Freezing temperatures in the root zone—effects on growth of containerized Pinus sylvestris and Picea abies seedlings

Roots of 1‐year‐old containerized seedlings of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) were experimentally frozen in December. The seedlings were then grown for 3 weeks in a growth chamber and evaluated with regard to root growth capacity (RGC) and shoot elongation. The subsequent RGC of Scots pine declined as root zone temperatures were lowered from ‐6°C to ‐11°C and from ‐11°C to ‐16°C. Almost no root growth was observed after exposure to ‐20°C. Shoot growth was also negatively affected by low root temperatures but less than root growth. Low root temperatures did not affect Norway spruce as much as Scots pine, although root and shoot growth of Norway spruce were reduced after exposure to the lowest test temperatures (‐16°C and ‐20°C). The length of exposure, ranging between 1 and 8 hours had no effect on subsequent growth.     

Fine root morphological adaptations in Scots pine, Norway spruce and silver birch along a latitudinal gradient in boreal forests

Variability in short root morphology of the three main tree species of Europe's boreal forest (Norway spruce (Picea abies L. Karst.), Scots pine (Pinus sylvestris L.) and silver birch (Betula pendula Roth)) was investigated in four stands along a latitudinal gradient from northern Finland to southern Estonia. Silver birch and Scots pine were present in three stands and Norway spruce was present in all stands. For three fertile Norway spruce stands, fine root biomass and number of root tips per stand area or unit basal area were assessed from north to south. Principal component analysis indicated that short root morphology was significantly affected by tree species and site, which together explained 34.7% of the total variability. The range of variation in mean specific root area (SRA) was 51-74, 60-70 and 84-124 m(2) kg(-1) for Norway spruce, Scots pine and silver birch, respectively, and the corresponding ranges for specific root length were 37-47, 40-48 and 87-97 m g(-1). The range of variation in root tissue density of Norway spruce, Scots pine and silver birch was 113-182, 127-158 and 81-156 kg m(-3), respectively. Sensitivity of short root morphology to site conditions decreased in the order: Norway spruce > silver birch > Scots pine. Short root SRA increased with site fertility in all species. In Norway spruce, fine root biomass and number of root tips per m(2) decreased from north to south. The differences in morphological parameters among sites were significant but smaller than the site differences in fine root biomass and number of root tips.     

Height growth of planted conifer seedlings in relation to solar radiation and position in Scots pine shelterwood

Seedlings of different provenances of Scots pine (Pinus sylvestris L.), lodgepole pine (Pinus contorta Dougl., var. latifolia Engelm.) and Norway spruce (Picea abies (L.) Karst.) were planted in three Scots pine shelterwoods (125, 65 and 43 stems ha⁻¹) and a clear-cut, all in northern Sweden. The sites were mounded and planting took place during 2 consecutive years (1988 and 1989). The solar radiation experienced by the individual seedlings was determined using a simulation model. Height development of the seedlings was examined during their first 6 years after planting. During the final 3 years of the study, height growth of Norway spruce was relatively poor, both in the shelterwoods and the clear-cut area. Height growth of lodgepole pine was significantly greater than that of Scots pine, both in the shelterwoods and the clear-cut. In contrast to Norway spruce, Scots pine and lodgepole pine displayed significantly greater height growth in the clear-cut than in the shelterwoods. For all three species in the shelterwoods, regression analyses showed that height growth was more strongly correlated with the distance to the nearest tree than with the amount of radiation reaching the ground, i.e. growth was reduced in the vicinity of shelter trees. Therefore, we conclude that the significant reduction in height growth of seedlings of Scots pine and lodgepole pine in Scots pine shelterwoods was partially caused by factors associated with the distance to the nearest shelter tree. Because the substrate was a nitrogen-poor sandy soil, we suggest that root competition for mineral nutrients, especially nitrogen, accounts for the reduction in height growth.     

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.     

