Bacteria inhabiting artificially inoculated xylem of Picea abies

Bacteria inhabiting the xylem of Norway spruce (Picea abies (L.) H. Karsten) were investigated. The trees had been wounded and artificially inoculated with fungi and bacteria obtained from wounds of naturally infected spruce. One and five growing seasons after inoculation the Gram‐negative bacterial population present in the stem of inoculated trees were analysed.

The Gram‐negative bacteria isolated from the trees were identified on the basis of morphological, biochemical and physiological tests and whole‐cell fatty acid composition. The predominant strains were Enterobacteriaceae fermenter strains (E. agglomerans or E. sakazakii), fluorescent and yellow pigmented Pseudomonas, Acinetobacter and Moraxella spp. All Gram‐positive bacteria were Bacillus species.

The Gram‐negative bacteria of Norway spruce differed from the Gram‐positive species in possessing stronger lipolytic activity and in their ability to utilize pine resins for growth. Gram‐positive bacteria were generally able to utilise cellulose and hemicellulose, whereas among the Gram‐negative bacteria only one xylanolytic (yellow Pseudomonas) strain was found.     

Basic density,longitudinal shrinkage and tracheid length of juvenile wood of Picea abies (L.) Karst

Variation in the growth ring width, basic density, longitudinal shrinkage and tracheid length was investigated in the juvenile wood of Norway spruce samples taken from different heights in the stem. Annual height increments were cut from the ten youngest shoots from trees of three different heights. By this method the properties of an individual growth ring could be analysed without taking samples from each ring. Sixteen tree tops of an average stem height of 8, 15 and 25 m were analysed. Basic density was low in the first shoot, highest in the second or third one and decreased gradually thereafter. Longitudinal shrinkage was highest close to the pith and decreased to about 0.2% in the outer rings. Tracheids were only ca. 1 mm long close to the pith and their length increased sharply towards the tenth shoot. The properties of juvenile wood varied with the height in the stem. Longitudinal shrinkage around the pith seemed to increase with increasing height and basic density was highest at 25 m stem height. The applicability of the method for the calculation of basic density and tracheid length in individual growth rings close to the pith is discussed.     

Antifungal properties of terpenoids in Picea abies against Heterobasidion parviporum

The antifungal properties of 14 major oleoresin‐constitutive terpenoids of Norway spruce (Picea abies) against Heterobasidion parviporum were evaluated in vitro at three gradient concentrations, 0.1, 0.2 and 0.4 μmol/cm², on potato dextrose agar medium. Eight monoterpene hydrocarbons (+)‐ and (?)‐α‐pinene, (?)‐β‐pinene, (+)‐3‐carene, myrcene, (+)‐ and (?)‐limonene, terpinolene; four oxygenated monoterpenes α‐terpineol, terpinen‐4‐ol, 1,8‐cineole, bornyl acetate; and two resin acids abietic acid and dehydroabietic acid were selected. Abietic and dehydroabietic acids showed the highest antifungal activities at all concentrations. Among oxygenated monoterpenes, bornyl acetate and α‐terpineol showed antifungal activity at the highest concentration. Among monoterpene hydrocarbons, (+)‐α‐pinene showed similar activity to terpinen‐4‐ol and 1,8‐cineole at the highest concentration; however, it was lower than α‐terpineol and bornyl acetate. Other monoterpene hydrocarbons inhibited mycelial growth by <10%. Re‐extraction of medium surfaces after the test period revealed that most α‐terpineol and terpinen‐4‐ol remained unchanged on the surface but monoterpene hydrocarbons completely disappeared. However, notable fungal transformed products were observed on surfaces applied with 1,8‐cineole and bornyl acetate. Thus, mycelial growth inhibition of monoterpenoids might be caused by complex functions such as biodegradation and/or detoxification by interaction between mycelium and compound. These results provide a basis for future studies considering the role of chemodiversity in the comprehensive chemical defence of P. abies.     

