Methyl jasmonate and oxalic acid treatment of Norway spruce: anatomically based defense responses and increased resistance against fungal infection

To study the effect of chemical pretreatment on conifer resistance, 13-year-old Norway spruce (Picea abies (L.) Karst.) trees were treated with methyl jasmonate (MJ) or oxalic acid (OxA) on the outer bark and inoculated with the pathogenic blue-stain fungus Ceratocystis polonica (Siem.) C. Moreau 4 weeks later. Both chemicals significantly reduced symptoms of fungal infection, but MJ was more effective than OxA (51 versus 18% reduction in length of necrotic lesions in the phloem relative to untreated control trees). Anatomical examination of treated stem tissues showed that MJ induced extensive formation of traumatic resin ducts in the xylem and extra polyphenolic parenchyma (PP) cells in the secondary phloem between the cambium and the regular annual PP cell layer. No traumatic resin ducts were formed after treatment with OxA, and the coverage of extra PP cells in OxA-treated tissues was not significantly higher than in the controls. The anatomically based defense reactions induced by MJ were similar to the reactions observed after pathogen infection, mechanical wounding and bark beetle attack. Neither MJ nor OxA had apparent phytotoxic effects on Norway spruce at the concentrations used, with needle and stem tissues of all trees appearing normal without visible symptoms of toxicity. However, trees treated with MJ had 30% less radial sapwood growth than control trees. In conclusion, MJ treatment of Norway spruce appears to have practical potential as a tool for increasing plant resistance to fungal infection, but with a modest reduction in sapwood growth.     

Induced resistance of Norway spruce,variation of phenolic compounds and their effects on fungal pathogens

Three clones of Norway spruce (Picea abies) were studied for their response to mass‐inoculation with the blue‐stain fungus Ceratocystis polonica. The effect of different pretreatments (fungal inoculation and wounding) before mass‐inoculation was investigated for their possible role in an acquired resistance reaction. Pretreated trees showed enhanced resistance to the subsequent mass‐inoculation relative to control trees that received no pretreatment. Furthermore, the fungal colonization of inoculated trees was less than that of wounded trees. The phenolic content of the bark, analysed by RP‐HPLC, was compared in trees receiving different treatments. Trees inoculated with C. polonica had higher average concentration of (+)‐catechin, taxifolin and trans‐resveratrol than wounded trees. Both inoculated and wounded trees had higher average concentrations of these compounds than control trees. The effect of the phenolic extract of Norway spruce bark on the growth of the root rot fungus Heterobasidion annosum and the blue‐stain fungi C. polonica and Ophiostoma penicillatum were investigated in vitro. Heterobasidion annosum was not negatively affected, and the extracts had fungistatic effects on the blue‐stain fungi. The growth of O. penicillatum was more inhibited than the growth of the more aggressive C. polonica.     

Quantitative assessment of fungal colonization in Norway spruce after green pruning

In July and in October of 1978, 1979, 1980 five Picea abies stands in Baden-Württemberg, Germany were green-pruned up to a tree height level of c. 10 m. In 1993–94, 30 pruned trees and five control trees from each stand were harvested for investigation. The quality of pruning proved to be excellent because only few branch collar, bark injuries or too long branch stubs were detected. Based on isolations from more than 6000 wood specimens of c. 20 mm³ obtained from 175 trees, specific infection rates are given for different tree compartments. Wood decay fungi as well as important blue stain fungi were only rarely present. Tolerable discolourations were limited to the stub containing core of a few trees. Wood formed after pruning showed neither more fungal infections nor other related disadvantages compared with the respective increment from unpruned control trees. Nectria fuckeliana, proved to be the most abundant fungus in pruned trees especially in the branch stubs. Respective infection rates in and close to dry branches were compared. Bacteria could be found in the pruned branch stubs, however, they did not penetrate into the heartwood or sapwood of the stems. Both summer and autumn pruning involve only very low risk of wood deterioration. However, summer pruning shows some advantages with respect to lower infection rates by N. fuckeliana.     

