Development of biomass proportions in Norway spruce (Picea abies [L.] Karst.)

This study tests the hypotheses that (1) the above-ground structure of Norway spruce (Picea abies [L] Karst.) is derivable from the functional balance theory, and that (2) crown ratio is a key source of structural variation in trees of different age and social position. Twenty-nine trees were measured in three stands (young, middle-aged, and mature), with three thinning treatments (unthinned, normal, and intensive) in the two older stands. There was a strong linear relationship between the total cross-sectional area of branches and that of stem at crown base. Foliage mass was linearly related with stem basal area at crown base. Also an allometric relationship was found between foliage mass and crown length. The mean length (weighted by basal area) of branches obeyed an exponential function of crown length. The parameters of most of these relationships were independent of slenderness (tree height/breast height diameter) and tree age However, total branch cross-sectional area per stem cross-sectional area in the young trees was greater than in the older trees. The young trees also had slightly shorter branches than predicted by the mean branch length equation. This was probably caused by branch senescence which had not yet started in the young stand. The older trees had a relatively long lower crown segment which was growing slowly and senescing. It was proposed that a segmented crown structure is characteristic of shade tolerant tree species, and that the structural model could be further developed by making the two segments explicit.     

Seasonal accumulation of ultraviolet-B screening pigments in needles of Norway spruce (Picea abies (L.) Karst.)

Conifer needles are highly effective in screening ultraviolet-B radiation (280–320 nm). This ability is mainly attributed to the presence of flavonoids and hydroxycinnamic acids in the epidermal tissue. In two field cabinet experiments with two different clones of Norway spruce we assessed the seasonal accumulation of UV-B screening pigments under near-ambient, and close-to-zero UV-B irradiation. At the beginning of needle development, i.e. in June, kaempferol 3- O -glucoside was the dominant UV-B screening pigment. It was replaced during needle differentiation by the more effective diacylated flavonol glucosides, particulary kaempferol 3- O -(3",6"- O -di- p -coumaroyl)-glucoside, which reached highest concentrations in July. In addition to the soluble pool of diacylated flavonol glucoside derivatives, a cell wall-bound UV-B screen in the epidermal cell walls was formed during needle differentiation, consisting mainly of p -coumaric acid and kaempferol 3- O -glucoside. An effect of UV-B radiation on the accumulation of diacylated flavonol glucosides was only observed in 1996 with clone 2, when the concentrations of kaempferol 3- O -(3",6"- O -di- p -coumaroyl)-glucoside were significantly higher in July and August under field, and near-ambient than under close-to-zero UV-B irradiance. For wall-bound p -coumaric acid and kaempferol 3- O -glucoside UV-B radiation enhanced the concentrations of these compounds by approximately 20% in relation to the concentrations in close-to-zero UV-B-treated plants in both field cabinet experiments.     

Influence of cultivation method on root grafting in Norway spruce (Picea abies (L.) Karst.)

Root grafting is the process by which a functional union of two or more roots subsequent to their formation is formed. The above- and below-ground parts of three Norway spruce stands (natural stand, Umbric Luvisol; row-culture and group-culture, Planosol; stand ages 40, 43 and 43, respectively) of high site quality (I) were investigated. Stand densities were 1550, 1783 and 1722 stems ha^-1, respectively. In all investigated stands, root grafting was most sensitive to tree spacing. Grafts were observed in case the distance between the trees was 0.7–1.2 m. Grafts occurred always in areas of higher rooting density, in a row of the row-culture and within a tree group in the group-culture. Root grafting was enhanced in case of a narrower humus horizon in the group culture compared with the row-culture, 16.5 and 30 cm, respectively; the humus horizon contained 99% and 95% of conducting roots with d ≥ 5 mm, respectively. Root graftings occurred in 75% of excavated trees in the group-culture, in 37.5% of excavated trees in the row-culture and in 33.3% of excavated trees in the natural stand. Stand age was 24 years in the row-culture and 22 years in the group-culture at the beginning of root grafting. No grafts occurred between two suppressed trees, whereas in 86–100% of all cases, at least one tree was dominant or codominant. In row- and group-cultivated Norway spruce stands, the initial minimum diameter of the grafted root without bark was from 1 to 3 cm in 63% of cases. Grafting of roots with d < 1 cm or d > 10 cm was rare or absent. Root grafting had usually begun at the root age of 10–20 years (46% of cases). This revised version was published online in June 2006 with corrections to the Cover Date.     

