Mechanical injury and fungal infection induce acquired resistance in Norway spruce

Norway spruce trees (Picea abies (L.) Karst.) pretreated by wounding and fungal infection showed highly enhanced resistance to a subsequent challenge inoculation with the pathogenic bluestain fungus Ceratocystis polonica (Siem.) C. Moreau. This is the first time the effectiveness of the constitutive and inducible defenses has been shown to depend on prior wounding and infection in conifers, although such acquired resistance has previously been found in several angiosperms. Trees that were pretreated with a combination of 12 bark wounds (1.6 x 10 cm), four fungal inoculations and four sterile inoculations 1-15 days before mass inoculation with C. polonica at 400 inoculations per square meter over a 0.8 m stem section had significantly shorter necroses in the phloem, less bluestained sapwood, and less dead cambium than untreated control trees. Pretreatment with four fungal or sterile inoculations alone did not lead to enhanced resistance. Pretreatment by bark wounding alone seemed to provide an intermediate degree of resistance compared to bark wounding, fungal inoculations and sterile inoculations combined. All trees had a marked increase in the number of resin ducts in the year of inoculation compared with previous years, suggesting that formation of traumatic resin ducts play an important role in the development and maintainance of enhanced resistance.     

Effect of thinning on anatomical adaptations of Norway spruce needles

Conifers and other trees are constantly adapting to changes in light conditions, water/nutrient supply and temperatures by physiological and morphological modifications of their foliage. However, the relationship between physiological processes and anatomical characteristics of foliage has been little explored in trees. In this study we evaluated needle structure and function in Norway spruce families exposed to different light conditions and transpiration regimes. We compared needle characteristics of sun-exposed and shaded current-year needles in a control plot and a thinned plot with 50% reduction in stand density. Whole-tree transpiration rates remained similar across plots, but increased transpiration of lower branches after thinning implies that sun-exposed needles in the thinned plot were subjected to higher water stress than sun-exposed needles in the control plot. In general, morphological and anatomical needle parameters increased with increasing tree height and light intensity. Needle width, needle cross-section area, needle stele area and needle flatness (the ratio of needle thickness to needle width) differed most between the upper and lower canopy. The parameters that were most sensitive to the altered needle water status of the upper canopy after thinning were needle thickness, needle flatness and percentage of stele area in needle area. These results show that studies comparing needle structure or function between tree species should consider not only tree height and light gradients, but also needle water status. Unaccounted for differences in needle water status may have contributed to the variable relationship between needle structure and irradiance that has been observed among conifers.     

Invasion potential of Agrilus planipennis and other Agrilus beetles in Europe: import pathways of deciduous wood chips and MaxEnt analyses of potential distribution areas

Bark‐ and wood‐boring beetles in the genus Agrilus (Coleoptera: Buprestidae) can survive wood‐chipping, and Agrilus planipennis has established in North America and European Russia with devastating impacts on forest ecosystems. The work presented in this paper combined import statistics of deciduous wood chips, Maximum entropy modelling (MaxEnt) of climatic similarities, and the distribution of potential tree hosts to predict the likelihood of four selected North American Agrilus species to become introduced and established in Europe. In agreement with the EU's energy‐policy target of increased use of wood chips, there was a linear or exponential increase in European imports of deciduous wood chips during the past 10 years from countries harbouring potentially harmful Agrilus species. MaxEnt showed high environmental suitability in Europe for the four selected Agrilus species, particularly in Eastern Europe and European Russia for A. anxius, A. bilineatus and A. planipennis and in southern Europe for A. politus. Documented susceptible host trees are widely distributed in the predicted areas of Agrilus distribution in Europe, and these areas receive large quantities of deciduous wood chips from countries where these and other Agrilus species are present. Thus, it was concluded that the fundamental conditions for introduction and establishment of Agrilus species in Europe are in place.     