Interference of bracken (Pteridium aquilinum L. Kuhn) with Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies L. Karst.) seedling establishment

The nature of interference of bracken with Scots pine and Norway spruce seedling establishment was considered in three field experiments. In a seeding experiment, it was found that Scots pine germination was highest on exposed mineral soil and lowest when intact bracken litter and humus were present, suggesting adverse effects of litter and humus on pine regeneration probably due to phytotoxicity. In a second experiment, smothering by bracken caused high mortality of Scots pine seedlings while Norway spruce seedlings were relatively unaffected. Mortality for both Scots pine and Norway spruce seedlings was low when planted in a adjacent Scots pine-bilberry stand with no bracken. Annual shoot growth of Norway spruce was higher in bracken than in Scots pine-bilberry vegetation while no differences in shoot growth between these two vegetation types occurred for Scots pine. In a third experiment, activated carbon was added to the ground under Norway spruce seedlings planted in bracken to adsorb possible phytotoxic compounds released by bracken. The addition of carbon had no effect on seedling mortality or growth rate, indicating that the seedlings were not susceptible to allelochemicals released by bracken. Since large Norway spruce seedlings were relatively unaffected by bracken interference in this study, artificial regeneration with containerized Norway spruce seedlings is suggested to achieve an acceptable conifer tree establishment on clear-cuts invaded by bracken.     

Responses of Pinus sylvestris and Picea abies Seedlings to Limited Phosphorus Fertilization and Treatment with Elevated Ozone Concentrations

Scots pine ( Pinus sylvestris L.) and Norway spruce [ Picea abies (L.) Karst.] seedlings were exposed to high phosphorus (HP) or low phosphorus (LP) availability for one growing season in the open field, and to combined P availability and elevated ozone (O 3 ) concentrations (0, 55, 110 and 210 ppb for Scots pine and 0, 40, 75 and 150 ppb for Norway spruce, respectively) for 28 days in controlled laboratory chambers. Compared with HP, the LP treatment reduced Scots pine current-year (C) shoot and root dry masses and Norway spruce total dry mass, whereas the highest O 3 concentrations increased the magnesium concentration of Scots pine C needles and P concentrations of the C needles of both tree species. Chlorophyll a, a+b and carotenoid concentrations of Scots pine C needles were significantly higher in the LP treatment compared with HP under the highest O 3 concentration (210 ppb). In the mesophyll tissue of C needles of both tree species, LP treatment increased the size of mitochondria and elevated O 3 -induced granulation of chloroplast stroma and disintegration of cytoplasm. Exposure to elevated O 3 concentrations increased swelling of chloroplast thylakoids and reduced the amount of vacuolar tannin in the LP Scots pine C needles. The results suggest disturbances in needle photosynthetic machinery due to acute exposure to the combination of elevated O 3 and low P availability. However, clear additive effects were found only in needle P concentrations < 1 mg g -1 in short-term O 3 exposure.     

Long‐term effects of liming on the fine‐root standing crop of Picea abies and Pinus sylvestris in relation to chemical changes in the soil

The long‐term effects of lime application on fine roots of Norway spruce, Picea abies (L.) Karst, and Scots pine, Pinus sylvestris (L.), have been studied in five experimental forest stands subjected to different lime applications 5 to 18 years before the present study was undertaken. The effects of liming does not seem to significantly influence fine‐root development in forest stands in the long term. The only response to liming in measured root variables was a tendency to increased specific root length (SRL = fine‐root length/fine‐root dry weight, m/g). A correlation between increased SRL, decreased root biomass and increased stem volume growth was indicated. Changes in water extractable amounts of mineral elements—P, K, Ca, Mg, Mn, S, Al and Fe‐in bulk soil and rhizosphere soil from the mineral soil layers were studied in a control area and an area treated with 3830 kg CaCO₃ ha^‐1. Few significant differences were found between treatments, and then mainly in the case of Ca.     

Forest Fertilization Research in Finland: A Literature Review

This paper summarizes results from fertilization research carried out in Scots pine ( Pinus sylvestris L.) and Norway spruce [ Picea abies (L.) Karst.] stands growing on mineral soil in Finland. The use of needle and soil analyses to indicate nutrient status and the need for fertilization are discussed along with the possibilities of increasing tree growth in various site types and stages of stand development. Doses, types of fertilizer, application time, growth increase, wood quality and the profitability of fertilization are also discussed. The stand response in terms of soil acidity, biological activity, soil fauna and resistance of trees to nutrient addition are all reviewed. The effects of fertilization on understorey vegetation, mushrooms and berries, and leaching loss of nutrients are also included.     