Soil changes in different age classes of Norway spruce (Picea abies (L.) Karst.) on afforested farmland

The aim of this study was to test the hypothesis that afforestation changes the content and distribution of soil organic carbon, nutrients and pH in the A-horizon of land previously used in agriculture, and that such soil changes depend on stand development. The investigation was evaluated as a completely randomised design with three treatments representing different age classes of trees: 20 years (Y20), 40 years (Y40) and 55 years (Y55). Eighteen trial plots, six per treatment, were established in plantations of Picea abies (L.) Karst. on soils of similar texture and mineralogy. Tree volume was 220 m³ ha^-1 in Y20, 400 in Y40 and 440 m³ ha^-1 in Y55.Concentrations of carbon (C) and nitrogen (N) were significantly higher in the uppermost part of the soil in the older stands Y40 and Y55 than in Y20. The total amount of organic C in the litter layer plus the top 15 cm of the soil differed between age classes, with Y40 and Y55 having the largest amounts. A reference layer (15–20 cm) was used in calculating the amount of soil C that had accumulated in the horizon since afforestation, being about 10 tonnes ha^-1 of C in Y20 and 19 tonnes ha^-1 in Y40 and Y55.Cation exchange capacity (CEC) and base saturation (BS) was higher in the older stands. Carbon contents and CEC were strongly correlated. In Y40 and Y55, pH was significantly lower than in Y20 in the lower part of the soil horizon. There was a general decrease with depth of C, N, CEC, K⁺ and Mg²⁺ in the soil horizon. BS, Ca²⁺, Na⁺ and pH showed a somewhat different pattern of distribution, with deceasing values in the upper part of the soil horizon and increasing values in the lower part of the soil horizon.Abbreviations BD Bulk density - CEC cation exchange capacity - BS base saturation - Ca²⁺ calcium ion - Mg²⁺ magnesium ion - K⁺ potassium ion - Na⁺ sodium ion - C carbon - Ca _a accumulated carbon content - C _t total carbon content - N nitrogen - Y20 age class 20 years - Y40 age class 40 years - Y55 age class 55 years     

Root distribution in a Norway spruce (Picea abies (L.) Karst.) stand subjected to drought and ammonium-sulphate application

Results of the spatial distribution of fine roots are reported from a Norway spruce (Picea abies (L.) Karst.) in SW Sweden stand subjected to drought (D) and ammonium-sulphate application (NS). The sampling was carried out by excavating monoliths in segments of 0.5 × 0.5 × 0.1 m to a depth of one meter. Root data also included in the study were obtained by excavating whole trees and soil coring.The data suggest a fairly deep distribution pattern of fine roots (< 1 mm in diameter) in the study area compared with other forest sites in SW Sweden. The weight fraction of living fine roots in the LFH-horizon amounted to 53, 36 and 55% of the total fine-root biomass and 12, 30 and 32% of the total fine-root necromass (dead fine roots) in the control, D and NS-treatment areas respectively. Drought seemed to result in a redistribution of fine roots to deeper mineral soil horizons. Ammonium sulphate application led to the reverse, viz, a concentration of fine roots to the LFH-horizon. A significantly smaller fine-root necromass was indicated in the LFH-horizon of the control areas compared with both the D and NS-treatment areas, suggesting a high mortality of fine roots in these areas. A heavy dry matter fraction accumulates in roots > 1 mm in diameter and in stumps. These root fraction, were frequently found between the trees, although the stump constitutes an important fraction in terms of dry weight.     

