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.     

Studies of the life cycle of Gremmeniella abietina on Scots pine in southern Sweden

Aspects of the life cycle of Gremmeniella abietina (Lagerb.) Morelet were studied from 1988 to 1990 in stands of Pinus sylvestris L., 16–32 years old, in southern Sweden, initiated in 1988 with a widespread outbreak of the disease. Pycnidia started to release conidia in late spring and apothecia began to release ascospores in summer. Latent infections could still be detected one year after their establishment by cultivation of healthy looking shoots on agar petri dishes. G. abietina appeared to have a mainly biennal life cycle, as most spores were released two years after infection of the shoot.     

Effects of nickel and copper on growth and mycorrhiza of Scots pine seedlings inoculated with Gremmeniella abietina

Scots pine (Pinus sylvestris) seedlings were planted in soil originating from two localities with different background levels of nickel and copper. In addition, some of the seedlings were exposed to additional nickel (20.5 mg Ni/l of soil) or copper (63.5 mg Cu/l of soil), or a combination of both Ni and Cu, via soil without direct shoot exposure during their second growing period. The seedlings were either irrigated with spring water (pH 6) or got only natural rain during the whole field experiment. All seedlings were inoculated with conidia of a shoot‐pathogen Gremmeniella abietina during their third growing season, and harvested the following spring. Lengths of shoots of different year‐classes were used as growth estimates. In roots, the proportion of fungal (assumedly mycorrhizal) biomass was estimated by measuring ergosterol concentration. Guajacol peroxidase activity was measured. Short roots were classified into two groups according to their condition and the composition of the mycorrhizal community was expressed as a proportion of morphotypes in the roots. The seedlings exposed to additional Ni had higher shoot growth than the seedlings in the other treatments. The mean Ni concentration in the roots of seedlings exposed to additional Ni was 79 p.p.m. and in other seedlings 16 p.p.m. Additional Ni also decreased the frequency of clearly senescent short roots and the proportion of the mycorrhizal morphotype with the thinnest mantle. These results indicate that the Ni exposure levels used in this experiment had some positive effects on the seedlings. The relative fungal biomass was about 6% lower (p = 0.0981) in the fine roots of seedlings treated with additional Cu. The mean Cu concentration in the roots of seedlings exposed to additional Cu was 256 p.p.m. and in other seedlings 29 p.p.m. Peroxidase activity, which was used as a general stress indicator in this study, was not affected by any of the treatments. The shoot growth and the relative biomass of fungi in the fine roots were positively correlated in all seedlings, and this correlation was stronger in seedlings exposed to additional Ni that were not irrigated compared with seedlings not exposed to additional Ni that were irrigated. The frequency of asymptomatic infections of G. abietina was positively correlated with the proportion of senescent short roots in the irrigated seedlings but not in not‐irrigated seedlings. The general condition of seedlings may be an important factor for infection by G. abietina when moisture is high enough for the fungi to infect seedlings by conidia.     

Incidence of Gremmeniella abietina in planted seedlings of Picea abies and Pinus sylvestris in northern Finland

The ascomycete Gremmeniella abietina causes a disease (scleroderris canker) on conifers. The pathogen kills terminal shoots and branches of Pinus sylvestris and Picea abies, sporulates on host shoots and causes leader changes (i.e. dieback). Damage caused by G. abietina was investigated in young P. abies and P. sylvestris plantations in northern Finland. Side branches from seedlings of both species and the main stems of P. sylvestris were collected from three sites in two locations (Poksa 1 and 2, Kivalo) in northern Finland. The number of cankers and leader changes was counted in branches of each age (i.e. year) in a total of ca. 6300 shoots of P. abies and 1200 shoots of P. sylvestris. Cankers were common on both P. abies and P. sylvestris in Poksa 1 (2002–2008) and on P. abies in Poksa 2 (1998–2001). In Kivalo, cankers occurred only sporadically on P. sylvestris. Leader changes were most frequent in 1999–2009 in Poksa 1, in 2001–2009 in Poksa 2 and in 2003 in Kivalo. Cankers and other symptoms of infection were more obscure on P. abies than on P. sylvestris. According to both conidial morphology and molecular analysis, the strain of G. abietina infecting both P. abies and P. sylvestris was small‐tree type (STT). This is the first report of G. abietina, STT or B type, injuring P. abies in plantations.     

