Effects of artificial and western spruce budworm (Lepidoptera: Tortricidae) defoliation on growth and biomass allocation of Douglas-fir seedlings

Artificial defoliation has been used commonly to simulate defoliation by insect herbivores in experiments, in spite of the fact that obvious differences exist between clipping foliage and natural defoliation due to insect feeding. We used a greenhouse experiment to compare the effects of artificial and western spruce budworm (Choristoneura occidentalis Freeman) defoliation on the growth and biomass allocation of 3-yr old half-sib seedlings from mature Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco variety glauca] trees that showed phenotypic resistance versus susceptibility to budworm defoliation in the forest. Artificial clipping of buds mimicked the effects of budworm feeding on total seedling biomass when 50% of the terminal buds were damaged. However, artificial defoliation decreased seedling height, relative growth rate of height, and shoot: root ratio more than budworm defoliation, whereas budworm defoliation decreased stem diameter relative growth rate more than artificial defoliation. Half-sib seedling progeny from resistant maternal tree phenotypes had greater height, diameter, biomass, and shoot: root ratio than seedlings from susceptible phenotypes. We concluded that careful artificial defoliation could generally simulate effects of budworm defoliation on total biomass of Douglas-fir seedlings, but that the two defoliation types did not have equal effects on biomass allocation between shoot and root. Further, an inherently higher growth rate and a greater allocation of biomass to shoot versus root are associated with resistance of Douglas-fir trees to western spruce budworm defoliation.     

Variation in budburst phenology of Douglas-fir related to western spruce budworm (Lepidoptera: Tortricidae) fitness

Variation in budburst phenology among individual trees of interior Douglas-fir (Pseudotsuga menziesii var. glauca [Beissn.] Franco) may influence their susceptibility to western spruce budworm (Choristoneura occidentalis Freeman) defoliation. We tested the hypothesis that phenological asynchrony between Douglas-fir and the western spruce budworm is a mechanism of resistance using clones derived from parent trees that showed resistance versus susceptibility to C. occidentalis defoliation in the field. Susceptible clones had earlier budburst phenology compared with resistant clones when they were grown in a common greenhouse environment, demonstrating a genetic basis for parallel phenological differences exhibited by the parent trees. We tested the importance of phenological asynchrony as a factor influencing fitness of C. occidentalis using two different greenhouse bioassay experiments. One experiment compared western spruce budworm performance on equivalent phenological stages of susceptible and resistant clones by matching larval feeding to the columnar (fourth) bud development stage of each clone. Larvae reared on resistant clones had greater realized fitness (i.e., number of F1 offspring produced) than those reared on susceptible clones when the influence of variation in budburst phenology was minimized. In the other experiment, western spruce budworm larvae were placed on all trees on the same date when approximately 50% of all terminal buds in the population were in the yellow (second) budburst stage. Larvae reared on susceptible clones had greater realized fitness than those reared on resistant clones when the influence of phenological asynchrony was expressed. Our results suggest that resistant phenotypes of Douglas-fir have negative effects on survival and reproduction of C. occidentalis under the natural conditions that insects and trees experience in the field. Genetic variation among trees in budburst phenology has an important influence on interactions between the western spruce budworm and Douglas-fir.     

The role of sugars in western spruce budworm nutritional ecology

Abstract.

• 1 The western spruce budworm, Choristoneura occidentalis Freeman, and Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco, have been used to test the hypothesis that variation in levels of foliar sugars form part of the basis for plant resistance to herbivore attack.
• 2 Budworm population growth was evaluated on artificial diets with 2–45% sucrose using a three generation bioassay. Diets with 1.2% and 3.9% N were tested to determine if responses to sugar were dependent on levels of protein. The 3.9% N diets were supplemented with a mineral salt mixture, so they had high levels of N and minerals.
• 3 The response of budworm population growth to sucrose concentrations ≤20% was convex at 1.2% N and flat for 3.9% N. Population growth on the 1.2% N diet, which had levels of N and minerals similar to host foliage, was good with only 2% sugar, but optimal at the 6% sucrose level; the number of F₁, F₂ and F₃ larvae produced declined substantially when sugar was increased to 11% and 20%. On the 3.9% N diets, population growth was equivalent for diets with 2% and 11% sucrose. Sucrose concentrations ≥29% were detrimental to the budworm at both N levels.
• 4 Sugar concentrations in Douglas-fir foliage varied between 5.7% and 18.4%. Thus, results from the 1.2% N experiment indicated that budworm performance was best on diets with sugar concentrations near the lower limit observed for host foliage. This implies that plants with higher foliar sugar may be inferior hosts for the budworm. Field observations supported this conclusion, as putatively resistant Douglas-fir trees had significantly higher levels of sugars in their foliage than nearby paired susceptible trees. Variation in foliar sugars among individual trees may be part of the mechanism in Douglas-fir resistance to C.occidentalis damage.