Decomposition and nitrogen dynamics of fine roots of Norway spruce (Picea abies (L.) Karst.) at different sites

Long-term decomposition and nitrogen dynamics of Norway spruce finest (<1 mm in diameter) and fine (<2 mm in diameter) roots were estimated using the root litter-bag techniques. The seasonal decomposition of the finest roots was investigated in a 40-year-old high site quality stand grown on brown lessive soil at different depths as part of productivity studies. The fine root decomposition studies were conducted on 8 permanent plots in the Estonia with the aim to describe the site variation. The initial material was collected from one of stands (high quality site) and incubated at the depth of 10 cm in 1989 (at one site 1990). The bags were collected once or twice a year except for one site, where the seasonal dynamics was investigated. In all initial and decomposing root samples oven-dry weight, ash and energy content and nitrogen concentration was determined. After five years the finest roots had lost 40% of their initial dry weight, half of it during the first year. The initial concentration of nitrogen was 1.29%, the mean concentrations varied during the incubation from 1.47 to 1.78%. After the first year fine roots had lost 21.0 to 32.7% of their initial dry weight, after two years the weight loss was 22.5 to 43.2%. The initial N concentration in fine roots was 0.73% and in the first years it varied from 0.97 to 1.40% at different sites.     

Nitrogen and drought effects on ectomycorrhizae of Norway spruce [Picea abies L.(Karst.)]

Effects of N addition and drought on ectomycorrhizae of Norway spruce trees were investigated in an outdoor pot lysimeter study. Three levels of N were applied as ammonium nitrate in irrigation water for five years; ambient rainwater (N0) and 5 (N5) and 15 (N15) times this N concentration. Mean annual N addition during the five years corresponded to 5, 27 and 82 kg·ha^-1·y^-1 for the N0, N5 and N15 treatments, respectively. During the third and fifth growth seasons two levels (lengths) of drought were artificially induced in addition to a watered control. Soil cores taken from each pot lysimeter were analyzed for mycorrhizal colonization and ectomycorrhizae were categorized according to macroscopic morphology. Drought decreased mycorrhizal colonization significantly. There was a significant interaction of drought and N effects on reduction of the mycorrhizal colonization. Although all of the mycorrhiza types were influenced by drought, only Cenococcum geophilum showed a significant change. N treatment alone did not show any significant effect either on mycorrhizal colonization or mycorrhizal types.     

Concentrations of nutritional and trace elements in needles of Norway spruce (Picea abies [L.] Karst.) as functions of the needle age class

The endogenous concentrations of the essential elements Ca, Cl, Cu, Fe, K, Mg, Mn, N, P and Zn, and of the nonessential elements Al, As, Ba, Br, Co, Cr, Cs, Hg, I, La, Na, Rb, Sb, Sc, Si and Sr were determined in 5 successive needle age classes. 40 mature spruce trees from 6 different sites were investigated individually. A given element usually shows smooth changes with the needle age class t. Trees on a given site usually have a similar dynamic behaviour. The same holds for the different site means. The concentrations can be approximated by functions c=f(t). Three different types of functions are required to describe the dynamic behaviour of 3 groups of elements that increase with t, and one for the elements that decrease with t. A given element usually can be described by the same type of function at all sites, even if its concentration differs widely. Exceptions are Mn, Co and Zn, which change from a decreasing function at low concentrations to an increasing function at high concentrations. Further irregulatities are found at some sites with Ca, Sr and Ba. These findings are corroborated by a multivariate statistical analysis.     

In-situ soil solution chemistry in an acid forest soil as influenced by growing roots of Norway spruce (Picea abies [L.] Karst.)

A new approach for non-destructive monitoring of soil solution chemistry in high spatial and temporal resolution for rhizosphere studies is presented. In a 5×10 mm grid, 30 micro suction cups (1mm) were installed in a rhizotron with Norway spruce (Picea abies [L.] Karst.) growing in low pH B-horizon soil. Roots grew through the grid, closely passing the suction cups. Soil solution composition before, during and after root passage was determined. For K⁺ and Mg²⁺ a significant decrease of soil solution concentration near root tips and elongation zones was observed, indicating a marked uptake of these elements. Mg²⁺ concentration was also significantly lowered when the root system aged, suggesting that this ion might also be taken up in older parts of the root system. No influence of growing roots was found on Na⁺-concentrations.     

