Distribution of preservatives in thermally modified Scots pine and Norway spruce sapwood

Studying the impregnation and distribution of oil-based preservative in dried wood is complicated as wood is a nonhomogeneous, hygroscopic and porous material, and especially of anisotropic nature. However, this study is important since it has influence on the durability of wood. To enhance the durability of thermally modified wood, a new method for preservative impregnation is introduced, avoiding the need for external pressure or vacuum. This article presents a study on preservative distribution in thermally treated Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) sapwood using computed tomography scanning, light microscopy, and scanning electron microscopy. Secondary treatment of thermally modified wood was performed on a laboratory scale by impregnation with two types of preservatives, viz. Elit Träskydd (Beckers) and pine tar (tar), to evaluate their distribution in the wood cells. Preservative solutions were impregnated in the wood using a simple and effective method. Samples were preheated to 170 °C in a drying oven and immediately submerged in preservative solutions for simultaneous impregnation and cooling. Tar penetration was found higher than Beckers, and their distribution decreased with increasing sample length. Owing to some anatomical properties, uptake of preservatives was low in spruce. Besides, dry-induced interstitial spaces, which are proven important flow paths for seasoned wood, were not observed in this species.     

Growth rates of Heterobasidion annosum s.s. and H. parviporum in functional sapwood of Pinus sylvestris and Picea abies

Growth rates of H. annosum s.s. and H. parviporum were investigated in the functional sapwood of young Pinus sylvestris and Picea abies plants as an indicator of the relative susceptibilities of the hosts to these pathogens. The stems of 520 five‐year‐old P. abies and 321 four‐year‐old P. sylvestris plants were inoculated and the extent of infection determined 16 weeks later. H. annosum s.s. grew further than H. parviporum in P. sylvestris sapwood, while in P. abies, no differences between the two Heterobasidion spp. were found. Both H. annosum s.s. and H. parviporum spread faster in the sapwood of P. abies than in P. sylvestris. There was high within‐host species variation in growth rates for both P. sylvestris and P. abies suggesting it may be possible to identify tree genotypes with lower susceptibility.     

Effect of planting Scots pine around Norway spruce stumps on the spread of Heterobasidion coll.

Management of a Norway spruce stand planted on a site infected by Heterobasidion coll. is problematic because the fungus spreads vegetatively from the colonized stumps of the previous generation to the new seedlings. Growing of mixed stand with more resistant tree species has been suggested to decrease the economic losses caused by butt rot in Norway spruce trees. The mechanistic simulation model Rotstand describing the spread of Heterobasidion coll. in coniferous stands of southern Finland was used to study the effect of planting Scots pines around colonized clear‐felling stumps of Norway spruce of the previous generation. Planting of Scots pines in clusters around colonized stumps markedly decreased the butt rot of Norway spruce trees at the age of 20 years and at clear felling. If the same number of Scots pines were planted randomly, the effect was weak. When the average diameter of colonized clear‐felling stumps was 30 cm, a Scots pine circle with a radius of 3 m resulted in the highest soil expectation value (SEV) at 2% discounting rate, whereas with 40‐cm stump diameter, a 4‐m radius produced the highest SEV. When the proportion of Heterobasidion parviporum in the old colonized stumps was 50% instead of 95%, planting pines around colonized stumps still clearly decreased the butt rot at the age of 20 years and in final felling.     

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.     

Spatial distribution and persistence of Heterobasidion parviporum genets on a Norway spruce site

Spatial distribution of Heterobasidion genets over a period of ca 50 years in two successive generations of Norway spruce (Picea abies) was unravelled. The genets were first identified in 1993 in a naturally regenerated 43‐year‐old spruce stand that had been thinned the previous winter. Heterobasidion parviporum was found in 17.5% of the old stumps of the previous spruce generation. Nine genets were identified on the study plot; seven of them were present in old stumps of the previous tree generation and two only in the new spruce generation. Eighteen spruce trees of the new generation were infected, 15 of them by vegetative growth of genets originating from the old stumps. The study plot was investigated again in 2005. No new genets had been established after thinning, and three old genets had died out. The remaining genets had infected five new trees, most likely from the thinning stumps of diseased trees. At the age of 56 years, 16.1% of the residual spruces were infected by Heterobasidion. The results of this study suggest that if spore infection to stumps of spruce can be prevented, the decay frequency caused by H. parviporum will not necessarily increase in successive generations.     

Infection of roots of Norway spruce (Picea abies) by Heterobasidion annosum

Wounding of excised roots of Norway spruce [Picea abies (L.) Karst.] caused a rapid accumulation of phenolic compounds, decrease of starch content and of certain enzyme activities in the ray cells. Infection of the wounds by Heterobasidion annosum (Fr.) Bref. intensified these processes, caused oxidation of the phenols, and induced activity of acid phosphatase and β-glucosidase in the ray cells of the infected area. The reactions were oxygen-dependent. Intra- and extracellular enzymes were detected in the wood-infecting hyphae. Other wood-growing fungi caused similar reactions in the wood as H. annosum.     

