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.     

Snow disturbances in secondary Norway spruce forests in Central Europe: Regression modeling and its implications for forest management

Snow is an important ecological factor limiting vegetation distribution, growth, and regeneration, and the importance of snow in the latitudes of Central Europe is expected to increase in the future. We assessed snow damage to secondary spruce stands (not of native provenance therefore not adapted to local conditions) in a mountainous region of 14500 ha in Central Europe (Moravian-Silesian Beskids, Czech Republic). We used neural networks-based regression modeling to study the relationship between stand and environmental parameters and four types of snow damage (top tree, crown, and stem breakage, and uprooting) that occurred during heavy snowfalls in winter 2005/2006 and 2009. Almost 40000 trees were sampled in 345 plots after each of these two events.The results suggest that parameters that can be controlled by forest management (mainly stand density and taper) were not closely associated with spruce forest resistance to snow damage. Investigated snow damage types were primarily related to the developmental stage of the stand, as indicated by stand volume, age, height, and diameter. Damage in 2009, which was caused by shorter-lasting and lower snow loads than the damage in 2005/2006, was also associated with elevation and snow depth. The response of snow damage to all stand development-related variables was clearly unimodal.We infer that forest management can reduce snow damage to secondary spruce forests in Central Europe only to a limited extent, especially under heavy snow loads. This conclusion is supported by the heavy snows that have frequently fallen on forests in Central Europe in the past and the projected increase in winter precipitation in mid- and northern latitudes; thereby increasing snow damage to forest in the future. Therefore, managers of such spruce forests should not specifically consider forest resistance to snow damage but should apply general practices that maintain forest health and productivity.     

Effects of forest restoration treatments on the abundance of bark beetles in Norway spruce forests of southern Finland

Restoration of protected areas in boreal forests frequently includes creating substantial volumes of dead wood. While this benefits a wide range of dead wood dependent invertebrate species, some of these are regarded as forest pests. Therefore, the risk of elevated levels of tree mortality in surrounding commercial forests must be considered. In a large-scale field experiment in southern Finland, we studied the effects of restoration treatments on the abundance of bark beetles within and in the vicinity of restored areas, in particular focusing on Ips typographus and Pityogenes chalcographus. The treatments applied to managed Norway spruce forests were controlled burning and partial harvesting combined with retaining 5, 30 or 60 m³/ha of cut down wood. We found that the abundance of bark beetles increased by both burning and harvesting with down wood retention, being highest where burning and harvesting had been combined. The actual volume of down wood retention had no significant effect. The effect of burning on the number of bark beetles along host tree boles was negative which suggests that burnt spruces provided a less suitable resource for bark beetles than unburnt dead spruces. The abundance of bark beetles along host trees also decreased with increasing volume of down wood retention. The abundance of P. chalographus was slightly elevated up to 50 m outside restored areas but the abundance was very low compared to that within the areas. The abundance of I. typographus was extremely low outside restored areas. We conclude that restoration treatments increase the abundance of bark beetles via increased availability of resources, but that the effect of burning is likely to be counteracted by decreased resource quality. Thus, burning might be the “safest” way to produce large quantities of dead wood. Furthermore, the fact that only few beetles were collected in adjacent areas suggests that restored areas pose little threat of serving as refugia in which bark beetle populations increase in sufficient numbers to attack live trees in adjacent forests. However, restoration actions repeated at consecutive years within a small area might enable the populations to grow to outbreak levels.     

Effects of elevated CO(2) concentration and nutrition on net photosynthesis,stomatal conductance and needle respiration of field-grown Norway spruce trees

To study the effects of elevated CO(2) on gas exchange, nonstructural carbohydrate and nutrient concentrations in current-year foliage of 30-year-old Norway spruce (Picea abies (L.) Karst.) trees, branches were enclosed in ventilated, transparent plastic bags and flushed with ambient air (mean 370 &mgr;mol CO(2) mol(-1); control) or ambient air + 340 &mgr;mol CO(2) mol(-1) (elevated CO(2)) during two growing seasons. One branch bag was installed on each of 24 selected trees from control and fertilized plots. To reduce the effect of variation among trees, results from each treated branch were compared with those from a control branch on the same whorl of the same tree. Elevated CO(2) increased rates of light-saturated photosynthesis on average by 55% when measured at the treatment CO(2) concentration. The increase was larger in shoots with high needle nitrogen concentrations than in shoots with low needle nitrogen concentrations. However, shoots grown in elevated CO(2) showed a decrease in photosynthetic capacity compared with shoots grown in ambient CO(2). When measured at the internal CO(2) concentration of 200 &mgr;mol CO(2) mol(-1), photosynthetic rates of branches in the elevated CO(2) treatments were reduced by 8 to 32%. The elevated CO(2) treatment caused a 9 to 20% reduction in carboxylation efficiency and an 18% increase in respiration rates. In response to elevated CO(2), starch, fructose and glucose concentrations in the needles increased on average 33%, whereas concentrations of potassium, nitrogen, phosphorus, magnesium and boron decreased. Needle nitrogen concentrations explained 50-60% of the variation in photosynthesis and CO(2) acclimation was greater at low nitrogen concentrations than at high nitrogen concentrations. We conclude that the enhanced photosynthetic rates found in shoots exposed to elevated CO(2) increased carbohydrate concentrations, which may have a negative feedback on the photosynthetic apparatus and stimulate cyanide-resistant respiration. We also infer that the decrease in nutrient concentrations of needles exposed to elevated CO(2) was the result of retranslocation of nutrients to other parts of the branch or tree.     