Seasonal development of the photosynthetic performance of Norway spruce (Picea abies [L.] Karst.) under magnesium deficiency

In order to investigate the influence of different magnesium nutrition on photosynthesis, one hundred 6-year-old spruce trees derived from one clone were planted in October 1990 into a special out-door experimental construction, where they were cultivated in sand culture with an optimal supply of nutrients, except magnesium, via circulating nutrient solutions. Magnesium was added to the nutrient solutions in three different concentrations, varying from optimal to severe deficient supplies. During the first vegetative period in 1991, photosynthetic performance and carboxylation efficiency were measured under saturating light, controlled CO₂ conditions, optimal temperature and humidity, using a minicuvette system.During summer, the trees under moderate magnesium deficiency developed tip yellowing symptoms on older needles, while the youngest needles remained green with unchanged chlorophyll contents. Trees under severe magnesium deficiency showed yellowing symptoms on all needle age classes combined with decreased chlorophyll contents in the youngest needles as well. In comparison with the controls, the photosynthetic performance of the 1-year-old needles was significantly lower in both deficiency treatments. The same was observed in the youngest needles of the trees under severe deficiency. Trees under moderate deficiency treatment decreased in photosynthetic performance during the summer without reduction of chlorophyll contents. The reduction of photosynthetic rates corresponded to a decrease in carboxylation efficiency, which is taken as a measure of the activity of the enzyme ribulose-1,5-bisphosphate carboxylase. This reduction, together with the observed increase of carbohydrate contents in needles of trees growing under magnesium deficiency, led to the assumption that the photosynthetic carbonfixation is reduced as a consequence of the accumulation of carbohydrates.     

Identification of a RAPD marker linked to the pendula gene in Norway spruce (Picea abies (L.) Karst. f. pendula)

The pendula phenotype of Norway spruce [Picea abies (L.) Karst f. pendula] is characterized by narrow crowns and strong apical dominance and is controlled by a single dominant gene (P). This defined genetic control presents one of the few opportunities to map a single gene controlling a morphological trait in a forest tree. We used random amplified polymorphic DNA (RAPD) markers and bulked segregant analysis to identify one locus OPH10_720, linked to the pendula gene. The estimated recombination frequency (r) between OPH10_720 and P was 0.046 (SE _r =0.032). Mapping of the pendula gene is an important first step towards the ultimate identification and cloning of this gene.     

Enhancement of somatic embryogenesis in Norway spruce (Picea abies L.)

Summary Embryogenic callus developed in 55% of the mature embryo explants of Norway spruce (Picea abies L.) growing on a LP medium minus the amino acids and sugars (except sucrose). This is the highest reported yield of embryogenic callus from mature embryos of P. abies that has ever been reported. Callus induction from either the middle or the end of the hypocotyl of the embryos began after 2–3 weeks. Three types of calli were recovered: (a) globular, (b) light green-compact, (c) white mucilaginous. Only the white mucilaginous calli were embryogenic. The globular and light green-compact calli never become embryogenic, even after several subcultures. The development of somatic embryos was accomplished on half-strength macro-elements of NSIII medium containing 1 M -naphthaleneacetic acid, 1 M abscisic acid, and 3% sucrose. The addition of 10^–7 M buthionine sulfoximine to the medium increased the development of somatic embryos by three fold. These results suggest that there is a great potential for increasing the frequency and development of somatic embryos in P. abies. Careful selection of the genotype and modification of the culture medium is required.     