Fine root biomass in relation to site and stand characteristics in Norway spruce and Scots pine stands

Variations in fine root biomass of trees and understory in 16 stands throughout Finland were examined and relationships to site and stand characteristics determined. Norway spruce fine root biomass varied between 184 and 370 g m(-2), and that of Scots pine ranged between 149 and 386 g m(-2). In northern Finland, understory roots and rhizomes (< 2 mm diameter) accounted for up to 50% of the stand total fine root biomass. Therefore, the fine root biomass of trees plus understory was larger in northern Finland in stands of both tree species, resulting in a negative relationship between fine root biomass and the temperature sum and a positive relationship between fine root biomass and the carbon:nitrogen ratio of the soil organic layer. The foliage:fine root ratio varied between 2.1 and 6.4 for Norway spruce and between 0.8 and 2.2 for Scots pine. The ratio decreased for both Norway spruce and Scots pine from south to north, as well as from fertile to more infertile site types. The foliage:fine root ratio of Norway spruce was related to basal area and stem surface area. The strong positive correlations of these three parameters with fine root nitrogen concentration implies that more fine roots are needed to maintain a certain amount of foliage when nutrient availability is low. No significant relationships were found between stand parameters and fine root biomass at the stand level, but the relationships considerably improved when both fine root biomass and stand parameters were calculated for the mean tree in the stand. When the northern and southern sites were analyzed separately, fine root biomass per tree of both species was significantly correlated with basal area and stem surface area per tree. Basal area, stem surface area and stand density can be estimated accurately and easily. Thus, our results may have value in predicting fine root biomass at the tree and stand level in boreal Norway spruce and Scots pine forests.     

Compaction of till soils and growth tests with Norway spruce and scots pine

Five of the most common soils in forest areas in Sweden, viz., coarse sand, fine sand, coarse sandy till, medium sandy till and fine sandy till soil, were compressed at different loadings for 10 min. In the soils studied, it was found that the more fine material there was in the soil, the easier it compacted, and that till soils compacted much more than sediments of the same main particle class. The chosen compression time seemed to correspond well with an equilibrium level of static compaction. Influence of wall friction in compression tubes was found to be negligible for till soils.After compaction the soil cores were sown with Norway spruce (Picea abies (L.) Karst.) or Scots pine (Pinus sylvestris L.) and 16–20 days later the root lengths were recorded. Lengths of primary roots in per cent of the control were found to be almost inverse to the compaction of the soils; Norway spruce was more impeded than Scots pine. Thus, the bulk density alone is a poor indicator of the impact of soil compaction. Growth to 26, 33 and 120 days showed that the extension of secondary roots might be more impeded by compaction than the primary roots. Fertilization increased growth but did not decrease the growth difference between seedlings on unloaded and loaded soil cores.     

Damage by pine weevil Hylobius abietis to conifer seedlings after shelterwood removal

Abstract

Pine weevil (Hylobius abietis L.) damage to seedlings after overstorey removal was investigated in a survey study in six shelterwoods in the south–central part of Sweden. The shelterwoods predominantly consisted of Scots pine, except at one site where the shelter trees mainly consisted of Norway spruce. Before final cutting, 10 plots were laid out at each site and measurements of shelter trees and marked seedlings were taken. The seedlings were examined during the 2 years after final cutting. The study showed that removal of shelter trees increases the risk of severe damage by pine weevil and the variable that was most strongly correlated with the risk was the seedling root collar diameter. Both Scots pine and Norway spruce seedlings were severely damaged by pine weevil, and most of the feeding occurred during the first year after cutting. The amount of debarked area was significantly larger for Scots pine than for Norway spruce seedlings. Vitality (growth of the leading shoot before final cutting) of the seedlings also affected the probability of damage. Seedlings with high vitality were less damaged by pine weevil than seedlings with low vitality. For Scots pine the shelterwood density before final cutting was correlated to the intensity of pine weevil feeding after cutting. In conclusion, after the final cutting of a pine or spruce shelterwood, pine weevils will probably invade the area. To avoid serious damage, Norway spruce and Scots pine seedlings should have reached a diameter of at least 10–12 mm.     

