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.     

Root-associated fungi of healthy-looking Pinus sylvestris and Picea abies seedlings in Swedish forest nurseries

The aim of this study was to assess fungal communities in roots of healthy-looking Pinus sylvestris L. and Picea abies (L.) Karst. seedlings in nine forest nurseries in Sweden using a combination of traditional culturing and direct sequencing of internal transcribed spacer of fungal ribosomal RNA (ITS rRNA) from the roots. Culturing from 1800 surface-sterilised root segments resulted in 2387 fungal cultures representing 42 different taxa. Direct sequencing from 180 root segments resulted in 119 ITS rRNA sequences representing 25 different taxa. In total, 55 different fungal taxa were detected using both methods. Although direct sequencing was more efficient than culturing in detecting different fungal taxa, both methods provided complementary information about fungal communities in roots since each detected rather different groups of fungi. The most dominant taxa detected by culturing were Trichoderma viride Pers. (19.5%), Phoma mucivora Davey & Currah (19.1%), Phialocephala fortinii Wang & Wilcox (17.4%) and Meliniomyces variabilis Hambl. & Sigler (10.2%), while Thelephora terrestris Ehrh. (26.1%), Unidentified sp. NS126 (25.2%) and Heliotales sp. C20 (10.1%) were most commonly detected by direct sequencing. In conclusion, results showed that forest nurseries in Sweden harbour diverse communities of fungi associated with the roots of healthy-looking P. sylvestris and P. abies seedlings. Although fungal communities were often dominated by saprotrophs and endophytes, several facultative pathogens were also detected indicating that under suitable conditions they may be a potential threat to the plants.     

Susceptibility of different provenances of Pinus sylvestris,Pinus Contorta and Picea abies to Gremmeniella abietina

In a randomized block factorial experiment, 1200 seedlings from four provenances, each of Pinus sylvestris, Pinus contorta and Picea abies were inoculated with conidia (2 × 10⁴ and 1 × 10⁶ conidia/seedling) of Gremmeniella abietina (Brunchorstia pinea), isolated from P. contorta plantations in northern Sweden. A further 600 seedlings were left as controls. The occurrence of symptoms and the extension of dead tissues on the annual shoots were recorded 13 months after inoculation. Only the higher spore dose resulted in significant infection. P. sylvestris and P. contorta seedlings were equally susceptible (53% infected), and significantly more infected than P. abies seedlings (39%). The annual shoots of P. sylvestris and P. abies were affected to 43% and 37% of their length, which was significantly more than 15% of the length of P. Contorta shoots. The frequency of affected seedlings differed between the most southern and northern provenances of P. sylvestris and P. abies, and between the northwestern and the south-eastern provenance of P. contorta Twenty-six months after inoculation, a higher proportion of P. contorta seedlings than P. sylvestris and P. abies seedlings had recovered, and a lower proportion of P. contorta than P. sylvestris and P. abies seedlings had died. This paper discusses why P. sylvestris was found to be more susceptible to G. abietina than P. contorta in this experiment, while the reverse is found in plantations in northern Sweden.     

Incidence of Gremmeniella abietina in planted seedlings of Picea abies and Pinus sylvestris in northern Finland

The ascomycete Gremmeniella abietina causes a disease (scleroderris canker) on conifers. The pathogen kills terminal shoots and branches of Pinus sylvestris and Picea abies, sporulates on host shoots and causes leader changes (i.e. dieback). Damage caused by G. abietina was investigated in young P. abies and P. sylvestris plantations in northern Finland. Side branches from seedlings of both species and the main stems of P. sylvestris were collected from three sites in two locations (Poksa 1 and 2, Kivalo) in northern Finland. The number of cankers and leader changes was counted in branches of each age (i.e. year) in a total of ca. 6300 shoots of P. abies and 1200 shoots of P. sylvestris. Cankers were common on both P. abies and P. sylvestris in Poksa 1 (2002–2008) and on P. abies in Poksa 2 (1998–2001). In Kivalo, cankers occurred only sporadically on P. sylvestris. Leader changes were most frequent in 1999–2009 in Poksa 1, in 2001–2009 in Poksa 2 and in 2003 in Kivalo. Cankers and other symptoms of infection were more obscure on P. abies than on P. sylvestris. According to both conidial morphology and molecular analysis, the strain of G. abietina infecting both P. abies and P. sylvestris was small‐tree type (STT). This is the first report of G. abietina, STT or B type, injuring P. abies in plantations.     

