Infection of Norway spruce container seedlings by Gremmeniella abietina

First‐ and second‐year containerized Norway spruce seedlings were inoculated with conidia of type A (large tree type) and type B (small tree type) of Gremmeniella abietina var. abietina at different times during the summer. The appearance of symptoms after artificial inoculation and natural infection on spruce seedlings were recorded the following spring and compared with the disease symptoms on Scots pine seedlings. The proportion of diseased seedlings after inoculation reached as high as 80%. The susceptible period during the summer began later on the first‐year seedlings than on the second‐year seedlings, and was similar for the pine seedlings. Susceptibility of first‐year seedlings was highest in August and on second‐year seedlings in July. The accumulated temperature sum, relative humidity and height growth for first‐ and second‐year seedlings was assessed. Natural infection in 2002 caused more disease on pine than on spruce seedlings. Experimental thinning of seedlings had no effect on disease incidence. In a preliminary comparison between the ability of A and B types to cause disease in Norway spruce seedlings, type B caused more damage than type A after inoculation. However, type A caused a high disease frequency in other experiments in this study. Symptoms on Norway spruce seedlings often first occurred in the mid‐section of the shoot, and were similar to those observed on pine seedlings: needles turned brown, starting at the needle base, in the spring following inoculation. On first‐year spruce, diseased needles were shed rapidly, in contrast to a slower rate of shedding on first‐year pine seedlings. Pycnidia developed about 2 years after inoculation (on pine 1 year after inoculation). On Norway spruce seedlings the lower part of the shoot, including the lateral shoots, often remained alive. The experiments show that G. abietina can cause disease on containerized Norway spruce seedlings under nursery conditions in Finland. The coincidence of spore dispersal, seedling susceptibility and predisposing factors are important in disease development.     

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.     

Precipitation and temperature as factors in Gremmeniella abietina epidemics

The combination of temperature and precipitation during the growth season could be correlated with the occurrence of Gremmeniella abietina epidemics in Denmark. The Gremmeniella index value (GIV) is the result of the monthly rain in mm divided by average monthly temperature in centigrade (mm/°C) and it was created to facilitate easy prediction of epidemics in Denmark. The two most severe outbreaks of G. abietina in pine and other conifers in Denmark took place in 1963–64 and 1984–85, following years where low average temperatures and high precipitation in May combined with the same weather patterns in either August or September. These meteorological conditions were not fulfilled from 1874 to 1900, in which period Pinus nigra was pronounced unsuitable for Danish forestry because of recurring problems with fungal disease. However, the earlier epidemics seemed to coincide with cold and rainy weather (high GIV) in August. In addition, severity of epidemics may also depend on the presence of dense stands between 30 and 40 years of age, as high GIV in August has occurred without disease outbreak in periods with low frequency of susceptible stands. Gremmeniella abietina attacks in Denmark are initiated by the climatic conditions expressed by the Gremmeniella index value, although it does not explain the biological process behind this phenomenon. Nevertheless, the method could be useful for other regions, where epidemics of G. abietina may be related to weather.     

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 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.     

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.     

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.     

Virulence of two Finnish Gremmeniella abietina types (A and B)

Thirteen Finnish Gremmeniella abietina isolates of types A and B were compared for differences in their virulence. Three kinds of inoculations were made: one with conidia on young Scots pine (Pinus sylvestris) seedlings and two with mycelia on the stems and shoots of young Scots pine trees. Inoculations with conidia were carried out in August 1992 and inoculations with mycelia were carried out six times between 11 August and 20 October, 1992. The experiments were evaluated in the late spring and early summer of 1993. The results showed that there was a difference in virulence between the two types. In the conidial inoculations type A infected 34.7% and type B infected 11.0% of the inoculated seedlings. For mycelial inoculations with type A the mean canker (stems) and necrosis (shoots) lengths were 19.3 mm and 8.6 mm longer, respectively, than with type B inoculations. In shoot inoculations there also was a clear difference between the two types in the number of such inoculations where no symptoms were observed. For type B shoot inoculations there was no fungal growth in 21.5% whereas for type A inoculations the figure was only 3.7%.     