The chemical composition of needle and leaf litter from Scots pine, Norway spruce and white birch in Scandinavian forests

Needle litter from 14 stands of Scots pine (Pinus silvestris,L.), 13 stands of Norway spruce (Picea abies (L.) Karst.) andleaf litter from three stands of white birch (Betula pubescensEhrh.) were analysed for chemical composition. The concentrationsof the elements N, P, K, Ca, Mg and Mn as well as solid organiccomponents (lignin, cellulose and hemicelluloses) and solubleswere determined. When the average chemical compositions werecompared the Scots pine needle litter was clearly the most nutrient-poorlitter type. Of the solid organic-chemical components the ligninfraction dominated in the spruce and birch litter whereas thecellulose dominated in the pine needle litter. When Norway spruce and Scots pine were growing in adjacent standson soils with the same bedrock origin the spruce litter hadsignificantly higher concentrations of nutrients (N, P, K, Ca,Mg, Mn) than the pine needle litter. At sites where Norway spruceand white birch were growing in adjacent stands, the birch leaflitter had generally higher concentrations of nutrients. However, significant or nearly significant differences were onlyobtained for Mg (P = 0.002), K (P = 0.056) and N (P = 0.087),probably due to the few replicates of stands compared. Concerningorganic chemical components, the spruce needle litter had significantlyhigher concentrations of lignin and mannan than all the otherlitters and lower levels of ethanol-soluble substances, celluloseand galactan than the pine needle litter. Further, it had lowerconcentrations of water solubles, rhamnan and xylan than thebirch litter. No relationships were established between the nutrient statusof the conifer litters and the site index H100 (the dominantheight of the trees at a reference age of 100 years) of thestands. Concentrations of solid carbohydrates in the litterswere, however, positively correlated with site index (P <0.001). Further, the concentration of nitrogen in the pine needlelitter was negatively correlated with the latitude of the sites(P < 0.01). The influence of litter chemistry on the decompositionof litter and nutrient cycling of forests is discussed.     

Effects of photoperiod and temperature on the timing of bud burst in Norway spruce (Picea abies)

We examined the effects of several photoperiod and temperature regimes imposed during the winter-spring period on the timing of bud burst in rooted cuttings of Norway spruce (Picea abies (L.) Karst.) grown in a greenhouse in Finland. The treatments were initiated in November and December after the cuttings had been exposed to natural chilling and freezing events. Irrespective of the treatments applied, time to bud burst decreased with increased duration of previous exposure to natural chilling and freezing events. Fluctuating day/night temperatures and continuous lengthening of the photoperiod hastened bud burst. Shortening the photoperiod delayed bud burst, suggesting that little or no ontogenetic development toward bud burst takes place during mild periods before the winter solstice. In the case of climatic warming, this phenomenon may prevent the premature onset of growth that has been predicted by computer simulations with models that only consider temperature regulation of bud burst.     

Polyamines in embryogenic cultures of Norway spruce (Picea abies) and red spruce (Picea rubens)

Embryogenic cultures of red spruce (Picea rubens Sarg.) and Norway spruce (Picea abies (L.) Karst.) were initiated from dissected mature zygotic embryos. The tissues were grown on either proliferation medium or maturation medium. On proliferation medium, the embryogenic tissue continued to produce early stage somatic embryos (organized meristems attached to elongated, suspensor-like cells), whereas on maturation medium fully mature embryos developed from the embryonic tissue. Analysis of polyamines in tissues grown on these two media showed that: (1) both putrescine and spermidine concentrations were always higher in cultures grown on proliferation medium than in cultures grown on maturation medium; (2) in both species, spermidine concentrations declined with time in the tissues grown on maturation medium; and (3) spermine was present in only minute quantities and showed only a small change with time. The presence of difluoromethylornithine in the culture medium had little effect on polyamine concentration, whereas the presence of difluoromethylarginine caused a decrease in putrescine concentrations in both red spruce and Norway spruce tissues grown on proliferation medium or maturation medium.     

