Effect of Thinning and Phlebiopsis gigantea Stump Treatment on the Growth of Heterobasidion parviporum Inoculated in Picea abies

The spread of Heterobasidion parviporum Niemelä & Korhonen in roots of Norway spruce was studied in three unthinned first rotation stands of Norway spruce [Picea abies (L.) Karst.] on former agricultural land in south-western Sweden. Heterobasidion parviporum was inoculated at stump height into the trunk of 135 standing trees in a randomized block design. One year after inoculation, two-thirds of the trees were thinned out and one-third was left standing. Half of the stumps left by thinning were treated with spores of Phlebiopsis gigantea (Fr.) Jül and half were left untreated. The spread of H. parviporum was examined both 3 and 5 yrs after inoculation. The rate of spread of H. parviporum and the proportion of infected roots were found to be significantly higher in the root systems of the stumps than in those of the standing trees. Treatment with P. gigantea had no significant effect on the development of H. parviporum in the stumps. There was a tendency 5 yrs after inoculation, however, for a lower proportion of H. parviporum-infected roots in the stumps treated with P. gigantea than in the untreated stumps. In conclusion, thinning of infected Norway spruce was found to increase the rate of spread of H. parviporum in the root systems of the infected trees, which could increase the risk of a rapid build-up of infection in the remaining stand.     

Allozyme Variation of Picea abies in Poland

The aim of this study was to determine the levels and patterns of allozyme variation among 29 populations of Norway spruce [ Picea abies (L.) Karst.] from Poland. Thirteen investigated isoenzyme systems were encoded by 27 gene loci and, on average, 71% of the loci per population were polymorphic. The average and effective numbers of alleles per locus were 2.17 and 1.26, respectively, while the expected heterozygosity was 0.156. A relatively low allozyme differentiation among populations from north-eastern and southern Poland was observed ( F ST = 0.028, mean genetic distance D = 0.005). The results suggest that historical events and extensive gene flow played an important role in the distribution of the observed allozyme differentiation of Norway spruce in Poland.     

Effect of Nutrient Optimization on Branch Characteristics in Picea abies (L.) Karst

The number and size of branches are major determinants of log grading. To study the effect of fertilization on branch characteristics in Norway spruce, trees were sampled in a nutrient optimization experiment in northern Sweden. Data were collected from 24 40-yr-old trees from fertilized and control plots after 12 yrs of annual nutrient application. On the fertilized plots, all essential macronutrients and micronutrients were supplied in irrigation water every second day during the growing season. Fertilization increased the number of new branches, but its effect on branch angle and branch mortality was small. Fertilization increased the diameter of the branches that were formed after the initiation of the treatment, but also the diameter of the branches that were formed before the fertilization commenced. Fertilization enhanced the general growth of a tree, but there were no significant changes in the relationships between stem and branch characteristics. Improved growth rate of trees, as an effect of fertilization, may impair branching quality of logs. Such negative effects may, however, to some extent be counteracted by planting density at stand establishment and later thinning regimens.     

A probability model for root and butt rot in Picea abies derived from Norwegian national forest inventory data

Root rot in Norway spruce (Picea abies (L.) Karst.) causes substantial economic losses to the forestry sector. In this study, we developed a probability model for decay at breast height utilizing 18,141 increment cores sampled on temporary plots of the Norwegian National Forest Inventory. The final model showed a good fit to the data and retained significant relationships between decay and a suite of tree, stand and site variables, including diameter at breast height, stand age, altitude, growing season temperature sum (threshold 5°C), and vegetation type. By comparing model predictions with recorded decay at stump height in an independent data set, we estimated a proportionality function to adjust for the inherent underestimation of total rot that will be obtained by applying a probability model derived from increment cores sampled at breast height. We conclude that the developed model is appropriate for national and regional scenario analyses in Norway, and could also be useful as a tool for operational forestry planning. This would however require further testing on independent data, to assess how well the new model predicts decay at local scales.     

