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.     

Initial reactions in sapwood of Norway spruce and Scots pine after wounding and infection by Heterobasidion parviporum and H. annosum

The xylem surface of seedlings, stem material and roots of Norway spruce (Picea abies) and Scots pine (Pinus sylvestris) were inoculated with strains of Heterobasidion annosum s. str. and H. parviporum s. str. The depth of necrosis in wounded spruce increased at a linear rate for at least seven weeks of incubation, but the rate of necrotic spread was significantly faster in infected wounds. In wounded pine the necrosis was maintained at a more superficial level for several weeks. Both spruce and pine sapwood were initially infected by hyphae of both species. In spruce, the hyphae advanced at a constant rate behind the necrotic front. On the contrary, after 1–2 weeks living H. parviporum hyphae were rare in pine rays. Heterobasidion annosum hyphae survived in pine rays, phloem and tracheids, despite a heavy accumulation of phenolics and resins and were able to penetrate into the sapwood at a linear rate although slower than infections in spruce. Histochemistry and quantitative estimates demonstrated that peroxidase activity was initially higher in spruce sapwood than in pine. Within three days of incubation, the activity in spruce sapwood disappeared concurrently with deepening necrosis. However, in pine, in both control and infected samples, there was a significant increase in peroxidase activity in the area surrounding the superficial necrosis, up to the wound surface and in the cambium and phloem around the wound. After wounding and infection, the content of soluble protein increased significantly in wood of older trees but not in seedlings. Infection resulted in an increased formation of lipophilic extractives in both spruce and pine but to a significantly greater degree in the latter, whereas the amount of hydrophilic compounds decreased in both. High‐performance liquid chromatography (HPLC) analyses of lipophilic extracts showed that inoculation of pine with the two species of Heterobasidion increased the amounts of pinosylvin, its monomethylether and several other phenolics as also resinous compounds. The results obtained may be relevant in explaining the known higher resistance of Scots pine to H. parviporum.     

Blue‐stain infections in roots,stems and branches of declining Pinus sylvestris trees in a dry inner alpine valley in Switzerland

Increased mortality rates in Scots pine (Pinus sylvestris) forests have recently been observed in the inner alpine Swiss Rhone valley. Drought, in combination with stand competition, mistletoe infections as well as nematode and insect infestations, appears to be the main factor for the decline. In focus of this study was the occurrence and role of fungal pathogens in the decline dynamics. Branches, stems and roots of 208 trees in five different crown transparency classes were collected and inspected for blue stain and fungal infections. Neither Armillaria species nor Heterobasidon annosum s. str. were detected, but blue stain was commonly observed. Visible blue stain increased with increasing crown transparency. Among the recently dead trees, 80% showed visible blue stain in the branches, 90% in the roots and 100% in the stems. In the crown transparency classes 2 and 3 (25–60% crown transparency), five of the 103 trees showed visible blue stain in the roots, one of 130 trees in the stem but none in the branches. Blue‐stain fungi were isolated from all parts of the trees and from all crown transparency classes. Overall incidence of blue stain was highest in the roots and lowest in the branches. In class 2, roots of 60% of the trees were visibly blue‐stained or developed blue stain in culture, but stems of only 24% and branches of 14% of the trees. In the roots Leptographium species, mostly L. serpens, dominated. From stems and branches, mainly Ophiostoma species were isolated. The positive relationship between the incidence of blue stain and crown transparency, in combination with the high infection levels of roots of fairly vigorous Scots pines, indicates the pathogenic potential of the blue‐stain fungi. Hence, these fungi together with their insect vectors may well act as an important contributing factor involved in pine decline.     

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.     

Dry weight loss of wood after the inoculation of Scots pine stumps with Phlebiopsis gigantea

Artificial inoculation of stumps with Phlebiopsis gigantea (preparation‘PgIBL’) against Heterobasidion annosum in Scots pine (Pinus sylvestris) stands on former agricultural lands in Poland is being performed throughout the year. The practical efficiency of the biological control appears to be influenced by the moisture content in stumps and roots. In experiment 1, dry weight loss of wood 3 months and 6 months after P. gigantea inoculation in laboratory was investigated in the stumps after salvage cutting (felling of dead and dying trees) and after thinning (routine cutting of trees) and compared with the decay of the artificially inoculated stumps under field conditions. It was found that 6 months after inoculation the dry weight loss of the samples was about 3%, 17% and 22%, respectively. In experiment 2, the decay of wood from horizontal roots collected after thinning, inoculated with P. gigantea in laboratory, was evaluated. Three months after the inoculation, the loss of dry weight wood was about 22–52%, depending on initial moisture of the roots.     