     

Taste receptor activity and feeding behaviour reveal mechanisms of white spruce natural resistance to Eastern spruce budworm Choristoneura fumiferana

The pattern of feeding of Eastern spruce budworm Choristoneura fumiferana (Clem.) (Lepidoptera, Tortricidae) is compared on foliage from white spruce Picea glauca (Moench) Voss. (Pinaceae) trees previously determined to be susceptible and resistant to defoliation by budworm. No differences are observed in electrophysiological responses from taste sensilla to aqueous extracts of the two foliage types, nor is there a preference for either extract type in a choice test. Acetone extracts from the two foliage types are both preferred to a control sucrose solution, although neither elicits a preference relative to the other. These results suggest that there is no difference in phagostimulatory power of internal leaf contents of the two foliage types. Longer‐term observation of feeding behaviour in a no‐choice situation shows no difference in meal duration, confirming the lack of difference in phagostimulatory power. However, on average, intermeal intervals are twice as long on the resistant foliage, leading to an overall lower food consumption during the assay. This result suggests an anti‐digestive or toxic effect of the resistant foliage that slows behaviour and limits food intake. Previous research has shown that waxes of the resistant foliage deter initiation of feeding by the spruce budworm and that this foliage contains higher levels of tannins and monoterpenes. The data suggest that the resistant foliage contains a post‐ingestive second line of defence against the spruce budworm.     

Determinants of fruit and seed set in Pavonia dasypetala (Malvaceae)

Summary Measurements of foliage quality, physiological, and phenological condition of sample trees were used as independent variables in multiple correlation analyses to determine their effect on female and male spruce budworm larval dry weights. Female budworm from trees high in foliar concentrations of beta-pinene, myrcene and total nitrogen weighed less than those from trees lacking these characteristics. Male budworm from trees high in foliar concentrations of alpha-pinene, myrcene, terpinolene, citronellyl acetate, and bornyl acetate weighted less than those from trees lacking these characteristics. Additionally, relatively vigorous and productive trees tended to be less susceptible (as evidenced by reduced larval weight) to budworm of either sex.     

Interguild interactions between a non-native phloem feeder and a native outbreaking defoliator

1. Competitive and synergistic interactions directly or indirectly drive community dynamics of herbivorous insects. Novel interactions between non-native and native insects are unpredictable and not fully understood. 2. We used manipulative experiments on mature red spruce trees to test interactions between a non-native phloem feeding insect, the brown spruce longhorn beetle (BSLB), and an outbreaking native defoliator, the spruce budworm. We subjected treatment trees to defoliation by three densities of spruce budworm larvae. Treatment trees were: stressed by (i) girdling (to mimic beetle feeding) or (ii) girdling + BSLB before spruce budworm larvae were introduced on branches in sleeve cages. Budworm larvae then fed on foliage and developed to pupation. We assessed all branches for budworm performance, defoliation, shoot production and shoot growth. 3. Shoot length did not differ in response to stress from girdling or BSLB infestation. Neither stress from girdling, nor interactions with BSLB feeding affected spruce budworm performance or defoliation. Intraspecific impacts on performance and defoliation in relation to budworm density were stronger than the effects of tree stress. 4. Prior infestation of red spruce by BSLB in our experimental set-up did not influence spruce budworm performance. BSLB is a successful invader that has blended into its novel ecological niche because of ecological and phylogenetic similarities with a native congener, Tetropium cinnamopterum. 5. Outbreaks by BSLB will not likely impede or facilitate spruce budworm outbreaks if they co-occur. It would be useful to evaluate the reverse scenario of BSLB success after defoliation stress by spruce budworm.     