Annual Variation in Pollen Contamination and Outcrossing in a Picea abies Seed Orchard

A mature Norway spruce [Picea abies (L.) Karst.] seed orchard, established in southern Finland with 67 clones from northern Finland, was analysed in three different years in order to estimate the pollen contamination ratios. Allozyme-based paternity analysis revealed that the contamination rate was high, 69-71%, and did not differ between the years studied. It appears that, in areas where spruce is the dominant tree species, the contamination rate will be very high even in mature seed orchards. However, the contamination rate in the thinned parts of the orchard was significantly lower than that in the unthinned parts in two of the three years studied. The outcrossing rate was also high, 96-100% in all years, even though the ramets of each clone were planted using a clonal-row design, and there were no significant differences in the outcrossing rates between the different parts of the orchard.     

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.     

Potential Effects of Temperature on Early Reproductive Development and Progeny Performance in Picea abies (L.) Karst

In order to enhance cone production in Norway spruce [Picea abies (L.) Karst] seed orchards have been established in more southerly, warmer sites in Norway. This has led to concern and some evidence that seedlings obtained from parent trees grown at warmer sites may not be well adapted if planted in northern regions. Selective events during sexual reproduction in the warmer climate may select for traits not well adapted for cold climates. This study describes ovule and male gametophyte development, fertilization and proembryo and early embryo development under six different temperature regimens of warm or cold, warm-cold or cold-warm, and compares development in relation to temperature sums in degree days (dd) and calendar date. The outside treatment in a cold climate in Norway served as the control, and normal development and few ovule abortions and abnormalities were observed. In no treatments were abnormalities in male gametophyte development observed. In treatments in which trees were kept in the greenhouse throughout the study or moved during the study from the outside to inside the greenhouse, a higher incidence of megagametophyte, egg or embryo abortion was generally observed.     

Effects of Seedbed Substrates on Regeneration of Picea abies from Seeds

The effects of seedbed substrates on the emergence and mortality of Norway spruce [ Picea abies (L.) Karst.] on clear-cuts were studied in a sowing experiment. Seven intact substrates as well as scarified plots on the same types of substrates were considered. The experiment took place at two locations in south-east Norway, during three consecutive years. Of intact substrates, Sphagnum spp. was found to be the best for emergence and litter was found to be the second best. Microhabitats dominated by feather mosses, Dicranum spp., Vaccinium myrtillus or Deschampsia flexuosa were poor environments for emergence. However, after two to three seasons, survival in per cent of emerged seedlings was more or less equal among all substrates. Emergence was higher on scarified plots than on intact substrates, and except for Sphagnum substrates, survival was equal or better. For scarified plots, the original substrate did not influence the emergence of seedlings, but mortality after the second summer differed owing to original substrate. The best substrates for scarification were V. myrtillus and D. flexuosa , while mortality was highest for plots that were originally dominated by Sphagnum . Weather conditions influenced the suitability of the substrates as seedbeds, as did the location of the plots.     

细胞分裂素和硝酸铵对欧洲云杉不定芽伸长的影响

报道细胞分裂素和硝酸铵对欧洲云杉不定芽伸长过程的影响.采用5年生的欧洲云杉1个无性系不定芽丛来研究玉米素和硝酸铵对不定芽生长的影响.试验3种诱导类型(玉米素浓度分别为6.84、13.68、20.52 μmol·L-1,激动素浓度0.69 μmol·L-1,培养4周),玉米素最高浓度处理诱导的伸长芽数量最多,在4个无生长调节剂继代培养过程中,加入了1个硝酸铵浓度逐渐增加的继代培养过程.经不含生长调节剂的低浓度的硝酸铵(2.5 mmol·L-1)培养基的继代培养后转入含有细胞分裂素的诱导培养基,下一个继代培养基补充的硝酸铵浓度分别为2.5、7.5、12.5、17.5 mmol·L-1,8周(2个继代培养)后转入含有硝酸铵浓度为2.5 mmol·L-1的培养基培养.硝酸铵浓度为7.5 mmol·L-1的处理可形成数量最多的不定芽,这对于采用不同浓度的玉米素进行诱导的处理结果是一致的,但随着硝酸铵的浓度增至12.5 mmol·L-1和17.5 mmol·L-1时则不定芽的数量减少.欧洲云杉的胚轴外植体经1次或多次诱导后,细胞分裂素(BA、激动素、玉米素、2iP)促进了不定芽的分化和发育,其中BA由于可诱导产生大量的不定芽而成为最有效的细胞分裂素.玉米素虽然诱导不定芽相对弱一些,却在BA多次诱导后可促进芽的伸长.这对获得具有形成不定芽能力并可扩繁的芽丛非常重要.     

