Survival and Growth of Picea abies Regeneration after Shelterwood Removal with Single- and Double-grip Harvester Systems

Three stands in Sweden were followed for 7 yrs to study the survival and growth of Norway spruce [Picea abies (L.) Karst.] regeneration after shelterwood removal with single- and double-grip harvester systems. The height of all seedlings within circular plots was recorded before and after final cutting. The tallest seedling and one randomly selected seedling on each circular plot were permanently marked and their annual height growth was measured each year during the period 1993-1999. No significant differences between single- and double-grip harvester systems were found with respect to their effects on regeneration density, survival or annual height growth. The annual height growth of the individual seedlings after shelterwood removal was significantly correlated with seedling height before shelterwood removal. Significant differences were found in annual height growth between seedlings with high and low vitality. Logging damage did not affect the annual height growth. Several practical conclusions were drawn. First, the choice of harvester system does not affect seedling height growth or survival after shelterwood removal. Secondly, the removal of a 200 m³ shelterwood with 200 stems ha^-1 does not seem to constitute a serious problem for seedling survival and growth. Finally, estimated vitality and seedling height both seem to be fairly good predictors of postrelease survival and growth.     

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.     

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.     

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.     

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.     

Growth and Terminal Bud Formation in Picea abies Seedlings Grown with Alternating Diurnal Temperature and Different Light Qualities

The root collar diameter and the height:diameter ratio are of particular importance in container-grown seedlings where a high density in the containers may produce spindly seedlings. Temperature regimes and light quality are known to affect plant growth. The aim of this study was to identify responses in Picea abies (L.) Karst. seedlings grown with light providing different red:far-red ratios and under temperature regimes with alternating day (DT) and night temperature (NT) from negative (DT < NT) to positive (DT > NT) difference (DIF) between DT and NT. Experiments were conducted in controlled environment chambers and in a daylight phytotron. Only limited thermoperiodic responses appeared in P . abies seedlings with respect to seedling height and dry weight accumulation. The formation of terminal buds, however, was clearly delayed in seedlings grown at negative DIF. The results indicate a requirement for day extension light that is high in far-red, to prevent terminal bud formation under natural short-day conditions. An extended study should be conducted to clarify the minimum level of light intensity and the optimal light quality needed to prevent terminal bud formation under natural short-day conditions.     

Natural Regeneration in Clear-cuts: Effects of Scarification,Slash Removal and Clear-cut Age

The effects of soil scarification (mounding), slash removal and clear-cut age on the natural regeneration in clear-cuts was evaluated using data from four sites in southern Sweden. The treatments were carried out during a good seed and establishment year for birch ( Betula pubescens Ehrh. and B. pendula Roth). Scarification had the strongest positive effect on the density of naturally regenerated seedlings, especially in birch, but also in pine ( Pinus sylvestris L.) and spruce [ Picea abies (L.) Karst]. Slash removal had a positive effect on birch density. No statistically significant effect of clear-cut age was found. In addition, no statistically significant interactions between clear-cut age and scarification or slash removal were found. The ingrowth of field vegetation was the fastest in areas that were not scarified, less rapid in areas scarified on old clear-cuts, and the slowest after scarification in fresh clear-cuts. In conclusion, it may be possible to control the density of birch during a good establishment year for birch. If birch is desired, the best combination of treatments is to remove the slash and scarify; otherwise, these treatments should be avoided.     

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.     

Community of Aphyllophorales and Root Rot in Stumps of Picea abies on Clear-felled Forest Sites in Lithuania

The community of Aphyllophorales fungi in stumps of Picea abies (L.) Karst. and the occurrence of root rot caused by Heterobasidion annosum were investigated at 38 clear-felled sites in Lithuania. Fruit-bodies were recorded on 36.0% of the 3924 examined stumps. The most common were Bjerkandera adusta, Phlebiopsis gigantea, Fomitopsis pinicola, Gloeophyllum sepiarium, Stereum sanguinolentum, Trametes zonata and H. annosum. All species occurred in stumps both with and without established root rot, but B. adusta, F. pinicola, G. sepiarium, T. zonata and Trichaptum abietinum occurred less often in stumps containing root rot, while H. annosum was encountered more often. Greater numbers of fruit-bodies were found on stumps cut in summer than on those cut in winter. The fruiting became most abundant 3-4 yrs after the trees were felled. The abundance of nearly all Aphyllophorales correlated positively with the stump diameter, and on larger stumps, fruit-bodies of several different fungi were more common. The number of stumps containing H. annosum root rot in different sites varied from 9.8% to 68.8%, and was 27.6% on average. The incidence of root rot correlated neither with the age nor with the density of the felled stand, but it correlated negatively with the proportion of deciduous trees within a stand.     