Organic and inorganic sulfur transport in the xylem sap and the sulfur budget of Picea abies trees

Temporal changes in inorganic and organic sulfur compounds (sulfate, glutathione, cysteine, methionine) were analyzed in xylem sap of 40-year-old Norway spruce (Picea abies (L.) Karst.) trees growing on acidic soils at a healthy and a declining stand in the Fichtelgebirge (North Bavaria, Germany). Studies were carried out (1) to quantify glutathione (GSH) transport in the xylem of spruce, (2) to study the significance of reduced sulfur versus sulfate (SO(4) (2-)) transport in the xylem, and (3) to compare total sulfur (S) transport in the xylem with the amount of foliar uptake of SO(2) in an air-polluted environment. Glutathione was the main reduced S compound in the xylem ranging in concentration from 0.5 to 5 &mgr;mol l(-1). Concentrations of inorganic SO(4) (2-) in the xylem sap were up to 50 times higher than those of GSH ranging from 60 to 230 &mgr;mol l(-1). During the growing season, concentrations of all S compounds in the xylem were highest in May (up to 246 &mgr;mol l(-1)) and decreased during summer and fall (up to 21 &mgr;mol l(-1)). On average, SO(4) (2-) concentrations in xylem sap were 30% higher at the declining site compared with the healthy site. Diurnal changes in organic S compounds were significant for GSH and cysteine with high concentrations during the night and low concentrations during the day. Diurnal changes in inorganic concentrations were not significant. Xylem sap concentrations of SO(4) (2-) and cysteine were twice as high and GSH concentrations were tenfold higher in surface roots than in branches. At both sites, transport of organic S was low (up to 3% of total S) compared to transport of SO(4) (2-). Annual transport of total S in the xylem (SO(4) (2-) was the main component) ranged from 60 to 197 mmol tree(-1) year(-1) at the healthy site and from 123 to 239 mmol tree(-1) year(-1) at the declining site. Although gaseous uptake of SO(2) was estimated to be similar at both sites (38 mmol tree(-1) year(-1); Horn et al. 1989), the ratio between annual gaseous uptake of SO(2) and transport of S in the xylem was 1:4 and 1:5 at the healthy and declining sites, respectively.     

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.     

Water content and bark thickness of Norway spruce (Picea abies) stems: phloem water capacitance and xylem sap flow

To determine the relationship between phloem transport and changes in phloem water content, we measured temporal and spatial variations in water content and sucrose, glucose and fructose concentrations in phloem samples and phloem exudates of 70- and 30-year-old Norway spruce trees (Picea abies (L.) Karst.). Large temporal and spatial variations in phloem water content (1.4-2.6 mg mg(dw)(-1)) and phloem total sugar concentration (31-70 mg g(dw)(-1)) paralleled each other (r(2) = 0.83, P < 0.0001 for the temporal profile and r(2) = 0.96, P < 0.008 for the spatial profile), indicating that phloem water content depends on the total amount of sugar to be transferred. Changes in phloem water content were unrelated to changes in bark thickness. Maximum changes in phloem water content calculated from dendrometer readings were only 8-11% of the maximum measured changes in phloem water content, indicating that reversible changes in bark thickness did not reflect changes in internal water relations. We also studied the relationship between xylem sap velocity and changes in bark thickness in 70-year-old trees during summer 1999 and winter 1999-2000. Sap flow occurred sporadically throughout the winter, but there was no relationship between bark shrinkage or swelling and sap velocity. In winter, mean daily xylem sap velocity was significantly correlated with mean daily vapor pressure deficit and air temperature (P < 0.0001, in both cases). Changes in bark thickness corresponded with both short- and long-term changes in relative humidity, in both winter and summer. Under controlled conditions at > 0 degrees C, changes in relative humidity alone caused changes in thickness of boiled bark samples. Because living bark of Norway spruce trees contains large areas with crushed and dead sieve cell zones-up to 24% of the bark is air-filled space-we suggest that this space can compensate for volume changes in living phloem cells independently of total tissue water content. We conclude that changes in bark thickness are not indicative of changes in either phloem water capacitance or xylem sap flow.     

Microflora of sound-looking wood in Picea abies stems

Ascocoryne spp. (A. cylichnium and A. cf. sarcoides) were found in 48 % of 262 60-year-old Picea abies stems. They were isolated with increasing frequency towards the pith from the lower 2-3 m, more often from thicker than from thinner stems. Neobulgaria premnophila was present in 10 % of the stems, more evenly distributed, and with no difference in occurrence between thicker and thinner stems. Nectria fuckeliana was situated at random in 8 % of the trees.     

