Tree recovery during the aftermath of an outbreak episode of the Hungarian spruce scale in southern Sweden

In 2010, the first, and so far only, infestation of the Hungarian spruce scale (Physokermes inopinatus) and accompanying sooty mould occurred in Scania, southernmost Sweden. About 1000?ha of Norway spruce (Picea abies) were affected, and the trees suffered from the sucking of the insects as well as from the dense sooty mould that covered the needles. Salvage cuttings were carried out in many of the massively attacked forest stands, both in response to the fear that the trees otherwise would die, e.g. from secondary bark beetle attacks and to prevent spreading of the infestation. The aim of this study was to provide basic, quantitative knowledge on the aftermath response of trees that were heavily infested, but not exposed to salvage cutting. Growth characteristics, in terms of needle weight, shoot length and tree-ring size were measured on infested and uninfested trees to compare and contrast the spruce growth before, during and after the scale outbreak. The infestation resulted in dwarf annual shoots, stunted needles and thin tree rings. The needle weight returned to normal the following year, whereas shoot length and tree rings required one growing season before full recovery.     

Defoliation and growth relationships for mid-rotation Sitka spruce attacked by the green spruce aphid, Elatobium abietinum (Walker) (Homoptera: Aphididae)

Green spruce aphid (Elatobium abietinum) feeds primarily on the 1-year-old and older needle leaves of spruce (Picea spp.) and is a major defoliator of commercial Sitka spruce plantations in the UK and other maritime regions of north-west Europe. The impact of E. abietinum on mid-rotation (23-28-year-old) Sitka spruce in Radnor Forest, in mid-Wales, was determined by comparing aphid population densities, defoliation and growth between plots of trees that were either treated with insecticide or were left untreated. The experimental treatments were maintained for 5 years and over this period (2002-2006) peak E. abietinum densities in the untreated plots varied from 5 to 36 aphids per 100 needles. These densities, which were representative of low to moderate rates of infestation, were associated with low rates of defoliation (0-8%), but they had a significant (P ? 0.01) impact on mean radial increment (RI) and mean volume increment (VI). In 2005, the year with the highest aphid populations, peak densities in the untreated plots averaged 14 aphids per 100 needles and this rate of infestation reduced RI by 17%, VI by 10% and the dry weight of current-year needles by 10%. On average, across all years, infestation by E. abietinum reduced mean annual VI by 6%. Comparisons with previous studies on the impact of the aphid on 4-year-old, 7-year-old and 9-year-old Sitka spruce (Straw et al., 2005) indicate that defoliation by E. abietinum, on a per capita basis, decreases as trees grow older, but that the impact on VI increases. The different growth response of young and mid-rotation Sitka spruce to infestation is related to differences in canopy structure, particularly in the ratio of current-year needles to older needles, and the greater demands on photosynthetic production in older trees that arise from the need to support an increasing quantity of non-photosynthetic structural tissues in branches, stem and roots.     

红皮云杉幼树不同光环境下生理生态特征研究

为了探讨红皮云杉光舍特征与树木枝叶分布格局，生物量及其分配的关系，采用人工控制光照环境，在4个光照条件下进行红皮云杉生长试验，结果表明，红皮云杉幼树瞬时最大净光合速率以透光70％处理最大，10％处理最小，对数据进行方差分析，结果差异显著；并且净光合速率最大的处理其暗呼吸速率也最大，光补偿点和光饱和点也随透光率而变化，透光率小(10％)的处理光补偿点也小，说明云杉幼树对庇荫环境的适应；不同光照条件下幼树的形态特征也表现出明显差异，长期生活在相对较暗的光环境中，形成细而高的树干，侧枝数量增加、长度加长，侧枝角度增大；云杉小苗(9年生以前)以相对照度70％处理生物量最大，9年生以后以全光区生物量最大，说明云杉在9年生以前耐庇荫，9年生以后喜光。     

Photosynthesis in Norway spruce seedlings infected by the needle rust Chrysomyxa rhododendri