Larval experience induces adult aversion to rearing host plants: a novel behaviour contrary to Hopkins' host selection principle

1. Hopkins' host selection principle (HSP) states that insects should prefer foliage from their rearing host plant over that of an alternative host. 2. The current study tested whether eastern spruce budworm, Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae), that were laid and developed on, respectively, resistant and susceptible white spruce Picea glauca (Moench) Voss, showed differences in their feeding and oviposition preferences for these two hosts. 3. The data revealed that previous experience of spruce budworm on a host tree type does not influence the host acceptance and feeding behaviour of later larval stages. However, adult budworm reared on resistant white spruce needles preferentially selected susceptible white spruce needles as the host for their progeny, whereas those reared on susceptible needles showed no preference. 4. To the authors' knowledge, this is the first report of an insect showing an oviposition preference for the non‐rearing host plant. This would tend to increase mixing between insects from susceptible and resistant trees. The present results thus argue against Hopkins' HSP and suggest that learned aversion to resistant foliage experienced by larvae is carried into the adult stage.     

A dendroecological reconstruction of western spruce budworm outbreaks (Choristoneura occidentalis) in the Front Range,Colorado, from 1720 to 1986

Radial increment cores from Douglas-fir (Pseudotsuga menziesii) and blue spruce (Picea pungens), defoliated by western spruce budworm (Choristoneura occidentalis), were analyzed by means of dendrochronological methods and compared with cores from undefoliated ponderosa pine (Pinus ponderosa) and lodgepole pine (Pinus contorta) growing on the same sites in the Front Range, Colorado. Extensive deforestation during the gold and silver booms in the second part of the nineteenth century led to dense and almost pure stands of shadetolerant budworm host species. By using the skeleton plot method, the number of trees with clear growth reductions is obtained, thus representing an exact record of forest insect attacks. The analysis of abrupt growth reductions revealed at least nine outbreaks of western spruce budworm between 1720 and 1986, the majority occurring in the nineteenth century. The outbreaks were graphically compared with periods of attack in New Mexico and Colorado which were detected by other scientists employing tree-ring measurement techniques. No increase in the frequency of severe outbreaks during the twentieth century was observed, yet there is some evidence that the most recent outbreak might be the most severe ever recorded. Open Douglas-fir stands on higher sites were more susceptible to heavy budworm attack than dense stands on lower sites. Blue spruce was less frequently and less severely attacked than Douglas-fir. The spatial pattern of historical outbreaks generally was very patchy.     

Behaviors of Western Spruce Budworm Moths (Choristoneura occidentalis) as Defences Against Bat Predation

We investigated potential defense behaviors of adult western spruce budworm (Choristoneura occidentalis), a non-auditive lepidopteran, against bat predation. Although western spruce budworm moths started to fly before sunset, earlier than many species of moths, temporal isolation of flying moths from foraging bats was incomplete as moths were most active after sunset once bats were foraging. Flying C. occidentalis were most active close to their host trees, and thus were isolated from some bat activity because vegetation limits foraging by some bats. Moths mostly flew near the tops of trees, an area that may have a high predation pressure from bats. Resting western spruce budworm spent little time fluttering their wings or crawling, behaviors that are used as cues by gleaning bats. The outbreak nature of this species, in which large numbers of moths are active at one time, may allow dilution effects to reduce predation risk.     

Spruce budworm growth,development and food utilization on young and old balsam fir trees

Laboratory rearing of spruce budworm, Choristoneura fumiferana, in conjunction with field rearing, gravimetric analyses, a transfer experiment, and foliage chemical analyses at six dates during the period of budworm feeding activity indicated that the age of balsam fir, Abies balsamea, trees (70-year-old mature trees or 30-year-old juvenile trees) affected tree suitability for the spruce budworm via the chemical profile of the foliage. Insects reared on old trees had greater survival and pupal weight, shorter development times, and caused more defoliation than those reared on young trees. Young trees were more suitable for the development of young larvae (instars 2–5), while old trees were more suitable for the development of older, sixth-instar larvae. These results were confirmed by the laboratory transfer experiment. Young larvae fed foliage from young trees had higher relative growth rates (RGR), digestibility (AD), and efficiency of conversion of ingested foliage (ECI) than those fed foliage from old trees. These differences appeared to be related to the high N:tannins ratio, and the high contents of P present in young trees during the development of the young larvae. Old larvae fed foliage from old trees had higher relative growth rates, relative consumption rates (RCR), and digestibility of the foliage than those fed foliage from young trees. The high digestibility of the foliage of old trees was compensated for by a lower efficiency of conversion of digested food (ECD), which in turn resulted in no significant effect of tree age on the efficiency of conversion of ingested foliage by old larvae. The low relative consumption rate of old larvae fed foliage from young trees appeared to be related to the low N:tannins ratio, and the high contents of bornyl acetate, terpinolene, and °-3-carene present in young trees during the budworm sixth instar. Variations in these compounds in relation to tree age may serve as mechanisms of balsam fir resistance to spruce budworm by reducing the feeding rate of sixth instar larvae.     