Radial mechanical properties of high-temperature dried Norway spruce (Picea abies) wood

Abstract

Wood drying experiments were conducted in which the temperature and the drying rate were controlled independently. The mechanical properties of dried wood in radial loading were analysed in relation to drying parameters. Mass loss, due to thermal degradation of the wood structural components, occurred predominantly in slow high-temperature drying processes. Despite the higher mass loss, slowly dried wood showed similar radial strength and stiffness to rapidly dried wood. The formation of irreversible hydrogen bonds (hornification) within the wood structure may compete with the effects of mass loss on the radial mechanical behaviour of wood. However, both the mass loss and the hornification resulted in wood specimens with lower hygroscopicity. Application of slow high-temperature drying to reduce microscopic cell-wall damage, caused by anisotropic shrinkage of cell-wall layers, did not seem to affect the radial mechanical properties of wood. The effects of stress relaxation within the wood cell wall on the mechanical behaviour of wood may be offset by the degradation of structural components along with drying. Radial mechanical properties may be improved by rapid high-temperature drying up to high final dryness.     

Monolignol oxidation by xylem peroxidase isoforms of Norway spruce (Picea abies) and silver birch (Betula pendula)

We partially purified peroxidase isoform fractions from xylem extracts of a gymnosperm, Norway spruce (Picea abies (L.) Karst.), and an angiosperm, silver birch (Betula pendula Roth.), to determine the participation of xylem-localized peroxidases in polymerization of different types of lignin in vivo. Several peroxidase fractions varying in isoelectric point values from acidic to basic were tested for their ability to catalyze the oxidation of the monolignols coniferyl alcohol, sinapyl alcohol and p-coumaryl alcohol in vitro. All of the xylem peroxidases extracted from Norway spruce and most of those from silver birch showed the highest rate of oxidation with coniferyl alcohol in the presence of hydrogen peroxide. The exception was an acidic peroxidase fraction (pI 3.60-3.65) from silver birch that exhibited higher oxidation activity for sinapyl alcohol than for coniferyl alcohol. For the xylem enzyme fractions extracted from silver birch, the ability to oxidize the artificial phenolic substrate syringaldazine coincided with high specific activity for sinapyl alcohol. Therefore, we conclude that the acidic, neutral and basic xylem peroxidases of Norway spruce all function in the synthesis of guaiacyl-type lignin, whereas in silver birch the acidic peroxidases preferentially oxidize sinapyl subunits. The latter provides a mechanism for synthesis of guaiacyl-syringyl lignin typical of tracheid cell walls in angiosperm trees.     

Yield capacity of planted Picea abies in northern Sweden

Norway spruce (Picea abies (L.) Karst.) is the dominant species in the older forests of central northern Sweden. However, spruce has seldom been planted in the area, partly because existing tools for site classification have indicated a low yield capacity for the species. The aim of this study was to examine the yield capacity of spruce on the basis of existing plantations. In total, 91 operational and experimental plantations in the age interval 27–46 yrs were sampled. Stands were located between 62° and 65° N at altitudes 130–620 m a.s.l. Site index was estimated by height growth and site‐factor equations previously developed from old‐growth stand data. Height developments in the plantations indicate that site index for these stands is on average 4.6 m higher than predicted by site‐factor equations. The differences between the two methods are larger on poor sites than on rich sites. No systematic deviations of top height development from the site index curves could be detected on remeasured sample plots. Existing growth models were applied on measured stand data to predict future growth. Calculated mean annual increments were on average 20% lower when site index was predicted by site factors instead of height and age. The bias means that the yield capacity of planted spruce in northern Sweden has been underestimated by about 35%.     

Pathogenicity of ophiostomatoid fungi on Picea abies in Slovenia

Pathogenic fungi can survive and develop in living plants, often causing diseases in the host. Some theories speculate that pathogenic ophiostomatoid fungi provide benefits to its vectors – bark beetles – by overcoming the tree's defence mechanisms. This study reports the results of an experiment in south‐eastern Europe in which mature and seedling Norway spruce trees were artificially inoculated with various ophiostomatoid fungi. The aim of the experiment was to determine the relative virulence of ophiostomatoid fungi by assessing the ability of the fungi to stimulate host tree defence mechanisms through inoculation experiments. Experiments were performed by inoculation of Picea abies in seedling and mature trees. The following fungi were used in low‐density and seedling inoculations: Ophiostoma ainoae, O. brunneo‐ciliatum, Grosmannia cucullata and an unidentified Leptographium sp., O. bicolor, O. fuscum, O. piceae, G. penicillata and G. piceiperda. Endoconidiophora polonica was used in mass and seedling inoculations. Various characteristics such as host vitality, blue stain, lesion and resin outflow were measured before and after the trees were felled. E. polonica caused blue stain, induced large lesions and killed some of the mature trees and seedlings, confirming earlier reports that it is a strong wound pathogen. Only E. polonica, Leptographium sp. and O. ainoae caused blue stains in the sapwood of inoculated seedlings. In low‐density inoculations, G. piceiperda induced intense necrosis and had higher values for all the characteristics monitored. Some of the other ophiostomatoid fungi showed a moderate level of pathogenicity. Fungi with the capacity to stimulate a host defence mechanism could play a role in the establishment of bark beetle populations.     

