Polyamines in embryogenic cultures of Norway spruce (Picea abies) and red spruce (Picea rubens)

Embryogenic cultures of red spruce (Picea rubens Sarg.) and Norway spruce (Picea abies (L.) Karst.) were initiated from dissected mature zygotic embryos. The tissues were grown on either proliferation medium or maturation medium. On proliferation medium, the embryogenic tissue continued to produce early stage somatic embryos (organized meristems attached to elongated, suspensor-like cells), whereas on maturation medium fully mature embryos developed from the embryonic tissue. Analysis of polyamines in tissues grown on these two media showed that: (1) both putrescine and spermidine concentrations were always higher in cultures grown on proliferation medium than in cultures grown on maturation medium; (2) in both species, spermidine concentrations declined with time in the tissues grown on maturation medium; and (3) spermine was present in only minute quantities and showed only a small change with time. The presence of difluoromethylornithine in the culture medium had little effect on polyamine concentration, whereas the presence of difluoromethylarginine caused a decrease in putrescine concentrations in both red spruce and Norway spruce tissues grown on proliferation medium or maturation medium.     

Applicability of non-destructive substitutes for leaf area in different stands of Norway spruce (Picea abies L. Karst.) focusing on traditional forest crown measures

Since individual tree leaf area is an important measure for productivity as well as for site occupancy, it is of high interest in many studies about forest growth. The exact determination of leaf area is nearly impossible. Thus, a common way to get information about leaf area is to use substitutes. These substitutes are often variables which are collected in a destructive way which is not feasible for long term studies. Therefore, this study aimed at testing the applicability of using substitutes for leaf area which could be collected in a non-destructive way, namely crown surface area and crown projection area. In 8 stands of Norway spruce (Picea abies L. Karst.), divided into three age classes and two thinning treatments, a total of 156 trees were felled in order to test the relationship between leaf area and crown surface area and crown projection area, respectively. Individual tree leaf area of the felled sample trees was estimated by 3P-branch sampling with an accuracy of ±10%. Crown projection area and crown surface area were compared with other, more commonly used, but destructive predictors of leaf area, namely sapwood area at different heights on the bole. Our investigations confirmed findings of several studies that sapwood area is the most precise measure for leaf area because of the high correlation between sapwood area and the leaf area. But behind sapwood area at crown base and sapwood area at three tenth of the tree height the predictive ability of crown surface area was ranked third and even better than that of sapwood area at breast height (R² = 0.656 compared with 0.600). Within the stands leaf area is proportional to crown surface area. Using the pooled data of all stands a mixed model approach showed that additionally to crown surface area dominant height and diameter at breast height (dbh) improved the leaf area estimates. Thus, taking dominant height and dbh into account, crown surface area can be recommended for estimating the leaf area of individual trees. The resulting model was in line with many other findings on the leaf area and leaf mass relationships with crown size. From the additional influence of dominant height and dbh in the leaf area model we conclude that the used crown model could be improved by estimating the position of the maximum crown width and the crown width at the base of the crown depending on these two variables.     

Effects of wide spacing and thinning strategies on wood quality in Norway spruce (Picea abies) stands in southern Sweden

Abstract

The initial density of a stand influences both the prethinning growth and the quality of the harvested trees. This study investigated the effects of different square spacings (2.0, 2.5 and 3.0 m) with thinning from above and thinning from below on stand development and tree characteristics in a 33-year-old Norway spruce [Picea abies (L.) Karst.] plantation in southern Sweden. Before thinning the total standing volume was not significantly different between treatments. Diameter at breast height (dbh) and diameter of the thickest branch were positively correlated. Before thinning, dbh of the trees and their thickest branches were significantly higher in the stands with 3.0 m spacing. After thinning, the dbh and the diameter of the thickest branch did not significantly differ between the 3.0 m spacing with thinning from above and 2.0 and 2.5 m spacing with thinning from below. Spacing did not significantly influence the presence or size of spike knots. The results suggest that a combination of wide spacing with thinning from above may yield timber of similar quality to denser spacing with thinning from below.     