Conversion of Norway spruce forests in the face of climate change: a case study in Central Europe

Steadily increasing damage to Norway spruce forests in Europe has caused researchers and managers to consider whether these forests can be converted to more stable ecosystems. In a central European mountain region, we investigated whether management systems (MSs) specified by regional stakeholders provide sound alternatives to the currently applied management. We used the forest model Sibyla to explore whether the tested MSs differ in their sensitivity to climate change in terms of altered biomass production, stand structure, forest damage, and financial outcome. The tested MSs were no-management (NM), currently applied management (BAU), and management based on the preferences of forest managers (FM) or on the preferences of other stakeholders (OSH). With NM, spruce remained dominant during the simulation period 2010–2100, and the rate of damage significantly increased. Spruce also remained dominant with FM, while the abundance of non-spruce species significantly increased with BAU and OSH. The rate of salvage logging converged at 50% of the total harvest for all MSs up to 2050. Climate change reduced biomass production (?15%) with all MSs but had a negligible effect on biodiversity indicators. The average initial value of the simulated stands was 20,000 € ha^?1 and the nominal value in 2100 was between 1900 and 10,900 € ha^?1. The Net Present Value calculated with the 2% interest rate was negative during the whole simulation period (?5600 to ?18,500 € ha^?1 in 2100). Effect of climate change on all financial indicators was negative. Our findings indicate that secondary spruce forests are highly vulnerable and that the systems proposed by both forest managers and other regional stakeholders failed to significantly reduce forest damage and stabilize forest production.     

Persisting bark beetle outbreak indicates the unsustainability of secondary Norway spruce forests: case study from Central Europe

? Context

Secondary Norway spruce forests in the Western Beskids are among the most damaged forests in Europe. Although spruce bark beetle (Ips typographus) has been recently causing large-scale damage to these forests, our understanding of I. typographus dynamics in this environment is inadequate for evaluating forest sustainability.

? Aim

This study aims to evaluate the patterns of damage caused by I. typographus to spruce forests with compromised ecological stability.

? Methods

Forest infestation by I. typographus was inferred from sanitary felling data collected from 1998 to 2004. Stand and site data were obtained from forest management plans. Spatial-dependence analysis, ordinary kriging and neural network-based regression modelling were used to investigate the patterns of infestation and the casual relationships in the studied ecosystem.

? Results

I. typographus long-distance dispersal substantially decreased with outbreak culmination. The spread of infestation was only weakly related to stand and site parameters. Infestations spread isotropically at the stand and patch level but directionally at the regional scale.

? Conclusions

The large-scale spread of infestation can be explained by the uniform age and species composition of the investigated forests and by the ability of populations to overwhelm suboptimal trees. The observations presented here suggest that secondary spruce forests in Europe may be unsustainable due to unprecedented bark beetle outbreaks, which can be further amplified by changing climate.     

Effects of experimental drought on the fine root system of mature Norway spruce

Norway spruce (Picea abies (L.) Karst.) is an economically important, but relatively drought-sensitive tree species that might suffer from increasing drought intensities and frequencies, which are predicted to occur in parts of central Europe under future climatic change. In a throughfall exclusion experiment using sub-canopy roofs, we tested the hypothesis that enhanced drought leads to an increase in fine root mortality, and also to a higher, subsequent fine root growth. Fine root production and mortality were assessed using two independent approaches, sequential soil coring (organic layer) and direct observations in minirhizotrons (organic layer plus upper mineral soil). Six weeks of throughfall exclusion resulted in mild drought stress, i.e. a reduction in average soil moisture from 20 to 12 vol.% during the treatment. Based on the sequential coring data, experimental drought did not result in significant changes in fine root biomass during the 6-week treatment period, but caused an increase in fine root mortality by 61% in the 6 weeks following the drought treatment. Remarkably, fine root production showed a synchronous increase in this period, which more than compensated for the loss due to increased mortality. The minirhizotron data confirmed that the drought treatment increased fine root loss in the organic layer. Based on this method, however, root loss occurred during the drought period and was not compensated by increased root production. The mild drought stress was mainly restricted to the organic layer and did not significantly influence fine root dynamics in the mineral soil. We calculated that the drought event resulted in an extra input of about 28 g C m⁻² and 1.1 g N m⁻² to the soil due to increased fine root mortality. We conclude that even periods of mild drought significantly increase fine root mortality and the associated input of root-derived C to the soil organic matter pool in temperate Norway spruce forests.     