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.     

Element concentrations in the xylem sap of Picea abies (L.) Karst. seedlings extracted by various methods under different environmental conditions

We used a Scholander pressure chamber to assess the effects of various extraction methods under different environmental conditions on element concentrations in xylem sap of 3-year-old Picea abies (L.) Karst. seedlings. Sap from excised shoots contained higher element concentrations when extracted at low than at high over-pressures. When comparing plants differing in water status, we found that a high extraction over-pressure introduced a systematic error into the data. For example, in well-watered non-transpiring plants relative to unwatered transpiring plants, potassium concentrations were 70% higher in sap extracted at 0.1 MPa over-pressure, but only 10% higher in sap extracted at 1.0 MPa over-pressure. Moreover, treatment effects depended on the time of day when the sap was extracted. Increased water flux in transpiring plants relative to non-transpiring plants resulted in reduced xylem sap element concentrations when samples were collected after 9 h of transpiration, but not after 4 to 6 h of transpiration. Drought had little effect on xylem sap element concentrations, indicating that rates of element release into xylem conduits, element depletion by growing tissues, and water flow maintained a balance that may prevent nutrient stress during short-term drought.     

Development and variation of wood density, kraft pulp yield and fibre dimensions in young Norway spruce (Picea abies)

Important wood, pulp and fibre properties were investigated on small wood samples from two Rumanian, one German and one Norwegian provenance of Norway spruce (Picea abies) grown in Sørkedalen, Norway. Several samples were collected from inside each single tree, both in radial and transversal direction in the stem. Data were collected from a total of 59 trees, each 28 years of age. All investigated properties showed close relationship to ring number (RN) (cambium age). For basic density and fibre wall thickness (FWT), a fast decline was first observed when moving from pith to bark and a minimum value was found around RN 5–8. The declining trend was then followed by an increase. Kraft pulp yield, fibre length and fibre width (FW) also increased with RN, but the increase was most pronounced close to the pith. The pulp yield (PY) more or less stabilised outside of RN 5–6. FW showed a decrease outside of RN 10–12, but this was probably due to the remarkable simultaneous drop in ring width for the investigated trees. FW decreased, while basic density and fibre length increased with increasing height in the tree. PY and FWT were not affected by height in tree. Diameter at breast height (DBH) was the most important variable indicating differences between trees in the investigated material. Basic density, fibre length and FWT decreased, while FW increased with increasing DBH. Height to crown had a positive effect on basic density, but had no influence on any of the remaining properties. Differences between provenances were found for basic density, fibre length and FW. The analyses showed that it is possible to describe the variation inside and between trees satisfactorily for a range of important wood and fibre properties.     

Ultrasound emission after cycles of water stress in Picea abies

The relationships among rate of ultrasound acoustic emission (AE), xylem water potential and transpiration rate were investigated in 5-year-old potted saplings of Picea abies Karst. after cycles of water stress. Water-stressed plants displayed minimum xylem water potentials of -3.9 MPa, near-zero transpiration rates and up to 45 AE counts per minute. After rewatering, water-stressed plants no longer produced AEs. Well-watered control plants produced only a small number of ultrasonic AEs. After three cycles of water stress (lasting 24 days in total), it was estimated that about two-thirds of the functional tracheids were embolized. The concomitant reduction in hydraulic conductance was about 70%.     

Between- and within-site variation of density and bending properties of Picea abies structural timber from Norway

This study provides an analysis on the variability of structural timber of Norway spruce (Picea abies) grown in Norway. Density, modulus of elasticity (MOE) and bending strength were measured on 1188 boards from 205 trees, sampled from 14 sites throughout Southern Norway, Eastern Norway and Trøndelag. The area represents the procurement area for the majority of Norwegian sawmills. The variability of the timber properties was analysed in a linear mixed model where the random variance was divided into variance due to site, variance due to trees and within-tree variance. Models describing variance due to site based on site index, altitude and latitude were developed, and combined with data from the Norwegian National Forest Inventory to estimate mean values and variability of the timber properties. The results showed that major parts of the variance due to site are explained by altitude and site index, and for density also by latitude. Major parts of the variance due to site and the variance due to trees in bending strength and MOE were explained by density.     