Structure and stand development in three subalpine Norway spruce (Picea abies (L.) Karst.) stands in Paneveggio (Trento, Italy)

1. Three permanent plots (100×0 m) were established in the subalpine Norway spruce (Picea abies (L.) Karst.) forest of Paneveggio in the spring of 1993, to begin a long-term forest ecosystem research project. The main purpose of these plots was to provide information about subalpine Norway spruce stand dynamics and to provide suggestions for close-to-nature silviculture. 2. The three stands were selected to represent the most common forest structures in the Paneveggio forest. The first stand is close to forestry roads, has a relatively regular and continuous canopy, and thinning and cutting operations only ended in the 1980s; the second stand is far from forest roads and has developed without anthropogenic influence for several decades; the third one is located at the present upper limit of the pure spruce forest and, apparently, was heavily used in the past as a pasture. 3. The first step in the investigation was to describe the structure and to study the history of the three stands using both written evidence from manage- ment plans and biological archives from tree rings. 4. The stands in plots 1 and 2 began to establish after a disturbance that removed part of the previous stands according to dendroecological studies, which are partially supported by written evidence. The remaining parts of these stands were eliminated by two major disturbances that occurred during the following decades. Written records about the use of the forest lead us to assume that the initial disturbances that occurred in the two stands were logging activities as a part of a group shelterwood system. The stand in plot 2 has developed without significant human interference for about half a century as confirmed by the presence of many dead trees. The stand in plot 3 consists of old trees that were part of an open stand and a secondary population that established after cessation of grazing. 5. The study has confirmed that dendroecological techniques can be used to identify occurrence and intensity of previous disturbance in forests stands, although at Paneveggio it is difficult to distinguish between natural and anthropogenic disturbances in the tree ring record. The presence of human activity necessitates investigation of multiple lines of evidence. 6. Paneveggio's forest management plans were useful in the interpretation of the data obtained through dendroecological analysis, although events did not always correspond because data from the management plans (yearly thinning, felling, wind-throw damage) never gave stand-level details, but applied to areas of several hectares. Despite these limitations, the information included in the management plans is of crucial importance in studying stand history and only by using all these sources of information is it possible to delineate the most important features of the history and disturbance that affected the origin and subsequent growth of the forest stands.     

Growth of adult Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica L.) under free-air ozone fumigation

This study attempted to detect the impact of ozone on adult trees of Norway spruce ( Picea abies [L.] Karst.) and European beech ( Fagus sylvatica L.) in an experimental mixed stand in Southern Bavaria, Germany. The aim was to examine whether there is a decrease in growth when trees are exposed to higher than atmospheric concentrations of ozone. This exposure was put into effect using a free-air fumigation system at tree crown level. Growth analysis was carried out on a group of 47 spruce and 36 beech trees, where radial stem increment at breast height - a sensitive index for stress - was measured. The ozone monitoring system allowed values to be obtained for the accumulated ozone exposure (SUM00) of each individual tree, so that their radial increment over three years could be correlated with the corresponding ozone exposure for the same time period. Correlation and regression analysis were then carried out to test the influence of ozone on diameter increment. In both spruce and beech, the initial stem diameter was the most influential factor on radial increment in the following year. A linear model was applied, including the diameter of the preceding year and the ozone exposure of the current year as predicting factors. For spruce trees, a significant negative influence of ozone exposure was found. In contrast, no significant ozone effect on diameter increment of beech was detected. The effect of ozone stress on a large spruce tree can lead to a decrease in potential radial increment of 22 %. The results are discussed in relation to other stress factors such as drought and lack of light.     

Influence of UV-B radiation on photosynthetic activity and chlorophyll fluorescence kinetics in Norway spruce [Picea abies (L.) Karst.] seedlings

Norway spruce [Picea abies (L.) Karst.] seedlings were grown in greenhouses with two supplemental levels of ultraviolet-B (UV-B) radiation. Photochemical efficiency of photosystem II and vitality index were determined monthly. At the end of the experiment, growth, chlorophyll content and photosynthetic rates were measured. The data indicate that low temperature in winter affected light dependent processes in experimental plants including control, while the rise of ambient temperatures, moderate this effect. The synergistic effects of UV-B radiation and low temperatures could only be observed in the second winter period. Measurements of net photosynthetic activity in the second winter period showed significant differences between treated and untreated plants.