Anatomical-based defense responses of Scots pine (Pinus sylvestris) stems to two fungal pathogens

We investigated the cellular responses of stem tissues of mature Scots pine (Pinus sylvestris L.) trees to inoculations with two fungal pathogens. The bark beetle vectored fungus, Leptographium wingfieldii Morelet, induced longer lesions in the bark, stronger swelling of polyphenolic parenchyma cells, more polyphenol accumulation and increased ray parenchyma activity compared with the root rot fungus, Heterobasidion annosum (Fr.) Bref., or mechanical wounding. Axial resin ducts in the xylem are a general feature of the preformed defenses of Scots pine, but there was no clear induction of additional traumatic axial resin ducts in response to wounding or fungal infection. The anatomical responses of Scots pine to pathogen infection were localized to the infection site and were attenuated away from bark lesions. The responses observed in Scots pine were compared with published studies on Norway spruce (Picea abies (L.) Karst.) for which anatomically based defense responses have been well characterized.     

Pathogenicity of four blue-stain fungi associated with aggressive and nonaggressive bark beetles

ABSTRACT The pathogenicity of two isolates of each of four bark beetle-associated blue-stain fungi was evaluated after mass inoculation of about 40-year-old Norway spruce trees (Picea abies). Trees were inoculated with a different isolate of each fungus in 1995 and 1996 at a density of 400 inoculations per m(2) in a 1.2-m-wide band on the lower bole (about 270 inoculations per tree). Trees were felled 15 weeks after inoculation. In 1995, Ceratocystis polonica was the only fungus that had stained the sapwood (56.3% of cross-sectional sapwood area). It induced five times longer phloem necroses, 21 times more dead cambium, and 11 times more dead phloem than any other fungus. In 1996, C. polonica induced less extensive host symptoms and an unidentified Ambrosiella sp. induced comparable symptoms to C. polonica in the phloem and cambium. No trees showed any foliar symptoms 15 weeks after inoculation, but six out of eight trees inoculated with C. polonica in 1995 had only 0 to 25% functional sapwood and probably would have died if felling had been delayed. This study confirms that C. polonica, an associate of the aggressive bark beetle Ips typographus, is pathogenic to Norway spruce. The pathogenicity of the Ambrosiella sp., which is associated with a nonaggressive bark beetle, seems moderate and varies between isolates. The two remaining fungi included in this study (Ophiostoma piceae and a dark fungus with sterile mycelium), which are associated with nonaggressive bark beetles, were nonpathogenic in both experiments. These results are consistent with the hypothesis that aggressive bark beetle species vector virulent fungi that may help them kill trees, but the results also show that some nonaggressive bark beetles may vector phytopathogenic fungi.     

Fungal Infection and Mechanical Wounding Induce Disease Resistance in Scots Pine

Scots pine trees (Pinus sylvestris) recovering from a 90–100% defoliation 2–3 years previously were pretreated with small mechanical wounds or inoculations with the blue-stain fungi Leptographium wingfieldii and Ophiostoma canum. Pretreated trees were less susceptible to a subsequent massive inoculation with L. wingfieldii than untreated control trees, which were extensively colonised by the mass-inoculation. A low pretreatment dosage of L. wingfieldii was somewhat more effective in inducing disease resistance than a higher dosage. Pretreatment with L. wingfieldii, O. canum, and mechanical wounding were about equally effective inducers of resistance in Scots pine, even though L. wingfieldii is known to produce much more extensive phloem necrosis than the other pretreatments. Thus, the strength of the induced resistance response did not depend on the amount of host tissues that was destroyed by the pretreatment. Previously, induced disease resistance has been demonstrated in Norway spruce (Picea abies), and the present study shows that similar responses can be activated in Scots pine.     