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.     

Optimizing the Management of a Butt-rotted Picea abies Stand Infected by Heterobasidion annosum from the Previous Rotation

There are no instructions on the management of stands of Norway spruce [ Picea abies (L.) Karst.] in which butt rot caused by Heterobasidion annosum coll. (Fr.) Bref. was not discovered until the time of first thinning. A stochastic simulation model describing the spread of butt rot in a stand of Norway spruce was used with a new submodel describing the butt rot of young trees. Non-linear stochastic optimization was used to determine the most profitable management schedule of a young Norway spruce stand with butt rot. If the initial level of butt rot was 5% or 10%, or the stand was thinned in summer with stump treatment, one thinning with a rotation of 55 yrs was optimal. Two winter thinnings with a rotation of 61 yrs was optimal in an initially healthy stand. Optimizations indicated that silvicultural measures that decrease the transfer of H. annosum to next generation at clear-cuttings could be economically fulfilled.     

Effects of aluminium and calcium on Picea abies seedlings

Newly‐germinated Norway spruce seedlings were grown for 15 weeks in pots of iron podzol soil that received additions three times a week of nutrient solution containing six levels of Al (0, 0.37, 0.74, 1.5, 3.0 or 6.0 mM) and two levels of Ca (0.09 or 0.46 mM). The Ca concentrations in shoots and roots decreased with increasing Al levels. Growth was suppressed at 0.74 mM Al in the nutrient solution, which gave 0.5 mM Al in saturated soil extracts at the end of the experiment.     

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.     

Effects of early cleaning on young Picea abies stands

Young Norway spruce stands (Picea abies [L.] Karst) are typically cleaned of non-crop species on one or several occasions during young stand stage. In order to objectively evaluate the perceived benefits of early cleaning (EC), we studied the effects of EC on three study sites 2–2.5 years after receiving the treatment. Experiments were established as a randomised complete block design with a total of 40 blocks. Although height growth and mortality were not significantly affected, we found that EC substantially reduced canopy competition and, consequently, the mean diameter of released spruce grew 21–32% faster depending on the site. Furthermore, non-crop trees that were cleared during EC had fewer sprouts in two of the three sites and the sprouts were substantially smaller than the corresponding non-crop trees on control in all sites. EC can reduce the cost of pre-commercial thinning because EC reduced the estimated time needed for subsequent management by 18–49%. Thus, EC offers forest owners and managers an economically viable young stand management option.     

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.     

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.     

Effects of storage conditions on dormancy and vigor of Picea abies seeds

The dormancy and vigor of Picea abies seeds were studied after five months of storage at 25 and 75% relative humidity (RH) and 5 and 12 °C. Dormancy was evaluated by studying germination response to light and moist chilling at 12 and 21 °C. Dormancy causing germination reduction in the dark was induced in seeds during storage at 25% RH (5–6% moisture content, MC) in darkness. The dormancy was greater when seeds were germinated at 12 than at 21 °C and after storage at 12 than at 5 °C. The effective dormancy relief by light indicates that germination was under phytochrome control. Moist chilling could partly replace light. According to accelerated aging and leachate conductivity tests, dry seeds could be stored at 12 °C for five months without affecting their vigor. After storage at 75% RH (11% MC) and 5 °C, the seeds germinated slowly and incompletely. The decreased germination response to light indicates that other processes than those mediated directly by phytochrome restricted germination of these seeds. The positive germination response to moist chilling suggested that secondary dormancy was induced in the seeds. However, vigor tests gave some evidence of simultaneous decrease of vigor. Storage at 75% RH and 12 °C decreased germination nearly to 0%. Germination of seeds stored at 75% RH could be stimulated by a short accelerated aging period.     

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.