Chrysomyxa rhododendri (DC.) De Bary is a needle rust with a host shift between Rhododendron sp. and Norway spruce (Picea abies (L.) Karst.), penetrating only the new developing flushes of the conifer. Because little is known about its effects on trees, we investigated several parameters related to photosynthesis in artificially infected 3-year-old Norway spruce seedlings. The potential efficiency of photosystem II (PSII; derived from chlorophyll fluorescence measurements) was reduced in infected current-year needles as soon as disease symptoms were visible, about three weeks after inoculation. Two weeks later, photosynthetic O(2) evolution (P(max)) of infected needles was less than 20% of control needles, whereas respiratory O(2) uptake (R(D)) was about three times higher than that of control needles. Nonstructural carbohydrate concentrations were about 60% of control values in all parts of the shoots of infected trees. Photosynthetic inhibition was associated with marked decreases in chlorophyll concentration and chlorophyll a/b ratio but only a small reduction in carotenoid concentration. In infected trees, P(max) of noninfected 1-year-old and 2-year-old needles was 50 and 80% higher than in the corresponding age class of needles of control trees. Estimation of potential daily net dry mass production, based on P(max), R(D), specific leaf area, carbon content and needle biomass, indicated that seedlings infected once were able to produce 60%, and those infected twice only 25%, of the dry mass of controls. We conclude that afforestation and regeneration of Norway spruce is seriously impaired in regions where seedlings are frequently attacked by Chrysomyxa.     

Carbon and nitrogen cycling immediately following bark beetle outbreaks in southwestern ponderosa pine forests

Bark beetle infestation is a well-known cause of historical low-level disturbance in southwestern ponderosa pine forests, but recent fire exclusion and increased tree densities have enabled large-scale bark beetle outbreaks with unknown consequences for ecosystem function. Uninfested and beetle-infested plots (n = 10 pairs of plots on two aspects) of ponderosa pine were compared over one growing season in the Sierra Ancha Experimental Forest, AZ to determine whether infestation was correlated with differences in carbon (C) and nitrogen (N) pools and fluxes in aboveground biomass and soils. Infested plots had at least 80% of the overstory ponderosa pine trees attacked by bark beetles within 2 years of our measurements. Both uninfested and infested plots stored ∼9 kg C m⁻² in aboveground tree biomass, but infested plots held 60% of this aboveground tree biomass in dead trees, compared to 5% in uninfested plots. We hypothesized that decreased belowground C allocation following beetle-induced tree mortality would alter soil respiration rates, but this hypothesis was not supported; throughout the growing season, soil respiration in infested plots was similar to uninfested plots. In contrast, several results supported the hypothesis that premature needlefall from infested trees provided a pulse of low C:N needlefall that altered soil N cycling. The C:N mass ratio of pine needlefall in infested plots (∼45) was lower than uninfested plots (∼95) throughout the growing season. Mineral soils from infested plots had greater laboratory net nitrification rates and field resin bag ammonium accumulation than uninfested plots. As bark beetle outbreaks become increasingly prevalent in western landscapes, longer-term biogeochemical studies on interactions with other disturbances (e.g. fire, harvesting, etc.) will be required to predict changes in ecosystem structure and function.     

Stand characteristics and downed woody debris accumulations associated with a mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak in Colorado

Lodgepole pine (Pinus contorta Dougl. ex Loud.)-dominated ecosystems in north-central Colorado are undergoing rapid and drastic changes associated with overstory tree mortality from a current mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak. To characterize stand characteristics and downed woody debris loads during the first 7 years of the outbreak, 221 plots (0.02 ha) were randomly established in infested and uninfested stands distributed across the Arapaho National Forest, Colorado. Mountain pine beetle initially attacked stands with higher lodgepole pine basal area, and lower density and basal area of Engelmann spruce (Picea engelmannii [Parry]), and subalpine fir (Abies lasiocarpa (Hook.) Nutt. var. lasiocarpa) compared to uninfested plots. Mountain pine beetle-affected stands had reduced total and lodgepole pine stocking and quadratic mean diameter. The density and basal area of live overstory lodgepole declined by 62% and 71% in infested plots, respectively. The mean diameter of live lodgepole pine was 53% lower than pre-outbreak in infested plots. Downed woody debris loads did not differ between uninfested plots and plots currently infested at the time of sampling to 3 or 4–7 years after initial infestation, but the projected downed coarse wood accumulations when 80% of the mountain pine beetle-killed trees fall indicated a fourfold increase. Depth of the litter layer and maximum height of grass and herbaceous vegetation were greater 4–7 years after initial infestation compared to uninfested plots, though understory plant percent cover was not different. Seedling and sapling density of all species combined was higher in uninfested plots but there was no difference between infested and uninfested plots for lodgepole pine alone. For trees ≥2.5 cm in diameter at breast height, the density of live lodgepole pine trees in mountain pine beetle-affected stands was higher than Engelmann spruce, subalpine fir, and aspen, (Populus tremuloides Michx.), in diameter classes comprised of trees from 2.5 cm to 30 cm in diameter, suggesting that lodgepole pine will remain as a dominant overstory tree after the bark beetle outbreak.     