End grain water absorption and redistribution in slow-grown and fast-grown Norway spruce (Picea abies (L.) Karst.) heartwood and sapwood

Abstract

Wood is susceptible to decay by rot fungi if it is exposed to high-moisture contents during long periods of time and it is therefore important to limit the duration of such periods. Critical points in outdoor wood structures are, for example, end grain surfaces in joints where water can get trapped after a rain. It is therefore of interest to study both absorption and redistribution of moisture in wood. This paper presents moisture content profiles during end grain water absorption and redistribution in Norway spruce (Picea abies (L.) Karst.) measured by computed tomography with the specimens in individual climate boxes. Heartwood and sapwood of two provenances (slow-grown and fast-grown wood) were included. No major differences were seen between the water uptake of the slow-grown and the fast-grown wood since the densities were similar despite of the large difference in growth ring width. However, for the sapwood specimens, the moisture content was higher further into the specimens than for the heartwood specimens in agreement with previous studies. For the slow-grown wood, the redistribution was also generally more rapid for the sapwood specimens than for the heartwood specimens.     

Lophodermium piceae and Rhizosphaera kalkhoffii in fallen needles of Norway spruce (Picea abies)

Falling needles collected from under individual 70-year-old Norway spruce (Picea abies) trees for about four years were checked for fungal fruitbodies. The most common fungi were Lophodermium piceae and Rhizosphaera kalkhoffii. Maximum frequencies of fruitbodies occurred in late autumn for L. piceae (ca. 80% of the needles), in summer for R. kalkhoffii (ca. 60%) and in winter for the third most common fungus, Tiarosporella parca (ca. 10%). The frequencies of needles with fruiting fungi varied greatly within and between years. This is the first report of T. parca from Sweden.     

The influence of magnesium deficiency on carbohydrate concentrations in Norway spruce (Picea abies) needles

Both short- and long-term effects of Mg deficiency on carbohydrate metabolism were investigated in 6-year-old clonal Norway spruce (Picea abies (L.) Karst.) trees cultivated in sand culture with an optimal supply of nutrients, except for Mg which was supplied at 0.203, 0.041 and 0.005 mM to provide optimal, moderately deficient and severely deficient Mg supply, respectively. Annual changes in carbohydrate concentrations (starch, sucrose, glucose and fructose) were analyzed and diurnal changes were investigated on a single day during the summer. Older needles of trees in the moderate Mg-deficiency treatment developed tip-yellowing symptoms, whereas current-year needles remained green. The severe Mg-deficiency treatment led to pronounced yellowing symptoms in needles of all ages. Increased carbohydrate concentrations were observed before needle yellowing occurred. Diurnal and annual changes in carbohydrates were similar in all treatments; however, carbohydrate concentrations were influenced by Mg supply. In both Mg-deficiency treatments, starch concentrations increased in needles, especially during summer and autumn. Starch accumulation was more pronounced at the beginning of the Mg-deficiency treatments than at the end of the treatments. Sucrose, and to a minor extent, glucose and fructose concentrations tended to increase in response to Mg deficiency. The consequences of Mg deficiency on carbohydrate metabolism are discussed with respect to reduced plant growth and decreased transport rates of carbohydrates to sink organs.     

Photosynthesis,nutrient, growth and soil investigations of a declining Norway spruce (Picea abies) stand in the coastal region of Northern Germany

The research site, Wingst Compartment 123B, is a 68-year-old Norway spruce (Picea abies (Karst.)) stand located in the coastal area of northern Germany. This area receives high atmospheric inputs of ammonium and also has relatively high ozone concentrations (0.061 mg m⁻³).Ten trees were categorized as healthy to slightly damaged (3–29% needle loss) or severely damaged (49–71% needle loss). Apparent net photosynthetic rates were measured on detached branches at light saturation (1000 μE m⁻² s⁻¹). Needles were analyzed for chlorophyll, N, C, Ca, Mg, K, Zn, Mn and Fe.When compared to the healthy-to-slightly-damaged trees, the severely damaged trees tended to have higher rates of net apparent photosynthesis in the 1 and 2-year-old needles and similar rates in the current-year needles. All three needle ages from the severely damaged trees had higher average stomatal conductances to water vapor (g_s. Although the damaged trees had significantly less total chlorophyll in all needle ages sampled, there was no statistically significant difference in the chlorophyll a:b ratio between the healthy and severely damaged trees.Nitrogen contents of the current-year needles were slightly lower in the severely damaged trees. Carbon and calcium levels did not significantly differ between the damage classes, although the average Ca content of all younger needles was generally under the recommended sufficiency levels. The current-year needles of the severely damaged trees had significantly less magnesium (0.42–0.46 mg Mg g⁻¹) than those of the healthy trees, and all trees had Mg values in the deficiency range (< 0.7 mg g⁻¹). The severely damaged trees also had lower average potassium levels in the older needles.The annual volume increment per unit crown surface area declined with increasing crown damage. Trees with a 50% needle loss showed a 62% loss of volume increment.Soil investigations revealed conditions of high soil acidity and poor nutrient capacity. The low pH values (pH < 3.8) in 64% of all samples indicated a high risk of acid toxicity for plant roots in the investigated area.The significance of these results relative to the current ideas concerning forest decline is discussed.     