Effects of cone-induction treatments on black spruce (Picea mariana) current-year needle development and gas exchange properties

Both drought and root pruning (RP) increased the number of cones induced when black spruce (Picea mariana (Mill.) B.S.P.) grafts were injected with gibberellins A(4/7) (GA), but their effects on predawn shoot water potential and current-year needle development differed. Drought decreased predawn shoot water potential (Psi(pd)), but only during the period when irrigation was withheld, and it had no effect on the growth or gas exchange properties of current-year needles. Conversely, root pruning had little effect on Psi(pd), but it resulted in trees with smaller current-year needles that had lower nitrogen and chlorophyll concentrations and reduced rates of gas exchange up to the later stages of shoot elongation compared with needles of control trees. These findings are discussed in relation to potential effects on the development of induced cones in the following growth cycle.     

The effects of fertilization on the health status,nutrition and growth of Norway spruce forests with yellowing symptoms

We studied the effects of magnesium (Mg) NPK fertilizer applied in 2000 to maturing Norway spruce (Picea abies /L./ Karst.) stands with yellowing symptoms and situated on poor acidified soils in the ?umava mountains (the Czech Republic). Long-term defoliation, nutrient content in the foliage and diameter increment were evaluated in three specific variants: yellowing trees, trees without yellowing symptoms and fertilized trees. The results indicate differences in soil characteristics, nutrient content and radial growth with response to climate. Fertilization increased Mg content and base saturation in the soil and decreased the amount of exchangeable aluminum and hydrolytic acidity. Differences were significant especially in magnesium, calcium and phosphorus contents in the foliage of yellowing trees. There was also a significant difference in manganese content; yellowing trees showed deficiency in comparison with both fertilized and unfertilized healthy trees. Two years after the initial fertilization (to present), defoliation and yellowing symptoms in the unfertilized yellowing trees were more pronounced compared to other variants. These symptoms were more influenced by fertilization than foliation. The duration of significant positive effects of fertilization on diameter increment lasted for a shorter time than the effect on foliage. Simultaneously, higher sensitivity to climatic factors was observed in the fertilized trees.     

Infection of roots of Norway spruce (Picea abies) by Heterobasidion annosum

Wounding of excised roots of Norway spruce [Picea abies (L.) Karst.] caused a rapid accumulation of phenolic compounds, decrease of starch content and of certain enzyme activities in the ray cells. Infection of the wounds by Heterobasidion annosum (Fr.) Bref. intensified these processes, caused oxidation of the phenols, and induced activity of acid phosphatase and β-glucosidase in the ray cells of the infected area. The reactions were oxygen-dependent. Intra- and extracellular enzymes were detected in the wood-infecting hyphae. Other wood-growing fungi caused similar reactions in the wood as H. annosum.     

Effects of stand density on ecosystem properties of subalpine forests in the southern Rocky Mountains, USA

? Mixed coniferous, subalpine forest communities in the Rocky Mountains are historically dense and have experienced infrequent, high-severity fire. However, many of these high-elevation stands are thinned for a number of perceived benefits.

? We explored the effects of forest stand density on ecosystem properties in subalpine forests in Colorado, USA, 17–18 y after forests were managed for timber.

? Forest structure significantly altered the composition and chemical signature of plant communities. Previously managed stands contained lower density of overstory trees and higher ground cover compared to paired reference stands. Foliar phenolic concentration of several species was negatively related to basal area of overstory trees. Furthermore, reductions in stand density increased total foliar phenolic:nitrogen ratios in some species, suggesting that gap formation may drive long-term changes in litter quality. Despite significant changes in forest structure, reductions in stand density did not leave a strong legacy in surface soil properties, likely due to the integrity of soil organic matter reserves.

? Changes in forest structure associated with past management has left a long-term impact on plant communities but has only subtly altered soil nutrient cycling, possibly due to trade offs between litter decomposability and microclimate associated with reductions in canopy cover.