The effect of acid rain treatments on the susceptibility of Pinus sylvestris to Gremmeniella abietina

Scots pine (Pinus sylvestris) seedlings were planted in a nursery bed 20 cm thick. The substrate had been removed from an illuvial horizon in the soil underlying a stand of Scots pine in Finland, which had suffered from severe infection by Gremmeniella abietina. The seedlings were irrigated for 3 years, applying three pH levels (2.5, 3.5, and 4.5) using sulphuric acid, nitric acid, and a 3:1 mixture of sulphuric and nitric acid. The pH of the pure water used in the control treatment was 6.1–6.4. The seedlings were inoculated with the conidiospores of G. abietina to find out their susceptibility to infection with the Scleroderris canker disease. Because to the adequacy of its buffering capacity, the soil was observed to become more acidic only in the treatments at pH 2.5. The concentrations of extractable Ca and Mg cations decreased while the concentration of Al and Fe increased with soil acidification during the 3-year experiment. Acidification did not increase the susceptibility of Scots pine to infection by the Scleroderris canker. Seedling growth was at its maximum in the acidification treatments where most nitrogen was added.     

不同复混肥对湿地松中龄林的施肥效应研究

为了研究不同复混肥对湿地松中龄林的施肥效应,以8年生湿地松林为研究对象,分别以施用2种普通复混肥和自制松树专用复混肥为施肥处理,各处理的施肥量均为700 g·株^-1;同时以不施肥为对照,进行施肥对比试验。结果表明:3种不同施肥处理中,以自制松树专用复混肥对湿地松林分生长的促进作用最大;施肥3年后林分的平均胸径、树高、单株材积分别较对照增长3.1%、4.7%、10.0%;施肥3年后自制松树专用复混肥处理的平均胸径、单株材积与2种普通复混肥处理间均存在极显著差异,树高均有显著差异;施用有长效作用并具针对性的松树专用复混肥能获得较高的木材增产效益。     

Early infection of Fagus sylvatica by Heterobasidion annosum sensu stricto

Heterobasidion annosum sensu stricto is the most important damaging agent in Scots pine stands planted on the former agricultural lands in Poland. The routine action in pine stands which have lost stability because of H. annosum root rot is to change stand management, including species conversion. In many cases, the Fagus sylvatica is used for this purpose. This study was the first assessment of widespread infection by H. annosum in young F. sylvatica plantations. Disease symptoms included atrophy and yellowing of leaves, wilting and the presence of pathogen sporocarps around the root collars of young trees. Heterobasidion annosum s. s. was observed on both 4‐ and 17‐year‐old beech. Based on annual increments, the disease could be present for 3–4 years before tree death. A high incidence of H. annosum in pine stumps of previous stands (80–100%) and dry periods in recent years may be the main reasons for such common infection of F. sylvatica. This work also showed that mice and frost were not the main killing factors F. sylvatica in plantations.     

Responses of conifer species of the Great Lakes region of North America to inoculation with the pitch canker pathogen Fusarium circinatum

Five conifer species grown in the Great Lakes region of North America were examined for their susceptibility to Fusarium circinatum, (syns. Fusarium subglutinans f. sp. pini and F. moniliforme var. subglutinans), the causal agent of pitch canker. Three‐year‐old (3‐0) seedlings of red (Pinus resinosa), jack (P. banksiana) eastern white (P. strobus), Scots (P. sylvestris) and Austrian (P. nigra) pine were planted in 4 l pots in a greenhouse at Auburn University in November 1998. In April and June 1999, seedlings were inoculated by removing a needle fascicle approximately 5 cm from the terminal bud and placing a drop containing F. circinatum conidia on the wound. Resin production, canker length and seedling mortality were recorded 12 weeks later. Jack, Scots and eastern white pine were the most susceptible with Austrian and red pine more resistant to the fungus. F. circinatum was re‐isolated from 37% to 96% of inoculated seedlings. The susceptibility of jack, Scots and eastern white pine indicates a potential risk to these important species of the region if F. circinatum were to be introduced into the area.     

Susceptibility of four conifer species to Gremmeniella abietina

G. abietina causes severe dieback in pole-stage stands of Scots pine in Britain. The susceptibility of ll provenances of Scots pine and of three other conifer species was tested by inoculations. Significant differences in susceptibility were found amongst Scots pine provenances, with those originating from a native Scottish pinewood at Loch Maree showing the highest levels of disease. Corsican pine was shown to be the most susceptible species tested although sporulation was greatest on Scots pine. Norway spruce was also shown to be susceptible when inoculated late in the growing season. Infection in lodgepole pine was negligible. High levels of β-phellandrene were tentatively linked with resistance among trees in a single stand of Scots pine.     