White spruce regeneration following a major spruce beetle outbreak in forests on the Kenai Peninsula, Alaska

Between 1987 and 2000, a spruce beetle (Dendroctonus rufipennis) outbreak infested 1.19 million ha of spruce (Picea spp.) forests in Alaska, killing most of the large diameter trees. We evaluated whether these forests would recover to their pre-outbreak density, and determined the site conditions on which spruce germinated and survived following the spruce beetle outbreak in forests of the Anchor River watershed, Kenai Peninsula, Alaska. White spruce (Picea glauca) and Lutz's spruce (Picea × lutzii), a hybrid between white and Sitka spruce (Picea sitchensis), dominate the study area. We measured the pre- and post-outbreak density of spruce in 108 3 m × 80 m plots across the study area by recording all live trees and all dead trees >1.5 m tall in each plot. To determine the fine scale site conditions on which spruce germinated and survived, we measured ground surface and substrate characteristics within 20 cm circular plots around a subset of post-outbreak spruce seedlings. The density of post-outbreak spruce (855/ha) was adequate to restock the stands to their pre-outbreak densities (643/ha) for trees >1.5 m tall. We could not accurately estimate recovery for pre-outbreak spruce seedlings because dead seedlings may have decayed in the 5–18 years since the beetle outbreak occurred. At the fine scale, spruce that germinated post-outbreak grew on a wide variety of substrates including downed log, stump, mesic organic mat, peat, hummocks and mineral soil. They exhibited a strong preference for downed logs (53%) and stumps (4%), and most (91%) of the downed logs and stumps that spruce rooted on were heavily decayed. This preference for heavily decayed logs and stumps was especially evident given that their combined mean cover was only 2% in the 3 m × 80 m plots. Within the 3 m × 80 m plots, spruce seedling survival was negatively correlated with bluejoint (Calamagrostis canadensis) litter cover.     

Seasonal nutrient dynamics in white pine and white spruce in response to environmental manipulation

Seasonal retranslocation in white pine (Pinus strobus L.) and white spruce (Picea glauca (Moench) Voss) was examined in response to silvicultural treatments (scarification, annual fertilization application, and annual control of competing vegetation with herbicide) that changed both environmental conditions and the growth rate of the trees. Four years after plantation establishment and initial treatment, nutrient accumulation in current-year needles of white pine and retranslocation from 1-year-old needles were increased following the vegetation control treatment, which increased resource availability (nutrients, water and light) and, hence, growth rate. Nutrient accumulation also increased in current-year white spruce needles following the same treatment, whereas retranslocation decreased in 1-year-old white spruce needles. Correlations of retranslocation (N, P and K) with growth rate (shoot biomass increment) showed a strong positive relationship for white pine and a negative relationship for white spruce. Retranslocation of K was correlated with foliar and soil K concentrations; the availability of this nutrient was also significantly reduced by vegetation control. A general theory for the control of nutrient retranslocation in conifers, which is not based exclusively on either sink strength or soil nutrient availability, is proposed. We conclude that retranslocation response is species specific and related to the potential phenotypic growth response to changing environmental conditions and to short-term imbalance in the supply versus the demand for nutrients.     

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.     