Occurrence of Gremmeniella abietina damage on Scots pine along a pollution gradient from Monchegorsk nickel smelter to western Lapland

Gremmeniella abietina damage was studied at stand and tree level on the Kola Peninsula and in northern Finland. At stand level, damage was recorded in four areas in Finland, while only individual damaged trees were found in Russia. According to the results of branch analysis, there was no sign of endemic epidemics in any of the areas. There was a significant negative correlation between the average G. abietina damage and modelled SO₂ deposition and temperature sum along the gradient. The results suggest that G. abietina is distributed all over northern Finland and the Kola Peninsula in natural stands of Scots pine, and that the direct effect of SO₂ deposition from the Monchegorsk nickel smelter on the damage may be minimal.     

Morphological and Ecological Variation of Gremmeniella abietina var. abietina in Pinus sylvestris,Pinus contorta and Picea abies Sapling Stands in Northern Finland and the Kola Peninsula

The morphological and ecological variation of two types of Gremmeniella abietina var. abietina causing scleroderris canker on conifers was investigated in Pinus spp. and Picea sp. sapling stands in northern Finland and the Kola Peninsula. Small - tree type (STT or B type) of G. abietina was detected alone in 13 Scots pine, three lodgepole pine and two Norway spruce sapling stands out of 26 stands investigated, both STT and large - tree type (LTT or A type) were observed in six Scots pine stands, and LTT was detected alone in two Scots pine stands. For the first time, G. abietina was found to injure Norway spruce saplings in a respective plantation in northern Fennoscandia. STT isolates produced statistically significantly more conidia in vitro than LTT isolates. Morphological variation in conidia septation revealed that STT produced conidia with more than five septa more frequently than did LTT. There was a greater range in variation in septation in STT than in LTT, with overlapping between the types. Isolates of both types were equally associated with cankers, coloured wood, pycnidia or apothecia in the infected saplings.     

Survival and vitality of Gremmeniella abietina on Pinus sylvestris slash in northern Sweden

Survival and vitality of Gremmeniella abietina on Pinus sylvestris slash was studied in northern Sweden during 2003 and 2004. Once a month between September 2003 and April 2004, two to three trees were cut down and debranched. Shoots with pycnidia were sampled at the felling date and then at every consecutive month. The percentage of germinated conidia from each shoot was calculated after 24, 48 and 72 h incubation. The vitality of G. abietina pycnidia in the slash remained high the whole period. Intact pycnidia were found on slash several months after the time of conidial sporulation, which indicates that new pycnidia may be produced on dead pine branches. Sampling of shoots from slash on 13‐ to 18‐month‐old clear‐cuts showed conidial germination capacity as high as in pycnidia collected in fresh slash. Due to survival of G. abietina in slash it is recommended to postpone planting of P. sylvestris seedlings in northern boreal areas to the third vegetation period after sanitary clear‐cuts.     

Growth reactions of a Gremmeniella abietina isolate and Scots pine embryogenic tissue cultures differ in a host–parasite in vitro system

Summary The aim of this study was to screen Scots pine embryogenic tissue culture material to Gremmeniella abietina. According to the results Scots pine (Pinus sylvestris) embryogenic tissue lines (27 lines) could be ranked based on significant differences in G. abietina effects estimated by changes in glucosamine concentrations, growth, and dry matter contents of inoculated tissue pieces. The ranking order of the extreme tissue lines stayed basically similar, regardless of the variable measured. These findings imply a possibility to use the ranked embryogenic tissue lines in more detailed investigations of interactions between G. abietina and its host. This is the first report of using embryogenic tissue cultures in studying host–parasite interaction between Scots pine and G. abietina.     