Polymerisation of added coniferyl alcohol by inherent xylem peroxidases and its effect on fungal decay resistance of Norway spruce

It is known that active peroxidase isozymes exist in mature wood of Norway spruce (Picea abies L. Karst.) and that they remain active for years and are found even in the heartwood (in Scots pine), where all cellular activity has ceased. This peroxidase activity was utilised in the impregnation of wood blocks with a natural monolignol, coniferyl alcohol and hydrogen peroxide. The hypothesis was that the internal wood peroxidases would oxidise the added monolignol and bind it stably into the cell wall matrix, which could hinder fungal decay. Since coniferyl alcohol is not very soluble in water, the impregnation was done under vacuum with an acetone–water solution containing 10% coniferyl alcohol and 0.4 mM H₂O₂ at room temperature (ca. 0.02 g of coniferyl alcohol was added to 1 g of wood). After impregnation, dimers of coniferyl alcohol and free coniferyl alcohol were found in acetone extracts with GC–MS analysis. Penetration of coniferyl alcohol and non-extractable reaction products were studied from the wood blocks with FTIR PAS technique. The wood samples treated were also subjected to a fungal decay test with Coriolus versicolor. This treatment hindered fungal decay in a 60-day experiment and led to a dry weight loss of 8.8% in comparison with 19.9% in the control. The reactions of coniferyl alcohol and H₂O₂ in the presence of peroxidases are discussed as well as the use of monolignols to increase wood decay resistance.     

Quantification of fungal infection in spruce needles from different sourcesc

A large number of batches of green as well as of chlorotic spruce needles were tested for fungal infection. Using ergosterol as a biochemical indicator for fungal colonization, the highest ergosterol contents were always found in partially or uniformly necrotized needles. However, no correlation was found between the ergosterol concentration and the chlorophyll content i. e. the degree of bleaching.     

Effect of fungal exposure on the strength of thermally modified Norway spruce and Scots pine

Abstract

Thermal modification at elevated temperatures changes the chemical, biological and physical properties of wood. In this study, the effects of the level of thermal modification and the decay exposure (natural durability against soft-rot microfungi) on the modulus of elasticity (MOE) and modulus of rupture (MOR) of the sapwood and heartwood of Scots pine and Norway spruce were investigated with a static bending test using a central loading method in accordance with EN 408 (1995). The results were compared with four reference wood species: Siberian larch, bangkirai, merbau and western red cedar. In general, both the thermal modification and the decay exposure decreased the strength properties. On average, the higher the thermal modification temperature, the more MOE and MOR decreased with unexposed samples and increased with decayed samples, compared with the unmodified reference samples. The strength of bangkirai was least reduced in the group of the reference wood species. On average, untreated wood material will be stronger than thermally modified wood material until wood is exposed to decaying fungi. Thermal modification at high temperatures over 210°C very effectively prevents wood from decay; however, strength properties are then affected by thermal modification itself.     

The search for resistance to Heterobasidion root rot in Norway spruce - old and new approaches in studies of infection biology

The paper surveys the methods used for studying the root rot in Norway spruce (Picea abies) and summarizes the major results on Heterobasidion infection, the importance of the rhizosphere, preformed and induced resistance factors, search for trees with high resistance, and on virulence factors of the fungus. New approaches are discussed regarding where to look for resistance, how to study host-parasite interaction, the importance of stress factors, and the need for short-term experiments and standardized inoculum.     

Inducible anatomical defense responses in Norway spruce stems and their possible role in induced resistance

Norway spruce (Picea abies (L.) Karst.) trees were preinoculated with a sublethal dose of the blue-stain fungus Ceratocystis polonica Siem. (C. Moreau) 1 to 52 weeks before they were mass inoculated with the same fungus. Trees pretreated 1 week before mass inoculation had similar, severe symptoms of fungal infection as the control trees. Pretreatment 3, 6 or 9 weeks before mass inoculation resulted in effective protection of the trees, reducing pathogenic symptoms by 63-90% relative to the control trees, whereas pretreatment 52 weeks before mass inoculation gave intermediate protection (44-71% reduction in symptoms). Thus, pretreatment induced resistance to the blue-stain fungus in Norway spruce by a process that requires more than 1 week to become activated and protects trees for at least one year after pretreatment. Pretreatment induced formation of traumatic resin ducts (TDs) in the sapwood and swelling and proliferation of polyphenolic parenchyma cells (PP cells) in the phloem. Trees pretreated 3-9 weeks before mass inoculation had more TDs and showed greater swelling of existing PP cells than control trees or trees pretreated 1 week before mass inoculation. We conclude that induced disease resistance in Norway spruce is probably associated with PP cell activation and TD induction, because resistance was enhanced within the same time frame as the induction of these defense responses.     

Proteins in Norway spruce thermomechanical pulp

Two methods of cytochemical staining using Coomassie dye and Cu⁺-bicinchoninic acid, respectively, showed that there are proteins in thermomechanical pulp (TMP) of Norway spruce. Protein isolated from TMP was analyzed for amino acid composition. There was about twice the amount of acidic amino acid material compared with basic amino acids, and the presence of glucosamine indicated that the isolated polypeptides also contained glycoproteins. The presence of proteins in ray cells and fiber tracheids in TMP adds to the chemical heterogeneity of the structurally complex high-yield pulp.     