Amino nitrogen and phenolic constituents of bark of American beech,Fagus grandifolia,and infestation by beech scale,Cryptococcus fagisuga

Concentrations of amino acids, total amino nitrogen, and phenols, ratio of phenol to amino nitrogen, and pH in the bark of American beech, Fagus grandifolia, were determined to see if these chemical constituents were correlated with susceptibility of trees to infestation by the beech scale, Cryptococcus fagisuga. The relationship of these bark constituents with scale infestation levels (none, light, and moderate), tree size (large vs. small trees), and bark shading was determined. Levels of some individual amino acids and total amino acid content in moderately infested trees were significantly higher than in uninfested trees. Large uninfested beech trees, which tend to be more readily infested by scale, had significantly higher concentrations of aspartic acid. Concentrations of phenol in the outer bark of both infested and uninfested trees were significantly higher than in the inner bark. However in bark infected by Nectria, phenol levels in the inner bark were significantly higher than in the outer bark.     

Stand and landscape level effects of a major outbreak of spruce beetles on forest vegetation in the Copper River Basin,Alaska

From 1989 to 2003, a widespread outbreak of spruce beetles (Dendroctonus rufipennis) in the Copper River Basin, Alaska, infested over 275,000 ha of forests in the region. During 1997 and 1998, we measured forest vegetation structure and composition on one hundred and thirty-six 20-m × 20-m plots to assess both the immediate stand and landscape level effects of the spruce beetle infestation. A photo-interpreted vegetation and infestation map was produced using color-infrared aerial photography at a scale of 1:40,000. We used linear regression to quantify the effects of the outbreak on forest structure and composition. White spruce (Picea glauca) canopy cover and basal area of medium-to-large trees [≥15 cm diameter-at-breast height (1.3 m, dbh)] were reduced linearly as the number of trees attacked by spruce beetles increased. Black spruce (Picea mariana) and small diameter white spruce (<15 cm dbh) were infrequently attacked and killed by spruce beetles. This selective attack of mature white spruce reduced structural complexity of stands to earlier stages of succession and caused mixed tree species stands to lose their white spruce and become more homogeneous in overstory composition. Using the resulting regressions, we developed a transition matrix to describe changes in vegetation types under varying levels of spruce beetle infestations, and applied the model to the vegetation map. Prior to the outbreak, our study area was composed primarily of stands of mixed white and black spruce (29% of area) and pure white spruce (25%). However, the selective attack on white spruce caused many of these stands to transition to black spruce dominated stands (73% increase in area) or shrublands (26% increase in area). The post-infestation landscape was thereby composed of more even distributions of shrubland and white, black, and mixed spruce communities (17–22% of study area). Changes in the cover and composition of understory vegetation were less evident in this study. However, stands with the highest mortality due to spruce beetles had the lowest densities of white spruce seedlings suggesting a longer forest regeneration time without an increase in seedling germination, growth, or survival.     

Growth, allocation and tissue chemistry of Picea abies seedlings affected by nutrient supply during the second growing season

One-year-old Norway spruce (Picea abies (L.) Karst.) seedlings were grown hydroponically in a growth chamber to investigate the effects of low and high nutrient availability (LN; 0.25 mM N and HN; 2.50 mM N) on growth, biomass allocation and chemical composition of needles, stem and roots during the second growing season. Climatic conditions in the growth chamber simulated the mean growing season from May to early October in Flakaliden, northern Sweden. In the latter half of the growing season, biomass allocation changed in response to nutrient availability: increased root growth and decreased shoot growth led to higher root/shoot ratios in LN seedlings than in HN seedlings. At high nutrient availability, total biomass, especially stem biomass, increased, as did total nonstructural carbohydrate and nitrogen contents per seedling. Responses of stem chemistry to nutrient addition differed from those of adult trees of the same provenance. In HN seedlings, concentrations of alpha-cellulose, hemicellulose and lignin decreased in the secondary xylem. Our results illustrate the significance of retranslocation of stored nutrients to support new growth early in the season when root growth and nutrient uptake are still low. We conclude that nutrient availability alters allocation patterns, thereby influencing the success of 2-year-old Norway spruce seedlings at forest planting sites.     