     

Effects of manganese deficiency on soluble apoplastic peroxidase activities and lignin content in needles of Norway spruce (Picea abies)

Apoplastic peroxidase activities were investigated in manganese-deficient and manganese-sufficient needles of field-grown Norway spruce trees (Picea abies L.). In Mn-sufficient needles, two sets of peroxidases, one with an alkaline pI >/= 9 and another with an acidic pI 相似文献   

Effects of seed pre-soaking on the emergence and early growth of containerized Norway spruce seedlings

To study the effects of seed soaking on seedling emergence and development in Norway spruce [Picea abies (L.) Karst.] two commercial seed lots were soaked for 15 h in aerated water. After soaking, floating and sunken seeds were separated and analyzed prior to being sown in peat-filled containers. Seedling development was followed for one growing season. All sunken seeds were viable according to radiography whereas the floating seeds contained a considerable number of larvae filled and empty seeds. Mean time of emergence of the sunken seeds was approximately 1.5 days shorter than control (i.e. dry) seeds and floating seeds. Soaking had no effect on the time taken for most germinants to emerge in both seed lots, thus failing to synchronize seedling emergence. The later a seedling would emerge, the more likely it was for it to die during the growing season or to be culled at the end of the growing season. Soaking had an inconsistent effect on stem diameter at the end of the growing season and no effect on stem or root biomass. Seedling size uniformity was equal in all treatments. This study suggests that soaking and seed sorting does not guarantee a more uniform seedling crop, but this technique may be useful in upgrading poor quality seed lots of Norway spruce containing a high number of empty or larva infested seeds.     

Effects of wide spacing and thinning strategies on wood quality in Norway spruce (Picea abies) stands in southern Sweden

Abstract

The initial density of a stand influences both the prethinning growth and the quality of the harvested trees. This study investigated the effects of different square spacings (2.0, 2.5 and 3.0 m) with thinning from above and thinning from below on stand development and tree characteristics in a 33-year-old Norway spruce [Picea abies (L.) Karst.] plantation in southern Sweden. Before thinning the total standing volume was not significantly different between treatments. Diameter at breast height (dbh) and diameter of the thickest branch were positively correlated. Before thinning, dbh of the trees and their thickest branches were significantly higher in the stands with 3.0 m spacing. After thinning, the dbh and the diameter of the thickest branch did not significantly differ between the 3.0 m spacing with thinning from above and 2.0 and 2.5 m spacing with thinning from below. Spacing did not significantly influence the presence or size of spike knots. The results suggest that a combination of wide spacing with thinning from above may yield timber of similar quality to denser spacing with thinning from below.     

The influence of magnesium deficiency on carbohydrate concentrations in Norway spruce (Picea abies) needles

Both short- and long-term effects of Mg deficiency on carbohydrate metabolism were investigated in 6-year-old clonal Norway spruce (Picea abies (L.) Karst.) trees cultivated in sand culture with an optimal supply of nutrients, except for Mg which was supplied at 0.203, 0.041 and 0.005 mM to provide optimal, moderately deficient and severely deficient Mg supply, respectively. Annual changes in carbohydrate concentrations (starch, sucrose, glucose and fructose) were analyzed and diurnal changes were investigated on a single day during the summer. Older needles of trees in the moderate Mg-deficiency treatment developed tip-yellowing symptoms, whereas current-year needles remained green. The severe Mg-deficiency treatment led to pronounced yellowing symptoms in needles of all ages. Increased carbohydrate concentrations were observed before needle yellowing occurred. Diurnal and annual changes in carbohydrates were similar in all treatments; however, carbohydrate concentrations were influenced by Mg supply. In both Mg-deficiency treatments, starch concentrations increased in needles, especially during summer and autumn. Starch accumulation was more pronounced at the beginning of the Mg-deficiency treatments than at the end of the treatments. Sucrose, and to a minor extent, glucose and fructose concentrations tended to increase in response to Mg deficiency. The consequences of Mg deficiency on carbohydrate metabolism are discussed with respect to reduced plant growth and decreased transport rates of carbohydrates to sink organs.     

Indented rings (hazel growth) of Norway spruce reduce anisotropy of mechanical properties

This study presents three-point bending test results of Norway spruce clear wood specimens loaded on the radial-longitudinal plane in two different load cases. The tested samples were graded as resonance wood for instrument making and were characterised by narrow annual rings and relatively low density. The modulus of elasticity (MOE) and the corresponding modulus of rupture (MOR) are illustrated separately for the samples with straight grain and the group showing the specific growth pattern of indented rings (‘hazel growth’). With the longitudinal wood anatomical direction parallel to span width, the fibre deviation caused by the indents reduces MOE and MOR values, whereas a ‘reinforcing’ effect of the indents could be observed for the load case with span width parallel to the radial direction. Both aspects lead to a reduction in anisotropy for hazel-growth Norway spruce (anisotropy MOE: indented rings 11.6, straight grain 14.7, anisotropy MOR: indented rings 6.9, straight grain 8.9), which partly explains the exceptional position of this growth pattern for the construction of high-class musical instruments with outstanding mechanical and acoustical performance.