Canopy density in stands of Picea abies and Pinus sylvestris after different thinning methods

Altogether 82 plots (261 estimations) of Picea abies (L.) Karst, and 193 plots (360 estimations) of Pinus sylvestris (L.) stands were estimated by a vertical tube. The “crown free projection”, CFP, of stands thinned in three methods with different thinning grades was measured: unthinned, heavily and very heavily thinned, heavily thinned delayed first thinning, extra heavily thinned and thinned from the top. Basal area (m²ha^?1) density (stems ha^?1) and diameter sum (m ha^?1) were plotted against CFP. Basal area was the best practical measure of stand in this study. Generally Scots pine stands have higher CFP and the curves are steeper than in Norway spruce stands. Depending on the grade of thinning, heavily and very heavily thinned spruce stands, delayed first thinning included, have CFP values of 10–15% and stands thinned from the top, 20–40%, compared with 30–80% and 30–60% respectively in pine stands. Extra heavily thinned stands have the highest CFP, 20–80% in spruce and 50–90% in pine stands. The CFP levels after thinning are too high in pine stands for avoidance of sucker and sprout production of aspen and birch. In dense Norway spruce stands thinned from the top or heavily and very heavily thinned, the CFP values are low enough (≤30%) to diminish the production of suckers.     

Effects of low-dose liming and nitrogen fertilization on stemwood growth and needle properties of Picea abies and Pinus sylvestris

Stemwood growth, needle weight and nutrient concentrations in needles after liming and nitrogen fertilization were studied in two randomized block trials. The treatments were a single application of 500 kg lime per hectare, 1000 kg limer per hectare, 150 kg N per hectare and combinations of the two compounds. The trials are situated in the southern part of Sweden on mesic mineral soils. One of the experiments was located in 32–55-year-old highly productive Picea abies stands and the other in a 31-year-old Pinus sylvestris stand.As calculated by growth-ring analysis, liming alone did not significantly affect the growth during a 10-year period following treatment. Nitrogen fertilization increased growth in pine, but not in spruce. Lime in combination with N gave about the same effect in pine as the pure N treatment, whereas lime and N interacted in spruce and resulted in increased stemwood growth.Current-year needles were sampled during the winter, weighed and analysed for nutrient concentrations. In the N-treated plots the N concentration increased in both tree species, whereas the needle weight increased only in pine. In two cases, lower concentrations were indicated. In the first, there was a tendency of decreased N concentrations in spruce needles after liming. In the second, the magnesium concentrations decreased in needles of both species, especially in the treatments that included nitrogen. The concentrations of the measured nutrients were never below levels considered to imply a severe deficiency.     

Effects of aluminium and calcium on Picea abies seedlings

Newly‐germinated Norway spruce seedlings were grown for 15 weeks in pots of iron podzol soil that received additions three times a week of nutrient solution containing six levels of Al (0, 0.37, 0.74, 1.5, 3.0 or 6.0 mM) and two levels of Ca (0.09 or 0.46 mM). The Ca concentrations in shoots and roots decreased with increasing Al levels. Growth was suppressed at 0.74 mM Al in the nutrient solution, which gave 0.5 mM Al in saturated soil extracts at the end of the experiment.     

Long-term Growth Effects Following Forest Nitrogen Fertilization in Pinus sylvestris and Picea abies Stands in Sweden

Forest nitrogen (N) fertilization induces changes in the soil and soil microorganisms that could hypothetically affect the long-term productivity of the soil. The tree growth response following a normal (i.e. 150 kg N ha^?1) single shoot N fertilization has a duration of 7–10 yrs. The aim of this study was to investigate whether any residual effects on tree growth persist, which could be attributed to previous N fertilization. The study included six Pinus sylvestris L. and three Picea abies (L.) Karst. experimental sites, sampled for growth parameters 14–28 yrs after the last fertilization. Residual growth effects were on average small, and not statistically significant. Negative residual growth effects of varying duration could be discerned, especially at low-fertility P. sylvestris sites. However, there was an overall tendency for growth to increase in the long term after N fertilization. The main conclusion is that operational forest N fertilization with a normal N dose should not be regarded as a threat to long-term forest production.     

Induction of hairy and normal roots on Picea abies,Pinus sylvestris and Pinus cortorta by agrobacterium rhizogenes

Two‐ to three‐week‐old seedlings of Picea abies, Pinus sylvestris and Pinus contorta were inoculated with the supervirulent strain Agrobacterium rhizogenes R1600 2cm above the base of the hypocotyl; other treatments included inoculation with Agrobacterium tumefaciens strain A281 (pTVK291) or mock inoculation. Hairy or highly branched roots formed on 2–3% of seedlings of all 3 species that were infected by R1600 but on none of the other seedlings. Normal adventitious roots were also induced on 5–10% of Agrobacterium‐treated seedlings but none appeared on the other seedlings. Opine assays and growth rates of hairy and normal roots indicated that only roots with the hairy morphology were transformed.     