The effect of scalping on seedling establishment after seed tree cutting of Scots pine stands in drained peatlands in northern Finland

We studied the establishment of natural seedlings after seed tree cutting and scalping in two drained Scots pine peatland stands in northern Finland (Simo and Sievi). Approximately, 50?ha^?1 of Scots pine stems were retained on both sites. During the six subsequent years, five seedling surveys were conducted. The effect of site preparation was analysed using generalized linear mixed models (GLMM). The mean density of pine seedlings increased more quickly in the scalped plots and was ca. 1.1?m^?2 after six years compared to 0.87?m^?2 in the non-scalped plots, on average. Scalping increased the number of pine seedlings 2-fold compared to that in non-treated plots in Simo, however, in Sievi non-treated plots had 30% higher pine seedling density after six years. Downy birch seedling density was 9–10-fold compared to that of non-treated plots in both sites. In terms of seedling density, seed tree cutting led to satisfactory pine regeneration after just two growing seasons in drained pine peatlands, even without scalping. Scalping significantly increased the number of pine seedlings, but it also enhanced the germination of birch seedlings strongly.     

Scots pine stand establishment with special emphasis on uncertainty and cost-effectiveness,the case of northern Finland

The effect of uncertainty on the cost-effectiveness of alternative chains of stand establishment in northern Finland was examined. The data were from a reforestation study of Scots pine (Pinus sylvestris L.) consisting of 288 sample plots, which were measured with respect to regeneration success. The study design included four site-preparation methods (patch scarification, ploughing, prescribed burning and disk trenching) combined with three reforestation methods (sowing, planting with containerized seedlings and planting with bare-rooted transplants). Initial reforestation density was 2,500 spots or seedlings per hectare, and the regeneration success was modeled as probability with two thresholds, namely 500 and 1,100 saplings. On formerly spruce-dominated as well as pine-dominated sites the most cost-effective chain was ploughing and planting with containerized seedlings, when threshold was 1,100 saplings per hectare. However, with threshold of 500 saplings the best performer was ploughing and direct sowing on both sites.     

Summer temperature affects the ratio of radial and height growth of Scots pine in northern Finland

? Radial and height growth chronologies from 150-year-old and 50-year old Scots pine stands, both located near to the northern timberline in Laanila, Finland (68° 30′ N, 27° 28′ E), were cross-correlated with each other and with mean temperatures of various temperature periods defined as months, days or growing-degree-days.

? The height-growth chronology correlates significantly with radial growth at lags 1 and 2, and radial growth with height growth at lag 2 when the effect of temperature is omitted. On average, low and high growth years represent cool and warm average growing seasons. The summer temperatures (June, July and August) affect most on growth, but height and radial growth do have a different set of effective temperature periods. Furthermore, July temperature variation affects stronger height than radial growth.

? Those years with low height/radial-growth ratio, i.e. relatively higher current year’s radial than next year’s height growth, do have lower growth and cooler-than-average July temperature whereas those years with high height/radial-growth ratio do have clearly warmer mid-summer temperatures.

     

Effect of half-systematic and systematic thinning on the increment of Scots pine and Norway spruce in Finland

Diameter, height and volume increment of Scots pine (Pinus sylvestrisL.) and Norway spruce (Picea abies (L.) Karst.) stands wereinvestigated in long-term thinning experiments in southern Finland.The measurement period was on average 19 years, and thinningsconsisted of five different treatments: (1) unthinned; (2) selectivethinning from below where thinning intensity was determinedaccording to the number of stems; (3) selective thinning frombelow where thinning intensity was determined according to standbasal area; (4) systematic thinning where corridors were openedat regular intervals in the stand; and (5) half-systematic thinningwhere corridors were opened at regular intervals and the remainingstrips thinned from below, i.e. a combination of selective andsystematic thinning. The selective (basal area) thinning ofScots pine resulted in a volume growth reduction of about 12per cent, but in Norway spruce it resulted in no marked reductioncompared with the unthinned plots. In both tree species, volumeincrement in the half-systematic and selective (basal area)thinning differed only slightly from each other. The systematicthinning resulted in the lowest volume increment. In the systematicthinning, the total yield (m³ ha^–1) somewhat decreasedwith increasing corridor width. In half-systematic thinning,on the other hand, the total yield was not clearly related tocorridor width. A corridor accelerated the diameter incrementof the edge trees. The edge effect progressively diminishedwhen moving deeper into the strip and ended at a distance ofabout 3–4 m from the corridor edge. After the thinning,the height increment first decreased, but later on it recovered.The unthinned plots had more dead trees than the other treatments,but there were no significant differences between the othertreatments. It can be concluded that the prevailing thinningmethod in the Nordic countries, i.e. half-systematic thinning,most probably results in no significant growth and yield losses.     