Interactions among Norway spruce,the bark beetle Ips typographus and its fungal symbionts in times of drought

Journal of Pest Science - Resilience and functionality of European Norway spruce forests are increasingly threatened by mass outbreaks of the bark beetle Ips typographus promoted by heat, wind...     

Storm-induced tree resistance and chemical differences in Norway spruce (Picea abies)

Introduction   

Windstorm is one of the most destructive environmental disturbance factors on forests, but its influence on conifer defense chemistry and susceptibility to insects and diseases is not well understood.     

Stand responses to initial spacing in Norway spruce plantations in Norway

Abstract

Using data from nine spacing experiments of Norway spruce (Picea abies (L.) Karst.) in Norway, covering wide ranges of site index and initial spacing, this study evaluated stand basal area and volume responses to initial spacing and examined whether these responses varied by stand ages or site quality. We developed nonlinear regression models that described the standing basal area or volume responses to initial spacing along with site index and stand ages. The results show that closer spacings produced higher standing basal area and volume than wider spacings. The response curves are highly nonlinear in younger stands and become nearly linear in mid-rotation stands, indicating stronger responses at younger ages and weaker responses as age increases. Furthermore, for young stands, spacing effects are stronger at closer than at wider spacings. The basal area and volume responses to initial spacing tend to be similar across site indices. However, the interaction of site index and stand age on spacing responses makes it difficult to isolate the effect of site index on spacing responses. Mortality is higher and begins earlier at closer spacings than wider. The mean diameter of the largest 100, 400, 600, and 800 trees ha^?1 increased with spacing in three out of the nine experiments. Dominant height did not vary by initial spacing for any of the experiments. The findings suggest that the extra volume production in stands of closer initial spacing is restricted to early stand development.     

Using drill resistance to quantify the density in coarse woody debris of Norway spruce

To evaluate the mass of coarse woody debris (CWD), it is necessary to quantify its density. Drill resistance measurements are introduced as a approach to estimate the density of CWD in different stages of decay. Dead logs of Norway spruce [Picea abies (L.) Karst.] from a Central European mountainous site were used as a test system to compare the new method with conventional predictors of wood density such as fast quantitative field estimates (e.g., knife probe) and classification of decay classes based on a set of qualitative traits and quantitative estimates. The model containing only drill resistance as a predictor explained 65% of the variation in wood density and was markedly better than models containing one or more of several conventional predictors. However, we show that the relationship between drill resistance and gravimetric wood density relationship is sensitive to the decay status. Therefore, the best model combines drill resistance and decay class (adj. R2 = 0.732). An additional experiment showed that drill resistance is also sensitive to the moisture state (fresh vs. oven-dry) of the sample. The major potential of the method lies in its non-destructive nature which allows repeated sampling in long-term ecosystem studies or in protected areas where destructive sampling is prohibited. The limitations of the method are discussed and recommendations for applications are given.     

Boron retranslocation in Scots pine and Norway spruce

We previously traced 10B-enriched boric acid from shoots to roots to demonstrate the translocation of boron (B) in Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings. To gain a more detailed understanding of B translocation, we sought: (1) to demonstrate B retranslocation directly, by showing that foliar-applied 10B is located in the new growth after dormancy; and (2) to assess whether shoot-applied B affects growth in the long term. We applied 10B-enriched boric acid to needles of Scots pine and Norway spruce seedlings. After a dormancy period and 9 weeks of growth, small but significant increases in the 10B isotope were found in the new stem and needles of both species. In Scots pine, the total B concentration of the new stem was also increased. Both species contained polyols, particularly pinitol and inositol. Boron-polyol complexes may provide a mechanism for mobilizing B in these species. To determine the long-term effects of applied B, seedlings were grown for two growing seasons after the application of 10B to shoots. In Norway spruce, the proportion of 10B in the root systems and current needles of the harvest year was slightly higher than in the controls, and in Scots pine root systems, marginally so. The B treatment had no effect on growth of Norway spruce seedlings. In Scots pine seedlings, the B treatment caused a 33% increase in total dry mass and significantly increased the number of side branches.     