Dendroentomological and forest management implications in the Interior Douglas-fir zone of British Columbia, Canada

Douglas-fir (Pseudotsuga menziesii var. glauca Mirb. Franco) forests in the Interior of British Columbia, Canada, show periodic defoliation due to western spruce budworm (WSB) (Choristoneura occidentalis Freeman) outbreaks. Tree defoliation causes a reduction in radial growth and is therefore visible in tree rings. In this paper, we identify WSB defoliation history, and critically examine the potential for using dendrochronological analysis by comparing tree-ring estimates with insect surveys. WSB defoliation history was investigated using cores from Douglas-fir growing in the Lac du Bois region of the Kamloops Forest District. Years with an abrupt decrease in radial growth were considered as negative pointer years that potentially reflected WSB outbreaks. The comparison with ponderosa pine (Pinus ponderosa Dougl., ex P. & C. Laws.) (a non-host species) permitted differentiation between growth reductions in Douglas-fir due to climatic effects and those due to defoliation by WSB. The dendrochronological data were matched with information reporting visible damage in Forest Insect Disease Survey (FIDS) and British Columbia Ministry of Forest records. Our objective-based method using ring-width measurements from host and non-host chronologies was compared with qualitative techniques based on the software program OUTBREAK. We were able to distinguish seven distinct outbreak events in 300 years of record.     

Nodulating associations among rhizobia and legumes of the genus Glycine subgenus Glucine

Current and one-year-old foliage was collected from sixty-five red spruce trees growing in thirteen stands at different elevations in the Green Mountains of Vermont and Adirondacks of New York. Sample trees were randomly selected from visually healthy trees at each site. Foliage was analyzed for major and minor elements. In July 1984, foliar Ca, Mg, and Zn concentrations were significantly greater at low than at high elevations. In October 1984, Ca, Mg, and Zn concentrations were higher at low elevations and Ca and Mg concentrations varied significantly among locations within elevational groups. Nitrogen concentration was significantly higher in the high-elevation group in July but not in October. The average red spruce foliar Mg concentration at the end of the growing season in the high elevation stands (442 mg kg⁻¹) is much lower than values reported for other mature red spruce stands in the eastern United States.     

Dendroentomological and forest management implications in the Interior Douglas-fir zone of British Columbia,Canada

Douglas-fir (Pseudotsuga menziesii var. glauca Mirb. Franco) forests in the Interior of British Columbia, Canada, show periodic defoliation due to western spruce budworm (WSB) (Choristoneura occidentalis Freeman) outbreaks. Tree defoliation causes a reduction in radial growth and is therefore visible in tree rings. In this paper, we identify WSB defoliation history, and critically examine the potential for using dendrochronological analysis by comparing tree-ring estimates with insect surveys. WSB defoliation history was investigated using cores from Douglas-fir growing in the Lac du Bois region of the Kamloops Forest District. Years with an abrupt decrease in radial growth were considered as negative pointer years that potentially reflected WSB outbreaks. The comparison with ponderosa pine (Pinus ponderosa Dougl., ex P. & C. Laws.) (a non-host species) permitted differentiation between growth reductions in Douglas-fir due to climatic effects and those due to defoliation by WSB. The dendrochronological data were matched with information reporting visible damage in Forest Insect Disease Survey (FIDS) and British Columbia Ministry of Forest records. Our objective-based method using ring-width measurements from host and non-host chronologies was compared with qualitative techniques based on the software program OUTBREAK. We were able to distinguish seven distinct outbreak events in 300 years of record.     