Effect of Air-filled Porosity and Organic Matter Concentration of Soil on Growth of Picea abies Seedlings after Transplanting

The effects of air-filled porosity (AFP) and organic matter concentration (OMC) of soil on the growth of Norway spruce [Picea abies (L.) Karst.] seedlings were studied in a greenhouse experiment. One-year-old seedlings were planted into 250 pots filled with five different growth media based on low-humified sphagnum peat and fine sand. The media were mixed to achieve five levels of OMC (1, 25, 50, 75 and 97% by mass). Five AFP levels (5, 10, 20, 30 and 40%) were applied to the mixtures of growth media during irrigation. The growth attributes of the seedlings were measured after seedlings had grown in the greenhouse for 15 weeks. The shoot height and mass growth as well as root mass were significantly higher in 20, 30 and 40% AFP than in 5 and 10% AFP (p<0.05). The longest shoots were produced in growth media with 25, 50 and 75% OMC (p<0.05). The effect of OMC on root mass, while significant (p=0.03), was less pronounced than the effect on height growth and mass of the shoots (p<0.001). The results indicate that, for good seedling growth in pots in greenhouse conditions, AFP should be 20-40% and OMC 25-75% in the growth medium.     

Incidence of Root and Butt Rot in Consecutive Rotations of Picea abies

The incidence of butt rot in two consecutive rotations of Norway spruce [Picea abies (L.) Karst.] in 28 permanent sample plots at four different sites in Denmark was evaluated. Incidence of butt rot was estimated by visual examination of stumps at final felling of the previous rotation and by examination of bore cores taken at the butt from a random sample of trees before first thinning of the subsequent rotation. There was no correlation between the incidence of butt rot at final felling of the previous rotation of Norway spruce and the incidence of butt rot at first thinning of the subsequent rotation of Norway spruce. The incidence of butt rot at final felling was between 19 and 100%, and at first thinning between 0 and 20%. The S-form of Heterobasidion annosum (Fr.) Bref. was the most commonly found decay-causing organism at all sites. Root systems of 28 trees without decay at stump height in the present rotation were excavated to estimate the incidence of root rot. Heterobasidion annosum was found in only one root. Resinicium bicolor (Alb. & Schw. ex Fr.) Parm. was found in 25% of the excavated root systems. The result of the study shows that the incidence of butt rot at first thinning of Norway spruce is not necessarily higher on sites where the previous rotation was heavily infected than on sites where infection in the previous rotation was low.     

Reconstructing 100-150 years of logging history in coastal spruce forest (Picea abies) with special conservation values in central Norway

Coastal spruce forests of central Norway harbour a unique assemblage of epiphytic lichens and are given high priority with respect to conservation of biodiversity. To assess the historical impact of logging during the last 100-150 yrs, 31 remnant stands were studied by means of tree-ring analysis of 2199 trees and the decay stage of 1605 stumps. No stands had been clear-cut, but all had been selectively logged at least twice during the last 150 yrs. Total harvested timber volume ranged from 65 to 409 m³ha-¹ (31-124% of present-day standing volume) and the selective logging kept standing volume low (40-200 m³ha-¹) during 1890-1930. Present-day stand characteristics were strongly correlated with site productivity and topographic position within the ravine valleys. Low amounts of dead wood at sites with high historical logging activity was the only consistent relationship found after covariance of site productivity, topographic position and deciduous trees were taken into account. The results indicate that old-growth stand characteristics, such as reversed J-shaped age distributions and dead wood in advanced decay classes, can be obtained 100-150 yrs after intensive selective logging.     