Rapid living crown reduction caused by Gremmeniella abietina affects foliar nutrient concentrations of Scots pine

The foliar chemistry of diseased and healthy trees was studied one growing season after severe reduction in living crown caused by Gremmeniella abietina in four young Scots pine (Pinus sylvestris) stands. Sample trees were chosen pairwise on the basis of the living crown length: a diseased tree with about 50% live crown reduction and a healthy tree in each pair. Fifteen elements were determined in the youngest healthy needles on the lateral top shoots of each sample tree. Diseased trees had higher foliar boron, manganese and sodium concentrations and lower magnesium, iron, zinc, copper, potassium, nitrogen and sulphur concentrations compared to the healthy trees. Foliar calcium, aluminium, phosphorus, carbon and hydrogen concentrations did not differ between the diseased and healthy trees, except for P and A1 in two of the stands when the stands were analysed separately. Significant correlations between the needle element concentrations and crown ratio (length of the living crown/tree height) were found especially for B (increasing B with decreasing crown ratio) and for Mg, Fe and Zn (decreasing concentrations with decreasing crown ratio). The effect of G. abietina-induced living crown reduction on tree nutrient status and the role of these mineral nutrients in the susceptibility are discussed.     

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.     

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.     

帽儿山地区不同种源樟子松树轮对气候因子的响应

[目的]为了进一步了解不同种源的樟子松在帽儿山地区的生长差异,研究该地区气候因子对樟子松径向生长的影响,本文建立了来自于阿尔山、高峰、红花尔基、卡伦山、图强罕达盖和金山等7个种源樟子松树轮年表。[方法]通过年轮-气候响应分析、相关分析、主成分分析,探讨了不同种源樟子松径向生长与气候因子的响应差异。[结果]表明:7个种源地樟子松树轮年表整体变化趋势基本相似,但局部也有不同之处,除图强种源外,其余6个种源地樟子松在生长初期年轮宽度呈增加趋势。阿尔山、高峰、红花尔基和卡伦山种源地樟子松径向生长与当年2月平均温度显著正相关,罕达盖种源地樟子松径向生长与前一年4月平均温度显著正相关;高峰、图强、卡伦山种源地樟子松径向生长与当年6月降水显著正相关,阿尔山、高峰、金山、卡伦山、红花尓基、图强种源樟子松的径向生长主要与前一年8月降水显著负相关。[结论]多数种源的樟子松在初期径增长较快,与当年2月份的温度和6月份的降水量有正相关的响应性,而与前一年8月份的降水量有负相关的响应性。     

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.     

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.     

Infektionsversuche mit Ascocalyx abietina und Ascocalyx laricina

Infection experiments with Ascocalyx abietina and A. laricina, on cembra pine, mountain pine, and larch between 1979 and 1985 gave very varying results from one year to another. Moderate to very severe disease only occurred after incubation on favourable sites in the subalpine zone. The intensity of disease may depend on local site and microclimatic conditions for infection. Successful infections occurred especially in autumn. An annual infection rate of 10 to 15 percent can endanger an afforestation: in an afforestation in Avers, 50 percent of the trees were killed in some 5 years.     

Susceptibility of six pine species to Gremmeniella abietina isolates from Spain

Inoculation experiments were performed in order to evaluate the virulence of Gremmeniella abietina isolates from Spain on the main pine species planted in the Iberian Peninsula, as well as the influence of seedling age on this virulence. Two different experiments were carried out with four isolates of G. abietina from Spain. The greenhouse experiments consisted of seedling inoculations. One‐ and 2‐year‐old seedlings of the following five pine species were used: Pinus halepensis, Pinus pinea, Pinus pinaster, Pinus sylvestris and Pinus uncinata; also, 1‐year‐old seedlings of P. nigra were inoculated. The relative necrosis length (RNL) caused by the pathogen after 130 days was used as a response variable. The laboratory experiments were performed on 2‐ to 6‐year‐old internodes of the above pine species excluding P. uncinata. The necrosis length after 6 weeks of incubation was measured. The results have shown that all G. abietina isolates were pathogens on seedlings of these six pine species and seedlings of P. halepensis were consistently the most susceptible ones, although it is important to take into account that all the isolates used in the present work were isolated from P. halepensis, the only pine species in Spain where G. abietina has been recovered up to now. The susceptibility of the other pine species depended on the age of the seedlings.     

Infection of Scots pine seedlings by Gremmeniella abietina during summer under different inoculum potential

The connection between natural conidia dispersal of Gremmeniella abietina (A‐type) and the disease incidence and severity in first and second year conifer seedlings of Pinus sylvestris was studied in central Finland. The seedlings where exposed to natural infection for a 3‐week period throughout the growth season, followed by a 3‐week incubation period in a growth chamber to promote symptom expression. In second year seedlings the infection periods in May–June had a similar effect as regards disease outbreak and roughly half of the successful infections occurred during these periods. However, the first year seedlings were more severely diseased after the infection period in late July/early August, although the inoculum density was lower during this time compared with the period in late June/early July. This result is consistent with earlier studies. The effect of age and growth phase of the seedlings, temperature sum and the number of conidia on the disease occurrence is discussed and compared with the results of earlier studies in which artificial inoculation was employed.     