Effects of timing,duration, and intensity of simulated browsing on Scots pine growth and stem quality

This study examines a large number of structural and growth-related responses of young Scots pine in relation to simulated browsing from large herbivores. We combined different modes of intensity, timing, and duration of clipping for nine consecutive years from 2000 to 2008. Data on tree responses were collected yearly during this period, as well as in 2011, and were analyzed using ordination techniques and generalized linear models. A key result demonstrated is the serious effects of clipping on the youngest pines. Mortality was high when clipping started the first year, indicating that the trees are vulnerable at that stage, but after that increasing intensity and duration of clipping were the main causes of mortality. Among the different treatment factors, intensity of clipping had the largest influence on altering the final growth responses, some of the quality responses, and mortality. Timing of clipping was the most important factor for explaining the occurrence of the majority of responses related to stem structure. Duration of clipping had the largest effect on the responses that included repeated-measures data, such as height development and number of stems. We discuss the long-term consequences of browsing for the trees and the implications for forest management.     

Superoxide dismutase (SOD) activity in Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies L. Karst.) needles in northern Finland

The superoxide dismutase (SOD EC 1.15.1.1) activity in Scots pine (Pinus sylvestris) and Norway spruce (Picea abies L. Karst.) needles in urban and rural trees of northern Finnish populations was studied. Enzyme activity was higher in pine than in spruce needles. Two pine clones from eastern Finland and Lapland revealed great differences in specific SOD enzyme activity.     

Long-term effects of stump harvesting on soil properties and tree growth in Scots pine and Norway spruce stands

Abstract

Effects of stump harvesting on the properties of surface soil and on the density, structure and growth of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) stands were estimated in a field trial in western Finland. The experiment was established in 1977 and measured in 2010. Stems and logging residues were harvested after clear-cutting, and stumps were lifted and removed from half of the experimental area. Sixteen plots were planted with pine seedlings and 16 with spruce. The main effects of stump harvesting were improved survival of planted trees and an increase in natural regeneration. No clearly negative effects were noted in the stand development. Stump harvesting had no or minimal effects on the properties of the organic layer and those of the 0- to 10-cm mineral-soil layer. Soil properties did not differ between tree species. Pine production was higher on plots with stump removal compared to plots without soil treatment.     

Impacts of changing climate on the productivity of Norway spruce dominant stands with a mixture of Scots pine and birch in relation to water availability in southern and northern Finland

A process-based ecosystem model was used to assess the impacts of changing climate on net photosynthesis and total stem wood growth in relation to water availability in two unmanaged Norway spruce (Picea abies) dominant stands with a mixture of Scots pine (Pinus sylvestris) and birch (Betula sp.). The mixed stands were grown over a 100-year rotation (2000-99) in southern and northern Finland with initial species shares of 50, 25 and 25% for Norway spruce, Scots pine and birch, respectively. In addition, pure Norway spruce, Scots pine and birch stands were used as a comparison to identify whether species' response is different in mixed and pure stands. Soil type and moisture conditions (moderate drought) were expected to be the same at the beginning of the simulations irrespective of site location. Regardless of tree species, both annual net canopy photosynthesis (P(nc)) and total stem wood growth (V(s)) were, on average, lower on the southern site under the changing climate compared with the current climate (difference increasing toward the end of the rotation); the opposite was the case for the northern site. Regarding the stand water budget, evapotranspiration (E(T)) was higher under the changing climate regardless of site location. Transpiration and evaporation from the canopy affected water depletion the most. Norway spruce and birch accounted for most of the water depletion in mixed stands on both sites regardless of climatic condition. The annual soil water deficit (W(d)) was higher on the southern site under the changing climate. On the northern site, the situation was the opposite. According to our results, the growth of pure Norway spruce stands in southern Finland could be even lower than the growth of Norway spruce in mixed stands under the changing climate. The opposite was found for pure Scots pine and birch stands due to lower water depletion. This indicates that in the future the management should be properly adapted to climate change in order to sustain the productivity of mixed stands dominated by Norway spruce.     