Root growth capacity and field performance of Pinus sylvestris and Picea abies seedlings

Correlations between root growth capacity (RGC), at the time of planting, and field performance were studied for Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings. Before planting a gradient in seedling viability was generated through exposure to low root temperatures and different winter storage regimes. The hypothesis that high RGC values would improve field performance was to some extent verified for pine seedlings while no correlations could be registered for spruce. Reasons for these results are discussed.     

Evidence that longer needle retention of spruce and pine populations at high elevations and high latitudes is largely a phenotypic response

There is abundant evidence that evergreen conifers living at high elevations or at high latitudes have longer-lived needles than trees of the same species living elsewhere. This pattern is likely caused by the influence of low temperature in combination with related factors such as a short growing season and low nutrient availability. Because it is not known to what degree such patterns result from phenotypic versus genotypic variation, we evaluated needle longevity for common-garden-grown lowland populations of European Scots pine (Pinus sylvestris L.) of wide latitudinal origin and Norway spruce (Picea abies L.) of wide elevational origin. Nine-year-old trees of 16 Scots pine populations ranging in origin from 47 degrees to 60 degrees N were studied in Kórnik, Poland (52 degrees N) and 18-year-old trees of 18 Norway spruce populations ranging in origin from 670 to 1235 m elevation in southwestern Poland were studied near Morawina, Poland (51 degrees N, 180 m elevation). There was no tendency in either species for populations from northern or high elevation origins to retain needles longer than other populations. All of the Scots pine populations had between 2.5 to 3.0 needle age cohorts and all of the Norway spruce populations had between 6.4 and 7.2 needle age cohorts. Thus, extended needle retention in Scots pine and Norway spruce populations in low-temperature habitats at high elevations and high latitudes appears to be largely an environmentally regulated phenotypic acclimation.     

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.     

Growth response of coniferous stands to whole‐tree harvesting in early thinnings

A series of 15 field experiments was established to quantify the growth response of first‐thinning stands of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst) to whole‐tree harvesting and to estimate the need for nutrient compensation. The experiments were undertaken in Finland, Norway and Sweden and represent a wide range of site conditions. The site index (H ₁₀₀) of Scots pine stands varied from 19 to 29 m, and that of Norway spruce stands from 28 to 36 m. Total amounts of biomass and nutrients removed were calculated based on data obtained from felled sample trees. During the first 5‐yr period the growth response to the removal of logging residues varied considerably in both pine and spruce stands. Regression analyses did not reveal any functions that explained the variation in results satisfactorily. In cases where whole‐tree harvesting influenced tree growth negatively, this effect was counteracted by compensatory fertilization. It was concluded that to determine the response of remaining trees to harvesting intensity reliably, the post‐harvest period analysed must be longer than 5 yrs.     

Contrasting effects of elevated carbon dioxide concentration and temperature on Rubisco activity,chlorophyll fluorescence,needle ultrastructure and secondary metabolites in conifer seedlings

Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings were grown for 50 days in growth chambers in an ambient or twice ambient carbon dioxide concentration ([CO2]) at a day/night temperature of 19/12 degrees C or 23/16 degrees C. Although elevated [CO2] (EC) had only slight effects on the growth parameters measured, elevated temperature (ET) increased above ground dry mass of both species. Among treatments, biomass accumulation of both species was greatest in the combined EC + ET treatment. The EC treatment induced thylakoid swelling and increased numbers of plastoglobuli observed in Scots pine needles. Although EC had little effect on Rubisco protein or N concentration of needles, ET had a large effect on N-containing compounds and enhanced N allocation from 1-year-old needles. Terpenoids were more responsive to EC and ET than total phenolics. Generally, terpene concentrations were reduced by EC and increased by ET. Increased terpenoid concentrations in response to ET might be associated with thermotolerance of photosynthesis. In Norway spruce, EC decreased total phenolic concentrations in needles, probably as a result of increased growth. We conclude that, in seedlings of these boreal species, the effects of elevated [CO2] on the studied parameters were small compared with the effects of elevated temperature.     