Altered light conditions following thinning affect xylem structure and potential hydraulic conductivity of Norway spruce shoots

Trees must respond to many environmental factors during their development, and light is one of the main stimuli regulating tree growth. Thinning of forest stands by selective tree removal is a common tool in forest management that increases light intensity. However, morphological and anatomical adaptations of individual shoots to the new environmental conditions created by thinning are still poorly understood. In this study, we evaluated shoot morphology (shoot length, needle number, projected leaf area) and anatomy (tracheid lumen area, tracheid number, tracheid dimensions, xylem area, potential hydraulic conductivity) in three Norway spruce (Picea abies/L./Karst.) families exposed to different thinning regimes. We compared shoot characteristics of upper-canopy (i.e. sun-exposed) and lower-canopy (i.e. shaded) current-year shoots in a control plot and a plot thinned to 50 % stand density the previous year. One tree per family was chosen in each treatment, and five shoots were taken per canopy position. We found that upper-canopy shoots in both plots had higher values than lower-canopy shoots for all studied parameters, except lumen roundness and tracheid frequency (i.e. tracheid number per xylem area). Thinning had little effect on shoot morphology and anatomy 1 year after thinning, except for small but significant changes in tracheid dimensions. Needles were more sensitive to altered light conditions, as projected leaf area of shoot, needle number and leaf hydraulic conductivity changed after thinning. Differences between upper- and lower-canopy shoots did not seem to be influenced by thinning and were almost the same in both plots. Our results suggest that lower-canopy shoots require several years to modify their morphology and anatomy to new light conditions following thinning. The slow light adaptation of the lower canopy may be of practical importance in forest management: thinned stands may be predisposed to drought stress because newly exposed shoots experience increased illumination and transpiration after thinning.     

Methyl jasmonate treatment of mature Norway spruce (Picea abies) trees increases the accumulation of terpenoid resin components and protects against infection by Ceratocystis polonica, a bark beetle-associated fungus

When conifers such as Picea abies Karst. (Norway spruce) are attacked by insects or pathogens, they often respond by producing increased quantities of terpenoid oleoresin. This response can be mimicked in young P. abies seedlings by treatment with methyl jasmonate (MJ). In this study, we determined the effects of MJ on terpenoids and other chemical defenses of mature P. abies, and investigated whether this treatment protected trees against attack by the blue-stain fungus Ceratocystis polonica (Siem.) C. Moreau, the most important fungal associate of the spruce bark beetle Ips typographus L. Methyl jasmonate treatment induced the formation of traumatic resin ducts in the developing xylem, enhanced resin flow and stimulated increased accumulation of monoterpenes, sesquiterpenes and diterpene resin acids. However, only minor changes were detected in terpene composition in response to MJ treatment and no changes in soluble phenolic concentration were measured. There was much variability in the timing and degree of response to MJ among clones. The observed chemical and anatomical changes in response to MJ treatment were correlated with increased resistance to C. polonica, suggesting that terpenoid oleoresin may function in defense against this pathogen.     

Forest land-use history affects the species composition and soil properties of old-aged hillock forests in Estonia

Decisions regarding forest typology, management and protection are often based on the structures of present-day forests, ignoring their successional history. Forests growing on kames, eskers and various moraine hillocks common in regions with Holocene glaciation are good examples of this approach. In Estonia, these forests locally persist as fragments of continuous primary forest, but usually they are situated on former slash-and-burn areas (bushlands) or reforested agricultural land. Our aim was to elucidate the strength of the effect of long-term land-use history on the present-day vegetation compositions of mature hillock forests and their soil chemistry. It appeared that even the mature secondary hillock forests are still distinct from historically continuous stands in terms of species composition. We discovered connections between stand history and species content in hillock forests as well as transformed soil properties. The carbon and nitrogen contents in the humus horizons of secondary forests are lower while their carbon–nitrogen ratios are higher than in continuous forests. The relationship between vegetation and stand history is demonstrated by the higher proportions of anthropophytic and apophytic species in the herb layer of the secondary forests. The presence of species that are tolerant of anthropogenic impact on the secondary hillock forests floor can also be partly explained by the effect of different species in the tree and shrub layers, gaps in the tree canopy, and the boundary effect caused by the small areas of forest patches, neighboring grasslands or fields. The extinction debt in secondary communities should also be considered.