S and P intersterility groups in Heterobasidion annosum; Infection frequencies through bark of Picea abies and Pinus sylvestris seedlings and in vitro growth rates at different oxygen levels

Seedlings of Norway Spruce and Scots Pine were inoculated with S and P strains of the root rot fungus Heterobasidion annosum. Infection frequencies through the living bark layers were not related to subgroup affiliation in neither spruces nor pines. Growth rates in vitro were not related to growth rates in vivo. Oxygen concentration had a stronger effect on the growth rates of P than on S strains.     

Stand density and fertilization effects on aboveground allocation patterns and stem form of Pinus sylvestris in young stands

The increasing demand for forest biomass for energy generation could be partially met by growing denser stands and use of fertilizer. Before this is done at large scale, more knowledge of the effects of stand density and fertilization on aboveground allocation patterns and stem form is needed. Therefore, effects of pre-commercial thinning (PCT) to 3000 stems ha^?1, an unthinned dense control (C), and PCT combined with two levels of fertilization (100 kg ha^?1 of nitrogen applied either during the establishment of the field experiment (F1) or annually (F2)) were examined in 23- to 26-year-old Scots pine (Pinus sylvestris L.) stands six years after the establishment of the field experiment. In total, 114 sample trees were harvested using destructive biomass sampling. The growth allocation and stem form of trees with diameter at breast height (DBH; 1.3 m height) >5.0 cm were not affected by either the PCT or fertilization. Small trees (DBH < 5 cm) in denser, unthinned control plots had more slender stems (lower DBH/height ratios) and allocated less growth to branches and foliage than trees in PCT plots. Fertilization had little effect on the stem form and growth allocation of the smallest trees. Therefore, effects of stem density and fertilization on stem form and growth allocation to foliage were only found for small suppressed trees, and the treatments had very little influence on dominant and codominant trees.     

Field survival of containerized red and jack pine seedlings inoculated with mycelial slurries of ectomycorrhizal fungi

Red pine and jack pine seedlings growing in styroblocks were inoculated 8 wk after sowing with mycelium/agar slurries of 3 mycorrhizal fungi (Laccaria bicolor, Scleroderma citrinum, and an unidentified basidiomycete), and one suspected mycorrhizal fungus (Cantharellula umbonata). Seedlings inoculated with L. bicolor developed mycorrhizae earlier and in greater numbers than the other inoculation treatments, with red pine out-performing jack pine in both respects. At 34 wk following sowing, seedlings were outplanted on a cleared xeric site in Baraga Co., in Michigan's Upper Peninsula. Seedlings inoculated with C. umbonata failed to form mycorrhizae and were not outplanted. Inoculation treatments did not affect shoot or root weight at outplanting. Red pine inoculated with L. bicolor averaged 21% and 19% greater survival compared with control seedlings after one and two years in the field, respectively. Other inoculation treatments failed to increase seedling survival for either tree species. Jack pine demonstrated higher overall survival than did red pine for both years in all corresponding inoculation treatments.     

Nitrogen retranslocation, allocation, and utilization in bare root Larix olgensis seedlings

We quantified biomass accumulation and nitrogen (N) retranslocation, allocation, and utilization of Changbai larch (Larix olgensis) seedlings subjected to four fertilization treatments (24, 59, 81, 117 kg·ha-1 N) with an unfertilized control during summer and autumn 2009. Ammonium phosphate (18-46-0) was the fertilizer used in all treatments. On both sampling dates, the needles had greater biomass and N content than new (2009) stems and old (2008) stems, and coarse, medium and fine roots (diameters of >5, 2-5 mm, and 0-2 mm, respectively). Higher N concentration was observed in old stems and coarse roots than that in new stems and medium roots. In mid-summer, fine roots had higher N concentration than coarse roots. The treatment with 24 kg·ha-1 N had the greatest biomass and N content in needles and old stems, and highest net N retranslocation (NRA) and amount of N derived from soil. On September 21, no N translocation was observed, while the treatment with 24 kg·ha-1 N had the highest N utilization efficiency and fertilizer efficiency. Vector analysis revealed that all four fertilization treatments induced N excess relative to the control. The treatments with 59, 81, 117 kg·ha-1 N induce N excess compared with treatments at 24 kg·ha-1 N. We conclude that the traditional local fertilizer application rates exceeded N requirements and N uptake ability for Changbai larch seedlings. The application rate of 24 kg·ha-1 N is recommended.     

Intersterilitätsgruppen und Klone von Heterobasidion annosum-Isolaten aus Koniferen- Wurzel- und -Stammfäulen

Intersterility groups and clones of Heterobasidion annosum isolates from root and butt rots of conifers 69 H. annousum heterokaryons from scots pine, Douglas fir, Norway spruce and larch belonged to the P group, three heterokaryons from Norway spruce to the S group. The results of a study with clones of H. annosum isolates from roots of Scotch pine trees in close neighbourhood suggest colonisation by H. annosum (partly) via root contacts.     