Effect of season and soil treatments on carbohydrate concentrations in Norway spruce (Picea abies) mycorrhizae

We studied effects of season and soil treatments (watering, acidification, liming and combinations of these treatments) on soluble carbohydrates of mycorrhizal roots of Norway spruce (Picea abies (L.) Karst.). Arabinose, arabitol, fructose, glucose, inositol, lactose, mannitol, pinite, quinate, raffinose, shikimate, stachyose and trehalose were identified by HPLC. Concentrations of inositol, lactose and pinite were constant throughout the year, whereas concentrations of raffinose, stachyose and trehalose were higher in winter than in summer, and concentrations of glucose, fructose and mannitol increased from February to September. Soil acidification and liming had no effect on the annual mean concentrations of fructose, glucose, lactose, pinite, raffinose and stachyose. Liming increased quinate concentrations and decreased arabitol concentrations. Annual mean concentrations of arabinose and mannitol decreased in response to soil acidification. Annual mean concentrations of inositol increased in response to irrigation. None of the soil treatments affected the mean annual concentration of trehalose.     

Effects of photoperiod and temperature on the timing of bud burst in Norway spruce (Picea abies)

We examined the effects of several photoperiod and temperature regimes imposed during the winter-spring period on the timing of bud burst in rooted cuttings of Norway spruce (Picea abies (L.) Karst.) grown in a greenhouse in Finland. The treatments were initiated in November and December after the cuttings had been exposed to natural chilling and freezing events. Irrespective of the treatments applied, time to bud burst decreased with increased duration of previous exposure to natural chilling and freezing events. Fluctuating day/night temperatures and continuous lengthening of the photoperiod hastened bud burst. Shortening the photoperiod delayed bud burst, suggesting that little or no ontogenetic development toward bud burst takes place during mild periods before the winter solstice. In the case of climatic warming, this phenomenon may prevent the premature onset of growth that has been predicted by computer simulations with models that only consider temperature regulation of bud burst.     

Seasonal variation in photochemical activity and hardiness in clones of Norway spruce (Picea abies)

We investigated changes in photochemical activity and cold hardiness of detached needles of three clones of Picea abies (L.) Karst. by measuring variable chlorophyll fluorescence (F(v)/F(m)), before and after artificial freezing, from September to June. Photochemical activity varied considerably during the study, but only minor differences in photochemical activity among the clones were observed before freezing. Photochemical activity was high during early fall and then declined from November until April. Photochemical activity was at a minimum in April and then increased quickly to high values in May. During the period from late September to October, and also during the winter, differences in F(v)/F(m) ratios after artificial freezing to below -10 degrees C were observed among clones, indicating clonal differences in cold hardiness and hardiness development. The clone having an average height of 2.3 m after 11 years showed consistently lower cold hardiness than clones that had reached average heights of 4.0 and 5.0 m. There were also differences in the temperature requirement for bud flushing among clones.     

