Baltic Sea countries are not appropriate for the assessment of the biomass stock in Latvia’s forests because these lead to biased estimates. The biomass functions derived in our study are recommended for reporting the biomass stock in Latvia.     

Microbial Inoculants for Sustainable Forests

Abstract

Fungal root pathogens are widespread and may cause substantial seedling losses in conifer nurseries. Furthermore, poor seedling survival and growth on reforestation sites results in reduced forest regeneration. Use of microbial inoculants for disease control and plant growth promotion has become an important endeavour. A microbial culture collection of 500 strains was assessed for biological control of fungal root pathogens and/or plant growth promotion of conifer seedlings. Seven of these strains showed significant suppres-sive effects on various soil-bome fungal pathogens. On Douglas fir, two strains, RAL3 and 64-3, reduced disease caused by Fusariumby. 7-42% in repeated growth room assays. The same strains significantly increased healthy stand of white spruce seedlings inoculated with Fusariumand Pythiumin a conifer nursery, and increased the survival of bare-root white spruce seedlings planted on a reforestation site by 19-23%. Both strains also significantly increased new root and total plant dry weights. Strain RAL3 in commercial formulation maintained a viable population of about log 8-9 cfu/ml for over a year when stored at 5°C. Strain survival on seed varied with conifer species. No decreases in bacterial populations were observed on seeds of jack pine or Douglas fir after 37 to 44 days storage at 5°C, but decreases were observed on seeds of white spruce and Scots pine. This study has provided candidate beneficial microbial inocu-lants which offer promise for development of commercial inoculants for the forestry industry.