Characteristics of foliar chemistry in a commerical spruce-fir stand of northern New England,USA

Monthly foliage samples were evaluated for elemental composition from red spruce and balsam fir trees in a commercial, low elevation spruce-fir stand at Howland, Maine during the 1987 growing season. Balsam fir showed consistently higher concentrations in A1 and to a lesser extent N, Ca, Mg, and Fe when compared to red spruce. Red spruce exhibited consistently higher Mn and K concentrations. Both species showed a marked seasonal trend in the foliar concentrations of N, P, K, and Ca in current year foliage. For N, P, and K current year foliar concentrations declined rapidly during the early part of the growing season. Calcium concentrations steadily increased during the growing season for both species and age class of needles. Foliar concentrations of N and P in both species suggests that the availability of these nutrients is limited for this site.     

Contribution of adventitious vs initial roots to growth and physiology of black spruce seedlings

Black spruce (Picea mariana [Mill.] BSP) is a boreal tree species characterized by the formation of an adventitious root system. Unlike initial roots from seed germination, adventitious roots gradually appear above the root collar, until they constitute most of mature black spruce root system. Little is known about the physiological role they play and their influence on tree growth relative to initial roots. We hypothesized that adventitious roots present an advantage over initial roots in acquiring water and nutrients. To test this hypothesis, the absorptive capacities of the two root systems were explored in a controlled environment during one growing season. Black spruce seedlings were placed in a double‐pot system allowing irrigation (25 and 100% water container capacity) and fertilization (with or without fertilizer) inputs independent to initial and adventitious roots. After 14 weeks, growth parameters (height, diameter, biomass), physiology (net photosynthetic rate, stomatal conductance, shoot water potential) and nutrient content (N, P, K, Ca and Mg foliar content) were compared. Most measured parameters showed no difference for the same treatment on adventitious or initial roots, except for root biomass. Indeed, fertilized black spruce seedlings invested heavily in adventitious root production, twice as much as initial roots. This was also the case when adventitious roots alone were irrigated, while seedlings with adventitious roots subjected to low irrigation produced initial root biomass equivalent to that of adventitious roots. We conclude that black spruce seedlings perform equally well through adventitious and initial roots, but if resources are abundant, they strongly promote development of adventitious roots.     

Effects of induced water stress on leaf trichome density and foliar nutrients of three elm (Ulmus) species: implications for resistance to the elm leaf beetle

Seedlings of three elm species with variable susceptibility to the elm leaf beetle (Pyrrhalta luteola Müller) (Coleoptera: Chrysomelidae) were subjected to three water stress treatments (no stress, low stress, and high stress) in a greenhouse experiment. The species tested were Ulmus pumila L. (Siberian elm = highly susceptible), U. parvifolia Jacq. (Chinese elm = resistant), and U. americana L. (American elm = intermediate). The seedlings were analyzed for changes in the levels of selected host traits (trichome density, foliar concentration of nitrogen [N], phosphorus [P], potassium [K], calcium [Ca], magnesium [Mg], iron [Fe], and manganese [Mn]), some of which had previously been implicated in resistance to the elm leaf beetle. Density of leaf abaxial surface trichomes (simple, bulbous, and total trichomes) and foliar Fe and Mg concentrations increased significantly in the highly susceptible Siberian elms under water stress. In contrast, stress reduced trichome density in the moderately susceptible American elms, but it had no effect on levels of foliar mineral nutrients. The stress treatments had no influence on host traits in the resistant Chinese elms. The results suggest that environmental stress can alter plant traits that are likely involved in determining resistance of elms to the elm leaf beetle.     

Changes in the forest floor of Canadian southern boreal forest after disturbance

Abstract. The concentrations and contents of organic matter and nutrients in organic deposits on the forest floor were estimated along a 231-yr chronosequence following fire at the southern limit of the boreal forest in eastern Canada. The sampling design was stratified to take into account the variability related to the presence of the principal tree species as well as to the presence of large gaps created by a recent spruce budworm (Choristoneura fumiferana) outbreak. The forest floor showed a steady accumulation of organic matter and total nutrients with time-since-fire and a 50 % decrease in the concentrations of available P and K, but not N (as determined by aerobic incubation). The increase in forest-floor weight was accompanied by an increased storage of available N, Ca and Mg. The availability of N and Ca was more strongly affected by tree species and gaps than by time-since-fire. A high N-availability was observed under Betula papyrifera and in gaps, while high a Ca-availability was found near Populus tremuloides and Thuja occidentalis. In old sites, the forest floor of gaps, created by a recent spruce budworm outbreak, had a necromass similar to that of a young forest, but the low concentrations of available P and K of an old forest.     