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

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

Effect of uninucleate Rhizoctonia on Scots pine and Norway spruce seedlings

One‐year‐old container‐grown seedlings were planted in spring on clear cut areas: the Norway spruce (Picea abies) on a moist upland site (Myrtillus‐type) and Scots pine (Pinus sylvestris) on a dryish upland site (Vaccinium‐type). While still in the nursery, half of the seedlings of each species had been inoculated during the previous summer, with a uninucleate Rhizoctonia sp., a root dieback fungus. At outplanting all the seedlings appeared healthy and had a normal apical bud, although the height of the inoculated seedlings was less than that of the uninoculated control seedlings. At the end of the first growing season after planting, the mortality of inoculated Scots pine and Norway spruce seedlings was 25 and 69%, respectively. After two growing seasons the mortality of inoculated seedlings had increased to 38% for Scots pine and 93% for Norway spruce. The mortality of control seedlings after two growing seasons in the forest was 2% for Scots pine and 13% for Norway spruce. After outplanting the annual growth of inoculated seedlings was poor compared with the growth of control seedlings. These results show that, although Rhizoctonia‐affected seedlings are alive and green in the nursery, the disease subsequently affects both their survival and growth in the forest.     

Heterobasidion annosum root rot in Picea abies: Modelling economic outcomes of stump treatment in Scandinavian coniferous forests

Abstract

The economic outcomes of stump treatment against spore infections of the root rot pathogen Heterobasiodion annosum s. l. were analysed based on simulations in four stands typical of Swedish forestry and forest management: (A) Norway spruce [Picea abies (L.) Karst.] stand on former agricultural soil (SI?=?32), (B) Norway spruce stand (SI?=?26) on forest land; (C) Mixed stand of Norway spruce and Scots pine (Pinus sylvestris L.) (SI?=?24) with only H. parviporum present, i.e. no infection of Scots pine and no possibility of interspecies spread of disease between hosts; and (D) same as C, but H. annosum s. str. (Fr.) Bref and H. parviporum Niemelä & Korhonen present, i.e. interspecies spread of disease possible. Models for disease development, growth and yield and cross-cutting were used in the simulations. The simulated decay frequency in Norway spruce trees ranged between 2 and 90%. Stump treatment at the previous final felling and in all thinning operations was profitable at interest rates 1 and 3% in stands A, B and D, but not in stand C. In stand C, no stump treatment at all or treatment in thinnings only gave the highest net future value. Implications for stump treatment in practical forestry are discussed.     

Fast, nondestructive measurement of frost hardiness in conifer seedlings by VIS+NIR spectroscopy

Frost hardiness development from mid-August to mid-November was evaluated in seedlings of three provenances of Norway spruce (Picea abies (L.) Karst.) and three provenances of Scots pine (Pinus sylvestris L.) raised at nurseries in north, central and south Sweden. Measurements of the visible + near infrared (VIS+NIR) spectra of shoots were made simultaneously with estimates of frost hardiness based on electrolyte leakage following artificial freezing. Nine physiological variables known to influence frost hardiness were measured throughout the experiment. Multivariate analysis showed that VIS+NIR spectra explained 69% and 72% of the variation in frost hardiness in Scots pine and Norway spruce, respectively. Stem lignification, dry weight fraction, and starch, glucose, fructose, galactose, sucrose, raffinose and stachyose concentrations together explained 80% and 85% of the variation in frost hardiness in Scots pine and Norway spruce, respectively when used as independent X variables in a partial least squares model. These physiological variables could be related to varying degrees with variation in the VIS+NIR spectra. We conclude that VIS+NIR spectroscopy provides a rapid nondestructive technique for measuring frost hardiness in conifer seedlings based on causal relationships between the spectra and the physiology of seedling frost hardiness.