Butt-rot of Scots pine in Thetford Forest caused by Heterobasidion annosum: a local phenomenon

Heterobasidion annosum was found to be the main cause of decay in 50–60-year-old Scots pine in Thetford Forest in East England. On some sites, up to 15% of the trees could be suffering from H. annosum rot, with decay occasionally extending 2 m or more up the stem. Prior to this study, Scots pine had been considered as resistant to H. annosum butt-rot in Britain, except in overmature trees.     

Durability of thermally modified sapwood and heartwood of Scots pine and Norway spruce in the modified double layer test

In the present study, durability of untreated and thermally modified sapwood and heartwood of Scots pine and Norway spruce was examined using a modified double layer test. Base layer samples were partly on contact with ground where exposure conditions were harder than that in a double layer test above the ground. The base layer on ground contact gave results already after one year of exposure in Finnish climate, but the top layer of a double layer test element simulated more the situation of decking exposure.

Significant differences in durability and moisture content (MC) between the wood materials were detected after six years of exposure in the field. Thermally modified pine heartwood performed very well in all layers of the test element and only minor signs of decay were found in some of the base samples. Both sapwood and heartwood of thermally modified spruce were suffering only slight amounts of decay while thermally modified pine sapwood was slightly or moderately decayed. Untreated sapwood samples of pine and spruce were severely decayed or reached failure rating after six years in the field. Untreated heartwood samples performed clearly better. The highest MCs were measured from untreated and thermally modified pine samples. Thermal modification increased significantly the durability and decreased the MC values of all wood materials.     

Effects of vitality fertilization on the growth of Heterobasidion annosum in Norway spruce roots

The effects of vitality fertilization on the growth of Heterobasidion annosum in roots of Norway spruce (Picea abies) were studied in a 53-year-old, naturally regenerated spruce stand in southern Finland. The fertilizer treatments were: (1), unfertilized control; (2), a compound fertilizer containing P, K, Ca, Mg, S, Cu, Zn and B; (3), as 2 with additional nitrogen; (4), as 3 with additional lime; and (5), a mixture composed on the basis of needle analysis, containing N, P, K and Cu. Three growing seasons after fertilization, four roots of eight trees in each treatment were inoculated with four different strains of H. annosum. Spread of the fungus from the inoculation point was determined after 12 months. Mean spread rates upwards in roots were 18.2, 25.6, 21.3, 26.0 and 29.8 cm/year in treatments 1, 2, 3, 4 and 5, respectively. These results suggest a tendency towards faster growth by H. annosum in fertilized trees. However, there was considerable variation in fungal growth at both the tree and root level and differences between treatments were not statistically significant.     

Immunocytochemical localization of pathogenesis-related proteins in roots of Norway spruce infected with Heterobasidion annosum

The occurrence and accumulation of β-1, 3-glucanase and chitinase in seedling roots of spruce (Picea abies) following challenge by the root-rot pathogen Heterobasidion annosum were studied. Chitinase activity increased 2–3 fold following inoculation, whereas no significant increase in the activity levels of glucanase was recorded during infection. With TEM immunogold labelling, the enzymes were localized in protein aggregates in host tissues and in the cell walls of intercellular hyphae. Gold particles were sparse and irregularly distributed within host-cell walls. Only minor labelling was observed on hyphal walls coated with electron-dense materials. The labelling intensity increased with infection time and was always higher than in non-infected seedling roots. When this experiment was repeated using root samples inoculated with the saprophyte Phlebiopsis gigantea, a similar labelling pattern was observed. The cross reactivity of antisera raised against sugar-beet chitinase and glucanase with spruceroot enzyme extracts was demonstrated using dot-blot assays and ELISA.     

Effectiveness of some biocontrol and chemical treatments against Heterobasidion annosum on Norway spruce stumps

The biocontrol agents Trichoderma harzianum, Verticillium bulbillosum, Hypholoma fasciculare, Phanerochaete velutina, Vuilleminiia comedens together with the chemical treatments propiconazole and the culture filtrate of V. bulbillosum were tested on Norway spruce stumps during 2 years in order to evaluate their effectiveness against Heterobasidion annosum. The results were compared, in 1 year, with the effectiveness of Phlebiopsis gigantea. All the treatments reduced H. annosum infections. The best results were obtained with P. velutina, propiconazole and with culture filtrates and conidial suspensions of V. bulbillosum.     