Fungi in bark peeling wounds of Picea abies in central Sweden

A total of 210 Norway spruce trees with stem wounds resulting from bark peeling by moose (Alces alces) were examined in three 45–50-year-old stands that contained 20-30% of damaged trees. Injured stems were between 8 and 40 cm diameter at breast height (d.b.h.) and showed 1 to 23-year-old wounds. Wounds varied in size from 2 to 4815 cm². The size of injury correlated negatively with the age of the injury (r=-0.24; p=0.004), and positive correlation was established between the age of the injury and d.b.h. of the wounded tree (r=0.50; p=0.0001). Each wound was sampled once using an increment borer and fungi were isolated. Among basidiomycetes, Stereum sanguinolentum was the most common (isolated from 26.7% of the damaged stems). Other common species were Cylindrobasidium evolvens (23.8%), Amylostereum areolatum (5.2%), A. chailletii (0.5%), Heterobasidion annosum (5.2%), Peni-ophorapithya (1.4%), Sistotrema brinkmannii (1.0%). The ascomycete Nectria fuckeliana was the most common among all fungi (present in 35.7% of bark peeling wounds). The frequency of S. sanguinolentum infection correlated positively with the age of the injury (r=0.27; p=0.001) and the opposite relationship was revealed for C. evolvens (r=?0.30; p=0.0001). Furthermore, C. evolvens infection correlated positively with the wound size (r=0.30; p=0.0001) and negatively with the tree d.b.h. (r=?0.20; p=0.004). A positive correlation was found between tree d. b. h. and the occurrence in stems of H. annosum (r=0.23; p=0.001 and N. fuckeliana (r=0.23; p=0.0006). The spruce bark beetle Dendroctonus micans attacked 14.8% of wounded trees. Presence in stems of N. fuckeliana was associated significantly with the D. micans attack (r=0.190; p =0.006; χ² test: p=0.01). Except for a negative correlation between infections of S. sanguinolentum and H. annosum (r=-0.140; p=0.04), no significant relationship between fungal species was found.     

Effect of sulfur-dioxide on protein-SH in needles of Picea abies1

The effect of SO₂ on the SH-groups of proteins in needles of Picea abies is studied. The content of total protein SH and glutathione is estimated with 5.5′-Dithio-bis-nitrobenzoic-acid (Ellman reagent). In addition structural protein SH is measured microspectrophotometrically in sections stained with the SH reagent, mercurochrome. The results indicate that GSH and also total protein SH and structural protein SH are increased in needles of Picea abies growing in SO₂ polluted areas. Compared to needles of trees in relatively unpolluted areas average increases of 2. 33 (GSH), 1.22 (total protein SH) and 1.55 (structural protein SH) are found.     

挪威云杉幼树韧皮部挥发性物质的测定

通过GC-MS测定挪威云杉幼树主干上部与下部韧皮主要挥发性物质的化学成分与含量，结果表明挪威云杉幼树的韧皮部挥发性物质的主要成分为α-蒎烯、茨烯、β-蒎烯、月桂烯、3-蒈烯、柠檬烯、β-水芹烯7种单萜化合物。α-蒎烯、β-蒎烯、月桂烯、3-蒈烯、β-水芹烯的含量在上部韧皮与下部韧皮有明显差异，其中上部韧皮部α-蒎烯、β-蒎烯和β-水芹烯的含量明显高于下部主干韧皮部,这为解释松树皮象的取食习性，打下了基础。     

Genetic parameters of growth and wood quality traits in Picea abies

Genetic parameters were estimated for wood and growth traits in two 19-yr-old clonal trials and a 40-yr-old full-sib progeny trial of Norway spruce [Picea abies (L.) Karst.]. In the clonal trials high (>0.4) broad-sense heritabilities were found for wood density traits, lignin content, number of internal cracks, growth traits, spiral grain and number of resin canals. Moderate (0.2–0.4) heritabilities were found for tracheid lumen diameter and cell wall thickness, microfibril angle and tracheid length, while low heritabilities (<0.2) were found for pulp yield, fibre strength, wood stiffness and wood colour. Lignin content and pulp yield showed low genetic variation, whereas the genotypic coefficient of variation for most other traits ranged between 5 and 15%. Most traits showed low levels of genotype by environment interaction. Among the wood properties, latewood proportion, earlywood density and ring density showed significant, adverse correlations with volume in both clonal trials.