Effects of Partial Cutting and Scarification on Planted Picea abies at Mid-elevation Sites in South-east Norway

Mortality, injury and height growth of planted Picea abies (L.) Karst, were examined over a 6 yr period in eight stands at middle altitudes (430–620 m a.s.l.) in south-east Norway. There were four residual stand density treatments (shelterwoods of high, medium and low density, 25 × 25 m patch-cut) combined with three scarification treatments (unscarified, patch scarification, inverting) in a split-plot design. Mortality was very low during the experimental period, and did not differ significantly between the residual stand density treatments (mean = 11.1%). Both survival and plant height after 6 yrs were improved by inverting, while patch scarification was intermediate but not significantly different from the unscarified alternative. The patch-cutting resulted in the tallest plants, while only minor differences in height growth were found between the shelterwood treatments. The frequency of injured plants after 6 yrs was not significantly affected by the treatments (mean = 10.6%). The improved plant establishment at inverting is in agreement with previous studies on clear-cuts.     

Root growth capacity and field performance of Pinus sylvestris and Picea abies seedlings

Correlations between root growth capacity (RGC), at the time of planting, and field performance were studied for Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings. Before planting a gradient in seedling viability was generated through exposure to low root temperatures and different winter storage regimes. The hypothesis that high RGC values would improve field performance was to some extent verified for pine seedlings while no correlations could be registered for spruce. Reasons for these results are discussed.     

Genotypic variation in natural infection frequency of Heterobasidion spp. in a Picea abies clone trial in southern Sweden

Abstract

Natural root and butt rot infection with Heterobasidion spp. was recorded on three occasions in trees from a 20-year-old Norway spruce [Picea abies (L.) Karst.] clone trial. In the first assessment, carried out when the trial had been running for 15 years, 29.5% of the trees felled were infected with Heterobasidion spp., but no statistically significant differences were found between the different clones. In the second assessment, also at 15 years, bore cores were taken from 595 trees. Of these cores, 16.5% were infected, but again no statistically significant differences were found between the different clones. Combining the two first assessments also revealed no significant differences. The discrepancy in rot incidence between the two assessments was explained by the inaccuracy of the bore core method. The third assessment was carried out after the trial had been running for 20 years. At that time, 424 trees of 50 clones were felled and sampled. Visible infection was recorded in 46.5% of the trees. The differences between clones were statistically significant and the broad-sense heritability estimate was 0.18. There were no significant genotypic correlations between tree size and rot infection; this is important for the Norway spruce breeding strategy. Calculations show that economic viability would increase by 0.28 SEK per plant for every 10% decrease in root rot frequency at final felling.     

欧洲云杉、黑云杉、白云杉苗期对比试验

选择欧洲云杉3个种源、黑云杉3个种源、白云杉3个种源、青海云杉1个种源进行了不同种、种源的苗期生长性状变异试验。结果表明：云杉不同种、种源间苗期性状差异显著。4个种的1年生苗以苗高进行排序：欧洲云杉〉黑云杉〉白云杉〉青海云杉。欧洲云杉、黑云杉、白云杉各个种的不同种源间有很大的变异及选择潜力。因此在进行云杉引种时,应选择不同的种、种源,才能取得更好的效果。     

Critical Factors Affecting Ex Vitro Performance of Somatic Embryo Plants of Picea abies