Changes in host chitinase isoforms in relation to wounding and colonization by Heterobasidion annosum: early and strong defense response in 33-year-old resistant Norway spruce clone

We studied the defense reactions of 33-year-old susceptible and resistant clones of Norway spruce (Picea abies (L.) Karst.) to the major root-rot fungus Heterobasidion annosum (Fr.) Bref. and determined if tissue cultures can be used as a model system for studying defense responses of mature trees at the molecular level. Quantitative PCR analysis of genomic DNA obtained from samples taken at different times along the lesion length in living bark indicated that the fungus was present in higher amounts and extended further into the host tissue in the susceptible clone than in the resistant clone. In protein extracts from the same lesion samples, there were differences in temporal and spatial changes in host chitinase isoform profiles between the resistant and susceptible clones. Host chitinase isoforms with pI values approximately 4.8, 4.4 and 3.7 increased more during the first 7 days after wounding and inoculation and extended further along the lesion length in the resistant clone than in the susceptible clone. These results suggest that the time from wounding and infection to induction of defense-related expression is shorter in the resistant clone indicating a more efficient host defense response than in the susceptible clone. Tissue cultures from the same clones were not resistant to H. annosum and showed no difference in the timing of the increase in chitinase isoforms in response to the pathogen. However, tissue cultures from both clones showed an increase in chitinase isoforms within 6 to 24 h past inoculation, indicating that increased chitinase expression in response to the pathogen is part of a general defense response common to both mature clones and tissue cultures.     

Effect of uninucleate Rhizoctonia on Scots pine and Norway spruce seedlings

One‐year‐old container‐grown seedlings were planted in spring on clear cut areas: the Norway spruce (Picea abies) on a moist upland site (Myrtillus‐type) and Scots pine (Pinus sylvestris) on a dryish upland site (Vaccinium‐type). While still in the nursery, half of the seedlings of each species had been inoculated during the previous summer, with a uninucleate Rhizoctonia sp., a root dieback fungus. At outplanting all the seedlings appeared healthy and had a normal apical bud, although the height of the inoculated seedlings was less than that of the uninoculated control seedlings. At the end of the first growing season after planting, the mortality of inoculated Scots pine and Norway spruce seedlings was 25 and 69%, respectively. After two growing seasons the mortality of inoculated seedlings had increased to 38% for Scots pine and 93% for Norway spruce. The mortality of control seedlings after two growing seasons in the forest was 2% for Scots pine and 13% for Norway spruce. After outplanting the annual growth of inoculated seedlings was poor compared with the growth of control seedlings. These results show that, although Rhizoctonia‐affected seedlings are alive and green in the nursery, the disease subsequently affects both their survival and growth in the forest.     

Durability of thermally modified Norway spruce and Scots pine in above-ground conditions

Abstract

One of the main objectives of thermal modification is to increase the biological durability of wood. In this study the fungal resistance of Norway spruce and Scots pine, thermally modified at 195°C and 210°C, was studied with a lap-joint field test. Untreated pine and spruce and pine impregnated with tributyl tin oxide (TBTO) and copper, chromium and arsenic (CCA) were selected as reference materials. The evaluations were carried out after 1, 2 and 9 years of exposure. After 1 and 2 years of exposure mainly discoloration was detected. Only the untreated pine was slightly affected by decay fungi. There were significant differences in the decay ratings of untreated and thermally modified wood materials after 9 years in the field. While the untreated wood materials were severely attacked by decay fungi or reached failure rating, only small areas of incipient decay were detected in the thermally modified samples. Thermally modified pine was slightly more decayed than thermally modified spruce. The only wood material without any signs of decay was CCA-treated pine, since some of the TBTO-treated pine samples were also moderately attacked by fungal decay. The results of the lap-joint test had a good correlation with mass losses in a laboratory test with brown-rot fungi.     

Decay resistance of sapwood and heartwood of untreated and thermally modified Scots pine and Norway spruce compared with some other wood species

Abstract

Thermal modification has been developed for an industrial method to increase the biological durability and dimensional stability of wood. In this study the effects of thermal modification on resistance against soft- and brown-rot fungi of sapwood and heartwood of Scots pine and Norway spruce were investigated using laboratory test methods. Natural durability against soft-rot microfungi was determined according to CEN/TS 15083-2 (2005) by measuring the mass loss and modulus of elasticity (MOE) loss after an incubation period of 32 weeks. An agar block test was used to determine the resistance to two brown-rot fungi using two exposure periods. In particular, the effect of the temperature of the thermal modification was studied, and the results were compared with results from untreated pine and spruce samples. The decay resistance of reference untreated wood species (Siberian larch, bangkirai, merbau and western red cedar) was also studied in the soft-rot test. On average, the soft-rot and brown-rot tests gave quite similar results. In general, the untreated heartwood of pine was more resistant to decay than the sapwood of pine and the sapwood and heartwood of spruce. Thermal modification increased the biological durability of all samples. The effect of thermal modification seemed to be most effective within pine heartwood. However, very high thermal modification temperature over 230°C was needed to reach resistance against decay comparable with the durability classes of “durable” or “very durable” in the soft-rot test. The brown-rot test gave slightly better durability classes than the soft-rot test. The most durable untreated wood species was merbau, the durability of which could be evaluated as equal to the durability class “moderately durable”.     

Responses of in vitro grown pine seedlings to infection by four strains of Heterobasidion annosum1

Scots pine seedlings growing in pure cultures were inoculated with 4 isolates of Heterobasidion annosum. Significant variation in resistance to this pathogen was observed within and among progenies of pine trees selected at two different locations. Mycorrhizal synthesis occurred more frequently in cases where seedlings were inoculated with less pathogenic isolates of the pathogen.