Effects on stem growth of Scots pine 33 years after thinning and/or fertilization in northern Sweden

Thinning and fertilization are two common and important stand treatments in forest management. In terms of area treated, thinning is the single most common form of stand treatment. The extent of forest fertilization on the other hand, has varied widely in recent decades and is currently not very common. Thinning is done primarily to promote stand properties while fertilization is done to increase growth before future final felling. After thinning stands of Scots pine, overall growth decreases, while growth of residual trees increases. An experiment was established outside Vindeln in northern Sweden where the long-term growth effects after thinning and/or fertilization were evaluated after 33 years. Experimental set-up was a randomized block design including 12 replications of four treatments. Treatments were control, fertilization, thinning, and thinning and fertilization combined. Thinning decreased overall and annual volume growth ha^?1, and increased green crown size and diameter growth at breast height (1.3 m, DBH) for the individual trees. No positive growth responses to fertilization could be seen after 33 years. In summary, this study showed that thinning can have long term effects on the growth of a Scots pine stand in northern Sweden. Possible reasons for the lack of positive response following fertilization are discussed.     

The effects of pine mistletoe (Viscum album subsp. austriacum) on the growth of Scots pine and Crimean pine in Turkey

In this study, the effect of pine mistletoe (Viscum album subsp. austriacum) on basal area increment of Crimean pine and Scots pine was investigated. Dendrochronological data were collected from 223 (71 uninfected and 152 infected) Crimean pines and 195 (77 uninfected and 118 infected) Scots pines located in Kastamonu province of Turkey in 2014. Infected sample trees were classified as light, moderate or severe infection levels. Growth trends and basal area increment loses were compared between uninfected and infected trees for the periods of the last 10, 20 and 30 years. In addition, infection status of forest stands was investigated using temporary sample plots; 27 plots in Crimean pine stands and 26 plots in Scots pine. Results demonstrated that basal area increments were negatively affected by pine mistletoe for both species. Mean basal area increment losses of infected trees for the last decade were determined as 24% for Scots pine and 26% for Crimean pine. Basal area increment losses varied by infection levels (light, moderate and severe) as follows: 25%, 20% and 28% for Scots pines and 20%, 32% and 9% for Crimean pines. Scots pine stands were more severely infected by pine mistletoe than Crimean pine stands. There were negative correlations between number of infected trees and stand density for both species, while positive correlation was detected between the number of infected trees and mean diameter for Scots pine. The results of this study indicate that the pine mistletoe infection has negative effect on radial growth of Scots pine and Crimean pine trees. The results can be an important contribution to the forest management and protection activities in mistletoe-infected stands.     

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.     

Impacts of inoculation with Herpotrichia pinetorum,Gremmenia infestans and Gremmeniella abietina on Pinus nigra subsp. pallasiana and Cedrus libani seedlings in the field

Herpotrichia pinetorum, Gremmenia infestans and Gremmeniella abietina were inoculated onto 2‐year‐old Anatolian black pine (Pinus nigra subsp. pallasiana) and Taurus cedar (Cedrus libani) seedlings planted in a high mountain forest (1800 m a.s.l) in south‐western Turkey, to determine the effects of these fungi during winter. In June, 8 months after inoculation, 39.9% of experimental plants were dead and 20.4% of the surviving plants failed to flush. Gremmeniella abietina and H. pinetorum caused the most fatalities. Prevention of new shoot formation on surviving plants, however, was mainly an effect of G. abietina infections, although many surviving plants inoculated with G. infestans or H. pinetorum also failed to flush. All three pathogens had the potential to severely damage young plants of P. nigra subsp. pallasiana and C. libani growing at high elevations near to forests with heavy inoculum loads. The implications of this finding for P. nigra afforestations at high altitudes in Turkey are discussed. This study is also the first to report that G. infestans can infect and cause disease on young C. libani plants.