Ectomycorrhizas and cadmium toxicity in Norway spruce seedlings

We studied the effects of ectomycorrhizal colonization by Laccaria bicolor (Maire) Orton S238 and Paxillus involutus (Batsch) Fr. 533 on cadmium (Cd) toxicity in Norway spruce seedlings (Picea abies (L.) Karst.). Both mycorrhizal and nonmycorrhizal seedlings were exposed to 0 (control), 0.5 or 5 &mgr;M CdSO(4) for 9 weeks in a sand culture system with frequent addition of nutrient solutions. In pure culture, P. involutus and L. bicolor showed similar Cd tolerance. However, in symbiosis, the Cd treatments decreased colonization by L. bicolor but not by P. involutus. Paxillus involutus ameliorated the negative effects of 0.5 &mgr;M Cd on shoot and root growth and chlorophyll content of old needles, whereas L. bicolor did not. Mycorrhizal colonization did not affect Cd concentrations of old needles and roots of seedlings. Despite differences between the ectomycorrhizal fungi in colonization and ability to alleviate Cd toxicity of seedlings, both species reduced Cd concentrations of young needles to a similar degree compared with nonmycorrhizal seedlings. However, in the 0.5 &mgr;M Cd treatment, the Cd content of needles of seedlings colonized by P. involutus was increased, whereas the Cd content of needles of seedlings colonized by L. bicolor was similar to that of needles of nonmycorrhizal seedings. When the amount of Cd translocated to needles was expressed on a root length basis to account for differences in the size of the root systems, the amount of Cd translocated to the needles was similar in seedlings mycorrhizal with P. involutus and in nonmycorrhizal seedlings. All mycorrhizal seedlings were similarly affected by 5 &mgr;M Cd, indicating that the amelioration efficiency of ectomycorrhizal fungi is dependent on the metal concentration to which the roots are exposed. Concentrations of P, K, Ca, Mg and Mn were decreased by 5 &mgr;M Cd to a similar extent in both nonmycorrhizal and mycorrhizal seedlings. In contrast to L. bicolor, P. involutus increased P uptake and altered patterns of root branching. We conclude that mycorrhizas alleviate Cd-induced reductions in growth of Picea abies seedlings. Although the two mycorrhizal fungi examined differed in their ability to alleviate Cd toxicity, these differences were not related to differences in Cd uptake or translocation to the shoot of the mycorrhizal seedlings. We suggest that amelioration of Cd toxicity by P. involutus may be a result of improved P nutrition or changes in root morphology, or both.     

Potential production of Norway spruce in Sweden

In this paper, the potential stem wood production for Norway spruce was estimated for different regions in Sweden. This was done by using basic physiological relationships of intercepted radiation versus biomass production and knowledge of how a water deficit reduces the potential production, derived from results of field experiments on nutrient optimisation. To scale these relationships up to regional and national levels, data of incident radiation and humidity during the growing season for all of Sweden were used. The figures for potential and attainable production indicate that the temperate to boreal climate allows considerably higher production than the current production, if availability of water and nutrients is non-limiting. In northern Sweden, the attainable production is ca. 300% higher than the current production and in southern Sweden, the yield can be increased by ca. 100%. In absolute numbers, as a mean for a whole rotation period, it is possible to achieve an annual stem-wood dry mass production of 7–9 Mg ha⁻¹ in southern and central and ca. 5–6 Mg in northern Sweden. This increased production would mean that rotation periods can be shorter than they are now by 20–30 years in southern Sweden and by ca. 50–60 years in northern Sweden.     

Fungi on spruce needles in Norway II

Tiarosporella parca has been found in different plots in Norway, mostly in small amounts. However, in Pasvik, East-Finnmark, this species was rather common and replaced the otherwise most common species, Lophodermiumpiceae.     

Fresh-stem bending of silver fir and Norway spruce

The bending and growth characteristics of large fresh stems from four silver fir (Abies alba Mill.) and three Norway spruce (Picea abies (L.) Karst.) trees were studied. Twenty logs taken from different stem heights were subjected to four-point bending tests. From the bending test records, we calculated stress-strain curves, which accounted for detailed log taper, shear deformation and self weight. From these curves we determined, among other parameters, the modulus of elasticity (MOE), the modulus of rupture (MOR) and the work absorbed in bending (W). No significant differences were found between species for the wood properties examined. Values of MOE, MOR and W generally decreased with stem height, with MOR in the range of 43 to 59 MPa and MOE ranging from 10.6 to 15.6 GPa. These MOE values are twice or more those reported for stems of young Sitka spruce (Picea sitchensis (Bong.) Carr.) trees. Based on the radial growth properties measured in discs from the logs, we calculated predicted values of MOE and MOR for the stem cross section. The predictions of MOE were precise, whereas those of MOR were approximate because of a complex combination of different failure mechanisms. Methods to test and calculate MOE, MOR and W for the stems of living trees are discussed with the aim of improving analyses of tree biomechanics and assessments of forest stability protection.