Vertical and horizontal water redistribution in Norway spruce (Picea abies) roots in the Moravian Upland

Hydraulic redistribution (HR) by roots of large Norway spruce (Picea abies (L.) Karst.) trees was investigated by means of sap flow measurements made with the heat field deformation method. Irrigation was applied to a limited portion of the root system to steepen gradients of water potential in the soil and thus enhance rates of HR. On completion of the sap flow measurements, and to aid in their interpretation, the structure of the root system of seven of the investigated trees was exposed to a depth of 30 cm with a supersonic air-stream (air-spade). Before irrigation, vertical redistribution of water was observed in large coarse roots and some adjacent small lateral roots. Immediately after localized irrigation, horizontal redistribution of water from watered roots to dry roots via the stem base was demonstrated. The amount of horizontal distribution depended on the position of the receiving roots relative to the watered roots and the absorbing area of the watered root. No redistribution from watered roots via dry soil to roots of neighboring trees was detected. Responses of sap flow to localized irrigation were more pronounced in small lateral roots than in large branching roots where release and uptake of water are integrated. Sap flow measurements with multi-point sensors along radii in large lateral roots demonstrated water extraction from different soil horizons. We conclude that synchronous measurements of sap flow in both small and large lateral roots are needed to study water absorption and transport in tree root systems.     

Variation in wood properties among five full-sib families of Norway spruce (Picea abies)

Genetic- and environmental variation and correlation patterns were characterized for modulus of elasticity (MOE), modulus of rupture (MOR) and related wood traits: latewood proportion, wood density, spiral grain, microfibril angle and lignin content in five full-sib families of Norway spruce. The families were evaluated on the basis of clearwood specimens from the juvenile -mature wood transition zone of 93 sampled trees at age 30 year from seed. Family-means varied significantly (p < 0.05) for all wood traits studied except lignin content. MOE varied between 7.9–14.1 GPa among trees and 9.4–11.0 GPa among families. MOR varied between 47–87 MPa among trees and 61–71 MPa among families. Families remained significantly different in an analysis of specific MOE (MOE/density) and MOR (MOR/density). Hence, solely relying on wood density as a wood quality trait in tree breeding would not fully yield the potential genetic gain for MOE and MOR. Correlations between wood structural traits and specific MOE and MOR are presented and discussed.     

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.     

Cupping of wooden cladding boards in cyclic conditions—a study of boards made of Norway spruce (Picea abies) and Scots pine sapwood (Pinus sylvestris)

The aim of the study was to examine how the thickness and species of softwood influence cupping during cyclic conditions. The study was conducted with full-scale test walls in laboratory conditions. On the basis of the results, the thickness and species of softwood both have a significant influence on cupping of the cladding board. The boards made of Norway spruce were considerably more resistant to cupping than those made of Scots pine sapwood. Boards made of Scots pine sapwood were more likely to remain curved after each drying period. It appears probable that the cupping sensitivity of Scots pine sapwood affects the durability of the cladding, e.g., causing cracking formation on the surface and peeling of the coating. From the curving results, it is possible to estimate surface elongation and durability of the wood surface and of the coating in cyclic conditions.     

Antagonists of Fomes annosus in the rhizosphere of grey alder (Alnus incana) and Norway spruce (Picea abies)

A comparison was made of the fungistatic effects of the root microflora of grey alder and Norway spruce with Fomes annosus as test fungus. The significantly higher frequency of actinomycetes in the alder rhizosphere probably constituted the main difference in this respect. The importance of fungi seemed to be about equal in the rhizosphere of the two tree species. Methods for studies of antagonism are tested and discussed.     

Crown architecture and stemwood production in Norway spruce (Picea abies (L.) Karst.)

The morphological data for Norway spruce (Picea abies (L.) Karst.) trees published by Burger (1953) were reanalyzed to examine the relationships between crown shape, needle and branch mass density of crown volume, needle efficiency in stemwood production, harvest index and stemwood production efficiency per unit of crown projected area. Production of stemwood per unit of crown projected area was higher in narrow-crowned trees than in broad-crowned trees because the narrow-crowned trees had (i) smaller horizontal space requirements for equal crown volumes, (ii) higher leaf area index due to both the geometrical crown shape and higher needle density per unit of crown volume, and (iii) higher harvest index indicating high allocation of dry matter production to stem.     