Horizontal transmission of the Picea glauca foliar endophyte Phialocephala scopiformis CBS 120377

We have studied the presence of the foliar endophtye of Picea glauca (white spruce) Phialocephala scopiformis CBS 120377 and its affect on the growth of Choristoneura fumiferana (spruce budworm). Here we examine the transmission of this fungus from 50 trees planted in a test field site to 250 P. glauca seedlings planted under the emerging canopies. After 3 y, the endophyte spread to 40 % of these trees (now 20–30 cm) with an average rugulosin (an anti-insect toxin) concentration of 1 μg g^?1. All woody plants within 2 m of the test trees were collected. These were all shown to be negative for P. scopiformis except for some spruce seedlings that arose from seeds (natural generation). This is positive evidence for the horizontal transmission of P. scopiformis and its apparent specificity to P. glauca under field conditions.     

A lagged, density-dependent relationship between jack pine budworm Choristoneura pinus pinus and its host tree Pinus banksiana

Abstract. 1. Survival of newly emerged jack pine budworm Choristoneura pinus pinus is related to the density of available pollen cones (microsporangiate strobili) produced by its host tree, jack pine Pinus banksiana. 2. A 7‐year time series of observations from a plot network in Ontario, Canada, compared the propensity of jack pine to produce pollen cones, τ, on trees that were either defoliated or undisturbed by the jack pine budworm. 3. Non‐defoliated jack pine trees have a high propensity to produce pollen cones. More than one‐third of these trees produced pollen cones in every year of the series. Propensity varied significantly among plots and trees. Temporal patterns in propensity were also highly variable but within a plot propensity was often autocorrelated in time. 4. Defoliation by the jack pine budworm was associated with forest plots composed of the oldest and the largest trees and with the fewest trees per hectare. Within a plot, outbreaks lasted 3 or 4 years although individual trees were only defoliated in 1 or 2 years. 5. The propensity to produce pollen cones in jack pine was reduced in the years after defoliation. The most pronounced reductions in propensity occurred where defoliation was most severe. 6. The reduction in propensity to produce pollen cones resulting from previous defoliation, coupled with the dependence of jack pine budworm survival on the availability of pollen cones, induces a lagged, negative feedback between the density of the consumer and that of its resource. 7. The lagged, density‐dependent relationship between jack pine budworm and its jack pine host contributes to oscillatory dynamics of the jack pine budworm. Comparison of the outbreak behaviour of jack pine budworm with that of the closely related eastern spruce budworm C. fumiferana suggests that differences in the strength of the host‐plant interaction may account for differences in the relative frequency of outbreaks in the respective systems.     

Macroscopic indicators for the retrospective assessment of Norway spruce crown response to stress in the Krkonoše Mountains

The synergistic effect of a number of natural and anthropogenic stress factors is probably responsible for forest decline in Central Europe. The macroscopic indicators of Norway spruce crown status, as well as branch structure transformation, were proposed to reconstruct the retrospective stress response of a tree to stress conditions over a 30-year period. Based on crown defoliation and the proportion of secondary shoots, five categories of tree response to stress factors were introduced: resistant trees (crown defoliation ≤35%, secondary shoots ≤65%); resilient trees (crown defoliation ≤35%, secondary shoots >65%); susceptible moderately transformed trees (crown defoliation >35%, secondary shoots ≤65%); susceptible strongly transformed trees (crown defoliation >35%, 65% < secondary shoots ≤90%); and exhausted trees to the environment (crown defoliation >35%, secondary shoots >90%). These stress response categories are discussed assuming a sink/source relationship on carbohydrate allocation. The relationship between stress response categories and current regenerative processes in the crown, assessed by macroscopic markers on bud development and shoot growth, is outlined. The research has been carried out mainly in three permanent research plots of old autochthonous Norway spruce forest stands in the Krkonoše Mts. (Czech Republic) under different historical stress loads.