Mechanisms of Douglas-fir resistance to western spruce budworm defoliation: bud burst phenology, photosynthetic compensation and growth rate

We compared growth rates among mature interior Douglas-fir (Pseudotsuga menziesii var. glauca (Beissn.) Franco) trees showing resistance or susceptibility to defoliation caused by western spruce budworm (Choristoneura occidentalis Freeman), and among clones and half-sib seedling progeny of these trees in a greenhouse. We also investigated bud burst phenology and photosynthetic responses of clones to budworm defoliation in greenhouse experiments. Resistant mature trees had a higher radial growth rate than susceptible trees, especially during periods of budworm defoliation. Clones from resistant trees grew larger crowns than clones from susceptible trees, whereas stem base diameter at the ground line and height did not differ. Half-sib seedling progeny from resistant trees had larger stem diameter, height, and total biomass than progeny from susceptible trees. Mean 5-year radial growth increment of mature trees was more strongly correlated with growth of seedlings than with growth of clones. Clones from resistant trees had later bud burst than clones from susceptible trees, and budworm defoliation of clones depended on the degree of synchrony between bud burst phenology and budworm larval feeding. Clones of resistant and susceptible mature trees showed similar responses of net photosynthetic rate to 2 years of budworm defoliation. We conclude that phenotypic differences in crown condition of Douglas-fir trees following western spruce budworm defoliation are influenced by tree genotype and that high growth rate and late bud burst phenology promote tree resistance to budworm defoliation.     

Effects of climate on radial growth of Norway spruce and interactions with attacks by the bark beetle Dendroctonus micans (Kug., Coleoptera: Scolytidae): a dendroecological study in the French Massif Central

Samples of Norway spruce (Picea abies [L.] Karst) were dendrochronologically investigated in order to detect infestations by Dendroctonus micans (Kug.), the great spruce bark beetle (Col. Scolytidae), a relatively recent introduction to France. Uninfested natural forests located in the north-eastern French Alps and heavily infested plantations in the Ardèche region (Massif Central) were compared. The penetration holes bored in trunks by the bark beetle induced visible marks on wood, such as extreme ring width reductions, locally missing rings and crescent-shaped resin patches between consecutive rings that make possible a post-infestation dating.The outbreak began in 1979, 5 years prior to first insect visual detection by foresters. In the infested forest, tree basal area growth was not as sustained as in uninfested natural stands, but showed an inflection point at an unusually young tree age (from 30 to 40 years). Ring widths showing extreme synchronous radial growth reductions were caused either by excessively cold periods (e.g. in 1948, 1980, 1984, 1992) or by summer drought (as in 1986). Most of these weak growth years were shared with uninfested sites. In healthy forests, the consequences of extremely cold years were usually recorded only in high elevation stands, especially near the timberline, whereas summer drought effects were mostly visible in low altitude forests. By contrast, both phenomena were recorded in the infected Ardèche plantation. An analysis of tree-rings and monthly climate confirmed that Norway spruce growth in Ardèche plantations was reduced by excessively low minimum temperature during most parts of the year prior to ring formation, by higher than average maximum temperature during current spring and summer, and by drought in winter, spring and summer. Thus, the regional Ardèche climate with both cold winters and dry summers (especially in July) seems to weaken spruce trees planted there. Moreover, tree sensitivity to climate was found to be greatly enhanced by insect infestation. Such interactions between climatic stress and insect outbreak led to forest dieback in a 15–20-year period, when trees were still young (less than 70 years), and without any tree recovery. Therefore, in that region spruce plantations should be replaced by non-host species of Dendroctonus micans, especially where soil conditions may exacerbate drought effects.     