Vulnerability of spruce (Picea abies) and beech (Fagus sylvatica) forest stands to storms and consequences for silviculture

The form and magnitude of storm damage and stand disclosure patterns were assessed in 332 randomly chosen pure and regular stands of spruce (Picea abies L.) and beech (Fagus sylvatica L.) after storm lothar, within a region of the Swiss Midlands. This data was analysed in relation to maximal wind speed, measured with Doppler radar techniques and other influential factors such as relief, allometric characteristics, silvicultural history, and neighbourhood. In addition, storm damage, assessed from aerial photographs over an extended perimeter (about 70,000 ha) was considered. A storm of the magnitude of lothar (December 26 1999), with an average maximal wind speed of 45 m s⁻¹ (160 km h⁻¹) appears to have a highly chaotic wind field structure, with great spatial and temporal variation of wind gusts. Wind speeds were not a significant predictor for damage in spruce stands and only weakly influential for beech. The consequences of this high randomness were analysed to estimate the return time of such a storm at the stand level. It lies between 86 and 113 years for spruce, 357 and 408 for beech. Only a few independent variables were significant and the overall explanatory strength of the model was unexpectedly low (R ²=0.07 for spruce and 0.30 for beech). Among the more reliable predisposing factors were mixture and aspect combined with gradient. An admixture of 10% or more broadleaved tree species or wind-firm conifers like Douglas fir [Pseudotsuga menziesii (Mirb.) Franco] significantly reduced the vulnerability of spruce stands (by a factor of more than three). On wind-exposed aspects, damage was more than twice the average. Steeper slopes caused a significant reduction in susceptibility (by a factor of six for slopes over 50%, in comparison to gentle slopes <20%). Other factors such as height to diameter ratio of trees or time since last thinning did not appear to be significant predictors.     

Bioenergy production and soil conservation from Norway spruce (Picea abies L. Karst) plantations in Denmark

This study analyses the trade-off between bioenergy production and soil conservation through thinning operations in Norway spruce (Picea abies L. Karst) plantations in Denmark. Thinning operations were evaluated under different regimes and intensities for a complete rotation period of sixty years and for different site qualities (site-classes I–VI). Applying a dynamic forest growth modeling tool, evolution of forest structure was predicted to observe the potentials for biomass production and inevitable soil degradation. Results showed thinning from below, with a higher utilization (maintenance of a minimum basal area of 25 m² ha^?1) could produce more bioenergy. However, these operations require simultaneous severe forest soil degradation. Therefore, the optimum thinning for bioenergy production under preservation constraints was thinning from above with a lower intensity (maintenance of a minimum basal area of 45 m² ha^?1). The ratio of bioenergy win (kWh) to soil-loss (m³ ha^?1) was calculated for this regime varying between 74,894 kWh m^?3 in a high quality site (site-class I) and 6,516 kWh m^?3 in a low quality site (site-class VI) with an average of 44,282 kWh m^?3. However, this could not always preserve the highest amount of growing stock essential for natural dynamics of forest ecosystem with an exception of the low quality sites (site-class VI). Thus, when aiming at bioenergy production through thinning operations, trade-offs with soil conservation and growing stock preservation should be regarded to prevent environmental degradation.     

Storage lipid dynamics in somatic embryos of Norway spruce (Picea abies): histochemical and quantitative analyses

Adequate storage compounds are a prerequisite for successful development during the later stages of somatic embryogenesis; however, the critical amount of reserves below which somatic embryos fail to mature and germinate has not been determined. We analyzed storage lipids during Norway spruce (Picea abies (L.) Karst.) somatic embryogenesis. As maturation progressed, lipids, which were stored as lipid bodies in the cytoplasm, were localized first in suspensor cells of the early embryos, and later in the embryonic root pole, superficial layers of the hypocotyl and in cotyledons. The concentration of total lipids exhibited marked variation, with values peaking during cotyledon development and then decreasing during maturation. Linoleic (18:2), oleic (18:1), palmitic (16:0) and 5,9-octadecenic (5,9-18:2) acids were the most abundant fatty acids in embryos. As embryos developed, linoleic acid concentration increased slightly, whereas oleic acid concentration decreased. Oleic acid was the most prominent component of the fatty acid spectrum in isolated dormant zygotic embryos and megagametophytes. Addition of 5% polyethylene glycol to the medium during somatic embryo maturation caused a shift in the fatty acid spectrum toward that of zygotic embryos. During maturation, changes in the exogenous carbohydrate supply had no significant effect on total lipid concentration in mature embryos. A marked decrease in lipid concentration was detected during desiccation, indicating the importance of adequate lipid reserves during this developmental stage. The lipid content of zygotic embryos differed considerably with harvest year and location, suggesting that zygotic embryo data cannot be an indicator of somatic embryo quality.