The impact of climate,drainage and fertilization on the survival and growth of pinus sylvestris L. in afforestation of open,low‐production peatlands

The objective of this study was to analyse the conditions for forest production on open, low‐production peatlands in Sweden with respect to climate, and water and nutrient regimes. The study focused on survival and growth of Scots pine (Pinus sylvestris L.) seedlings, planted 18 yrs ago in five experimental areas evenly distributed between south and north Sweden. Different ditch spacing and NPK fertilizer treatments were combined systematically in all experiments. Survival was positively correlated with temperature sum, fertilization and drainage intensity. Tree growth was not influenced by variations in temperature sum after merely draining, but in combination with fertilization, growth was strongly correlated with climate. In the southern experiments, fertilization increased stand growth eight to nine times, whereas stands on the northern sites did not respond to fertilization. The most important fertilizer element was P. The application of N had no effect on growth. More intensive drainage increased stand growth by 60%. In the southern areas, height development in the most intensive drainage and fertilizer treatments indicates a mean annual increment of 6–7 m³ ha^?1, and no sign of decline in growth was seen. Turf‐planting had positive effects on both survival and growth, especially in less intensively drained plots. A large proportion of damaged trees was observed in the experiment. The frequency of damaged trees was positively correlated with treatment intensity but negatively correlated with temperature sum. The results show clearly that merely draining is not sufficient for successful afforestation of low‐production peatland sites. Fertilization by P and K is a necessary prerequisite, but the effect of fertilization varies with climatic conditions, probably owing to the amount of plant‐available N.     

The effect of weed control and fertilization on survival and growth of four pine species in the Virginia Piedmont

The growth response of loblolly pine (Pinus taeda L.), shortleaf pine (Pinus echinata Mill.), Virginia pine (Pinus virginiana Mill.), and white pine (Pinus strobus L.) to weed control and fertilization in the Piedmont of Virginia was assessed. Four different silvicultural treatments were evaluated: (1) check (no treatment); (2) weed control; (3) fertilization; (4) weed control plus fertilization. The weed control treatment included a series of herbicide and mechanical treatments to eliminate competing hardwoods. The fertilizer treatments added N, P, K, and S. Survival and growth was measured annually through age 5. There were significant differences in survival and growth among species. Survival was greatest for loblolly pine, lower in shortleaf and Virginia pine, and lowest in white pine. Fertilization without controlling the competing hardwoods decreased survival in all planted pines due to the increased hardwood competition. Loblolly pine was tallest through the 5-year period, shortleaf and Virginia pine were shorter and white pine was shortest. Silvicultural treatments had no impact on tree height but significantly affected DBH. Weed control increased DBH while fertilization did not. When applied in combination with weed control, there was no additional increase in growth of the pines due to fertilization beyond that from weed control only. Fertilization stimulated the growth of the competing hardwoods which were significantly taller in the fertilized plots.     

Results of long-term K and Mg fertilizer experiments in afforestation

Results from long-term fertilizer experiments on Scots pine (Pinus sylvestris) planted on old arable land in the north-eastern German lowlands demonstrate the necessity for fertilization with KMg on poorly buffered sandy soils.Applying KMg to young Scots pine increased pole production over a 60 year period by a total of 129 m³ ha⁻¹ (26%). In other experiments the improvement was as much as 93%.KMg manuring on light soils ensures optimal K and Mg nutrition and creates the conditions for uninterrupted growth and healthy development of the plantation. On degraded soils, KMg treatment leads to improved utilisation of the low soil nitrogen supply.With increasing eutrophication due to atmospheric nitrogen, KMg fertilization overcomes the depletion of soil cations and resulting imbalance in tree metabolism.Adequate application of KMg is a prerequisite for success in underplanting with beech (Fagus silvatica) on soils with a land use classification below 30, increasing the growth in height of beech by about 30%.KMg fertilizer favours biomass production and thus the fixation of N deposited through the atmosphere by the vegetation. Thereby, KMg fertilizer use and other silvicultural measures reduce the negative effects of atmospheric N-input in forest ecosystems to a certain extent, especially the N-pollution of groundwater. However, they cannot replace the technologies to reduce the N emissions.     

Infection experiments with Gremmeniella abietina on seedlings of Norway spruce and Scots pine

Conidia of Gremmeniella abietina infected and caused disease symptoms in annual shoots of both Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) seedlings. In Norway spruce shoots the infection remained largely latent, with only a few seedlings showing symptoms. Mycelial growth inside the shoots was faster in Scots pine than in Norway spruce and was favoured by low temperature in both hosts. The shoots of Norway spruce seedlings had higher endophyte populations than those of Scots pine, and the populations were decreased by low temperatures. Reductions in the normal epiphytic or endophytic flora by acid mist treatments seemed to favour the development of G. abietina.