The potential to use somatic embryos for large-scale propagation of elite genotypes, for integration into breeding programmes and for connecting breeding and mass propagation, is receiving much attention. However, before the methods are applied it is important that the plants regenerated via somatic embryogenesis grow as expected, i.e. as seedlings or cuttings. Growth of somatic embryo plants is under a cumulative influence of a number of treatments given during the in vitro phase and during the ex vitro establishment phase. The aim of this study was to identify treatments with a negative influence on the subsequent growth of somatic embryo plants of Norway spruce (Picea abies L. Karst.). Based on the results, the time of contact with abscisic acid during somatic embryo maturation and the length of continuous light treatment (CLT) during the first growth period strongly affect the height growth during two successive growth periods. In both cases longer treatments exerted negative effects. Based on these results a new method was set up, which includes: (1) prematuration treatment of the suspension culture in a growth regulator-free medium, by which the maturation step is synchronized and contracted; and (2) a two-phase germination treatment, first on a solidified medium and then in a liquid medium. This treatment avoids extended CLT during the first growth period. Another advantage of the two-phase germination treatment is a better root-system development. Somatic embryo plants produced according to this method can be transferred directly from in vitro conditions to the greenhouse.     

Variation in Tree Size and Resistance to Ceratocystis polonica in a Monoclonal Stand of Picea abies

Tree resistance to the pathogenic blue-stain fungus Ceratocystis polonica was studied in a monoclonal stand of Norway spruce [ Picea abies (L.) Karst.] in relation to tree social status and diameter at breast height (DBH). The DBH distribution of the 33-yr-old stand ranged from 5 to 35 cm. There were clear differences in tree height between the suppressed (DBH 7.4-10.3 cm), codominant (DBH 11.8-17.4 cm) and dominant (DBH 18.6-23.9 cm) tree classes. The resistance was tested by mass inoculating trees with a low (400 inoculations m -2 , 60 cm inoculation belt) or high (400 inoculations m -2 , 120 cm inoculation belt) dosage. The small, suppressed trees were more susceptible to inoculation than the codominant and dominant trees, based on the amount of blue-stained and occluded sapwood, lesion length and dead cambium/phloem. A threshold in tree social status or tree size may be important in the overall resistance to fungal infection.     

Effects of Previous High N Addition on Nutrient Conditions in Above-ground Biomass of a Picea abies Stand in Sweden

The effect of different thawing procedures on seedling quality in Norway spruce (Picea abies) was evaluated. Freezer-stored seedlings were thawed rapidly for 20 h by immersion in water with an initial water temperature of 8°C or thawed more slowly by gradually increasing the temperature in storage over a period of 8 weeks. Seedlings from these treatments were also compared with seedlings thawed rapidly overnight at 15°C. Frost tolerance, carbohydrate content and the time of bud break were used as indicators of the vitality of the seedlings. Seedlings thawed rapidly in water or air showed significantly better frost tolerance after storage than slowly thawed seedlings. The contents of sucrose and raffinose in seedlings thawed rapidly were also significantly higher than in seedlings thawed more slowly. Bud break occurred later in seedlings thawed rapidly than in slowly thawed seedlings. Rapid thawing in water may be a good method to ensure the delivery of high-quality thawed seedlings throughout the planting season.     

Radial,Height and Volume Increment Variation in Picea abies (L.) Karst. Stands with Varying Thinning Intensities

Variation in radial increment at different stem heights was compared with stem volume increment and height increment using thinning experiments of Picea abies (L.) Karst. in southern Finland. The magnitude of growth variation was similar on trees from different crown classes and stand densities. However, there was higher autocorrelation in the codominant and intermediate trees and trees on dense plots. The relationship between short-term volume increment and radial increment variation at breast height was not affected by thinnings. While short-term growth variation was similar at different stem heights, some evidence was found for differences in medium-term variation between radial increment at breast height and volume increment. Height increment reacted to unfavourable climatic events later than radial and volume increment. Radial and volume increments were negatively correlated to temperature and positively correlated to precipitation of the current summer. Height increment was positively related to current early summer temperature, but negatively to temperature of the previous summer.