Microsite patterns of conifer seedling establishment and growth in a mixed stand in the southern Alps

In many forests of the Alps, permanent forest cover and, therefore, its continuous renewal is the main silvicultural goal. Regenerating these forests must be based on a sound understanding of the ecology of the tree species in question. The regeneration of silver fir (Abies alba Mill.) and Norway spruce (Picea abies (L.) Karst.) in the upper montane zone of the southern central Alps has so far received little attention. The aim of this study was, therefore, to characterize the ecological niche for the establishment and growth of silver fir and Norway spruce seedlings in this zone. The study was conducted as a case study on a plot of 1.5 ha in the upper montane zone of southern Switzerland, at an altitude of 1380 m. The basal area of the mixed Norway spruce–silver fir stand was 43.7 m² ha⁻¹. Seedlings were censused on 375 plots of 1.0 m² area. Seedling density was 1.30 ± 4.25 m⁻² (mean ± standard deviation) for Norway spruce and 0.87 ± 1.50 m⁻² for silver fir. Logistic regression models were used to test the effect of microsite characteristics on seedling occurrence, and general linear models for effects on seedling height growth and biomass increment. Most seedlings received less than 10% light (photosynthetic photon flux density) as compared to values in the open. Silver fir occurrence was positively related to microsites at the edge of canopy trees, but unrelated to ground cover type, light and micro-relief. Norway spruce occurrence was only, and positively, related to the presence of mosses. Height growth and biomass increment of seedlings of both species were only loosely correlated with microsite conditions. Seedlings without canopy cover grew faster than those under canopy cover, probably as a result of light and moisture limitations under the canopy of adult trees. Diffuse radiation was positively correlated with average annual biomass increment of silver fir, but not of Norway spruce seedlings. In general, the results suggest that silver fir seedlings have less specific microsite requirements than Norway spruce seedlings in terms of ground cover. They are also more shade tolerant, and therefore, grow faster than Norway spruce in low-light environments of the upper montane zone of the southern Alps. While the microsite concept can be helpful in designing silvicultural operations, it has limitations when only patch characteristics that are easy to assess are used, and others neglected. Finally, our study suggests that counting the number of green shoots is a promising method to quickly and non-destructively estimate the biomass of a great number of small seedlings.     

Variation in Elatobium abietinum Attack on Picea glauca and its Relation to Pissodes strobi Resistance

White spruce (Picea glauca (Moench) Voss) is host to several pests, including the white pine weevil (Pissodes strobi (Peck)) and the green spruce aphid (Elatobium abietinum (Walker)). The larvae of the white pine weevil damage spruce leaders by consuming the cortex while the green spruce aphid is a defoliator. White spruce emblings (seedlings produced by culturing tissues from seed embryos) from 18 families previously ranked for resistance to the white pine weevil were defoliated to varying degrees by the green spruce aphid in a natural outbreak that developed within a holding shadehouse. A strong relationship was shown between damage caused by the aphids and weevil resistance. Emblings ranked as highly weevil - resistant sustained significantly less aphid defoliation.     

Physiological responses of black spruce layers and planted seedlings to nutrient addition

We investigated effects of nutrient addition on several physiological characteristics of 60-cm-tall black spruce (Picea mariana Mill. B.S.P.) layers (i.e., rooted branches of overstory trees) and 20-cm-tall planted seedlings on a clear-cut, N-limited boreal site. After two growing seasons, current-year and one-year-old needles of fertilized trees (layers and seedlings combined) had higher net photosynthetic rates (A(n)) and maximum capacity of Rubisco for CO(2) fixation (V(max)) than unfertilized trees. One-year-old needles of fertilized trees had higher stomatal conductance (g(s)), higher water-use efficiency, and lower intercellular to ambient CO(2) ratio than unfertilized trees. Additionally, fertilized trees had higher predawn and midday shoot water potentials than unfertilized trees. Stomatal conductance of 1-year-old needles was 23% higher in seedlings than in layers, but there were no significant differences in g(s) of current-year needles between the regeneration types. For both needle age-classes, A(n) and V(max) of layers were 25 and 40% higher, respectively, than the corresponding values for seedlings. The higher values of A(n), V(max) and foliar N concentration of layers compared with seedlings after two growing seasons may be associated with the larger root systems of the layers compared with the transplanted seedlings.     