Effects of associated fungi Sclerophoma pythiophila and Cenangium ferruginosum on Gremmeniella abietina dieback in Spain

The interactions between Gremmeniella abietina and either Sclerophoma pythiophila or Cenangium ferruginosum, fungi frequently isolated from diseased twigs along with G. abietina, were studied under laboratory (dual cultures) and greenhouse conditions (double‐inoculations). Virulence of each species was also evaluated in greenhouse experiments by means of single‐inoculations. In vitro interactions were assessed on Petri dishes containing malt agar with pine needle extract, and greenhouse experiments were performed on 1‐year‐old Pinus halepensis seedlings. In vitro growth of G. abietina was inhibited by both fungi when grown in dual culture. In single‐inoculations, G. abietina caused the greatest necrosis length on P. halepensis seedlings, followed by S. pythiophila, whereas C. ferruginosum did not cause significant necrosis. In double‐inoculations, C. ferruginosum was able to reduce the length of necrosis caused by G. abietina on the P. halepensis seedlings. In contrast, necrosis length was greater in seedlings inoculated with both S. pythiophila and G. abietina than in those inoculated with G. abietina alone. Therefore, S. pythiophila seems to play a role in disease expression caused by G. abietina on P. halepensis in Spain.     

CO2 fluxes of a Scots pine forest growing in the warm and dry southern upper Rhine plain, SW Germany

The effects of the warm and dry weather in the southern upper Rhine plain in the southwest of Germany on the carbon balance of the Scots pine forest at the permanent forest meteorological experimental site Hartheim were analysed over a 14-month period. The investigation of the net ecosystem exchange of carbon dioxide (F _NEE) of the Scots pine forest started in the extraordinary hot and dry August 2003. Carbon dioxide fluxes were measured continuously using an eddy covariance system and analysed by use of the EDDYSOFT software package. After determining the temperature dependence of the forest ecosystem respiration and the daytime light dependence of the CO₂ exchange, monthly and annual carbon balances of the Scots pine forest were calculated. Mean peak daytime F _NEE rates observed in August and September 2003 (−6.5±3.6 μmol m⁻² s⁻¹) were drastically lower than in August and September 2004 (−11.8±5.2 μmol m⁻² s⁻¹), which did not show pronounced deviations from the mean long-term (1978–2002) climatic conditions. In August 2003, the Hartheim Scots pine forest was a distinct CO₂ source (35 g C m⁻²). The estimates of the annual carbon sink strength of the Scots pine forest ranged between −132 g C m⁻² (August 2003–July 2004) and −211 g C m⁻² (October 2003–September 2004). The main uncertainty in the determination of the carbon balance of the Hartheim Scots pine forest was introduced by the frequently low turbulence levels, i.e. the friction velocity corrected night-time F _NEE fluxes.     

Enzymatic interactions between Gremmeniella abietina var. abietina,European race,and two resistant hosts,Pinus banksiana and P. contorta

Ultrastructural observations showed that the European race of Gremmeniella abietina var. abietina can secrete several cell wall–degrading enzymes that help the invasion of Pinus banksiana and P. contorta shoot tip tissues. Alterations in the content or distribution of cellulose were obvious, indicating that colonization might be achieved primarily through the action of glucanases. Polygalacturonases seemed more abundant in the cortex and the phloem of the stem, suggesting they are among the first enzymes secreted by the pathogen during infection. Production of laccases and peroxidases was also revealed but always close to fungal cells or in areas where host wall degradation was obvious. In places where there was an accumulation of phenolic compounds in the infected zones of the shoot, particularly in or near the transition zone, immunolocalization of both enzymes was limited or absent. Laccases and polygalacturonases were regularly detected over the extracellular sheath of G. abietina, and this indicates that the sheath could play a significant role during host wall degradation. Finally, examination of G. abietina wall constituents, such as chitin and β‐1,3‐glucans, suggests that the composition of the pathogen wall changes during the infection process. This change may be associated with the secretion of fungal enzymes but could also be linked to host reactions altering the integrity of pathogen cell walls.