Reduction in turgid water volume in jack pine,white spruce and black spruce in response to drought and paclobutrazol

Significant reductions in needle water content were observed in white spruce (Picea glauca (Moench) Voss), black spruce (Picea mariana (Mill) B.S.P.), and jack pine (Pinus banksiana Lamb.) seedlings in response to a 10-day drought, although turgor was apparently maintained. When the seedlings were re-watered after the drought, jack pine needles regained their original saturated volume, whereas white spruce and black spruce needles did not. Significant drought-induced reductions in turgor-loss volume (i.e., tissue volume at the point of turgor loss) were observed in shoots of all three species, especially jack pine. Repeated exposure to 7 days of drought or treatment with the cytochrome P(450) inhibitor, paclobutrazol ((2RS,3RS)-1-(4-chlorophenyl)-4,4-dimethyl-2-(1,2,4-triazol-1-yl)-pentan-3-ol), reduced seedling height relative to that of untreated controls in all three species. The reductions in saturated and turgor-loss needle volumes in the paclobutrazol-treated seedlings were comparable with those of seedlings subjected to a 10-day drought. The treatment-induced reductions in shoot and needle water contents enabled seedlings to maintain turgor with tissue volumes close to, or below, the turgor-loss volume of untreated seedlings. Paclobutrazol-treated seedlings subsequently survived drought treatments that were lethal to untreated seedlings.     

Mistletoe effects on Scots pine decline following drought events: insights from within-tree spatial patterns, growth and carbohydrates

Forest decline has been attributed to the interaction of several stressors including biotic factors such as mistletoes and climate-induced drought stress. However, few data exist on how mistletoes are spatially arranged within trees and how this spatial pattern is related to changes in radial growth, responses to drought stress and carbon use. We used dendrochronology to quantify how mistletoe (Viscum album L.) infestation and drought stress affected long-term growth patterns in Pinus sylvestris L. at different heights. Basal area increment (BAI) trends and comparisons between trees of three different infestation degrees (without mistletoe, ID1; moderately infested trees, ID2; and severely infested trees, ID3) were performed using linear mixed-effects models. To identify the main climatic drivers of tree growth tree-ring widths were converted into indexed chronologies and related to climate data using correlation functions. We performed spatial analyses of the 3D distribution of mistletoe individuals and their ages within the crowns of three severely infested pines to describe their patterns. Lastly, we quantified carbohydrate and nitrogen concentrations in needles and sapwood of branches from severely infested trees and from trees without mistletoe. Mistletoe individuals formed strongly clustered groups of similar age within tree crowns and their age increased towards the crown apex. Mistletoe infestation negatively impacted growth but this effect was stronger near the tree apex than in the rest of sampled heights, causing an average loss of 64% in BAI (loss of BAI was ～51% at 1.3 m or near the tree base). We found that BAI of severely infested trees and moderately or non-infested trees diverged since 2001 and such divergence was magnified by drought. Infested trees had lower concentrations of soluble sugars in their needles than non-infested ones. We conclude that mistletoe infestation causes growth decline and increases the sensitivity of trees to drought stress.     

Boron retranslocation in Scots pine and Norway spruce

We previously traced 10B-enriched boric acid from shoots to roots to demonstrate the translocation of boron (B) in Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings. To gain a more detailed understanding of B translocation, we sought: (1) to demonstrate B retranslocation directly, by showing that foliar-applied 10B is located in the new growth after dormancy; and (2) to assess whether shoot-applied B affects growth in the long term. We applied 10B-enriched boric acid to needles of Scots pine and Norway spruce seedlings. After a dormancy period and 9 weeks of growth, small but significant increases in the 10B isotope were found in the new stem and needles of both species. In Scots pine, the total B concentration of the new stem was also increased. Both species contained polyols, particularly pinitol and inositol. Boron-polyol complexes may provide a mechanism for mobilizing B in these species. To determine the long-term effects of applied B, seedlings were grown for two growing seasons after the application of 10B to shoots. In Norway spruce, the proportion of 10B in the root systems and current needles of the harvest year was slightly higher than in the controls, and in Scots pine root systems, marginally so. The B treatment had no effect on growth of Norway spruce seedlings. In Scots pine seedlings, the B treatment caused a 33% increase in total dry mass and significantly increased the number of side branches.