Photosynthesis and root respiration of Pinus sylvestris and picea abies seedlings after different root freezing and storage temperatures

Photosynthetic performance and root respiration were measured for seedlings of Scots pine and Norway spruce under constant conditions in an open gas exchange system in the laboratory. Measurements were carried out after root exposure to ‐20, ‐5 and 0°C and subsequent longtime storage in darkness at +1 or +4°C. Stomatal conductance in relation to net photosynthetic rates was also investigated after the same treatment of seedlings. Root respiration was low for seedlings whose root system had been exposed to ‐20°C, Scots pine showing lower rates than Norway spruce. This was probably an indication of root damage. At least for one provenance of Scots pine, respiration rates were higher for seedlings stored at +1 than at +4°C. Photosynthetic performance was also lowest for seedlings whose roots had been exposed to +20°C compared to higher temperatures, the difference being more clear‐cut for Norway spruce than for Scots pine. Storage at +1 gave slightly higher photosynthetic rates than at +4°C. There was a close relation between stomatal conductance measured on individual needles and photosynthetic performance measured on the whole seedling.     

S and P intersterility groups in Heterobasidion annosum; Infection frequencies through bark of Picea abies and Pinus sylvestris seedlings and in vitro growth rates at different oxygen levels

Seedlings of Norway Spruce and Scots Pine were inoculated with S and P strains of the root rot fungus Heterobasidion annosum. Infection frequencies through the living bark layers were not related to subgroup affiliation in neither spruces nor pines. Growth rates in vitro were not related to growth rates in vivo. Oxygen concentration had a stronger effect on the growth rates of P than on S strains.     

Functions for Biomass Estimation of Young Pinus sylvestris,Picea abies and Betula spp. from Stands in Northern Sweden with High Stand Densities

New silvicultural regimes with high within-stand competition require new functions for estimation of standing stock and growth of biomass components, since the allometry of trees is changed by light competition. This paper presents functions for estimation of the aboveground biomass dry weights for stem wood, stem bark, branches and leaves of young (diameter at breast height <10 cm) Scots pine (Pinus sylvestris L.), Norway spruce [Picea abies (L.) Karst.] and birch (Betula pendula Roth. and Betula pubescens Ehrh.) trees growing in dense mixed stands. The functions were derived from a sample consisting of 84 Scots pine, 43 Norway spruce and 66 birch trees from six stands in northern Sweden with high stand densities (>10000 st ha^-1). The logarithmically transformed power function displayed a good ability to stabilize the variance of dry weights and showed a good fit to the material (0.37< R ² <0.99). A comparison with the most commonly used biomass functions in Sweden today showed that they overestimated the weight of stem wood and branches, while the weight of foliage was underestimated. The nature of these discrepancies suggested that the precision of biomass estimations might also be improved for young trees at wider spacing.     

Models for Predicting Wood Properties in Stems of Picea abies and Pinus sylvestris in Sweden

To study and model the variation of wood properties, sample trees were selected from 42 Norway spruce and 20 Scots pine stands covering a wide variation in climatic and site conditions, stand maturation and tree sizes. Plot and tree measurements were followed by sampling wood from different heights in each sample tree and laboratory measurements of wood properties. Mixed linear and non-linear prediction models were developed using diameters, number of annual rings and climatic indices as explanatory variables. The variation in spruce properties explained by these variables was: basic density 50%, latewood content 52%, juvenile wood diameter 85%, heartwood diameter 94% and bark thickness 76%. The corresponding values for pine were 59, 54, 79, 92 and 85%. Random among-tree variance was an important contributor to the remaining variation for density and latewood. In general, only a minor part of the random variation was related to variance between stands. Predictions derived from the models for density and juvenile wood in both species, and heartwood in pine showed good agreement when validated with data sets from two other studies. The resulting models may be used for predicting wood properties in forest planning and in bucking computers in harvesters, provided that the essential information is available.     

Snow-packing as a potential harmful factor on Picea abies,Pinus sylvestris and Betula pubescens at high altitude in northern Finland

Snow-packing, a combination of ice, hoarfrost and snow on trees, and the subsequent tree damage by snow, were estimated on Norway spruce (Picea abies), Scots pine (Pinus sylvestris), and pubescent birch (Betula pubescens) in the winter of 1993-94 in southern Lapland, northern Finland, near the local alpine timberline around 400 m a.s.l. Snow-packing on each fully sized tree increased from a few hundred kilograms at 150–250 m a.s.l. to a maximum of 3290 kg at 300–350 m a.s.l. At 300 m a.s.l., snow-packing per metre of stem increased from 30 to 50 kg on trees < 5 m in height up to 180–200 kg on 20-m trees. There was 300 000-480 000 kg/ha of snow accumulated on tree crowns. No stem breakage by snow-packing occurred at or below 250 m a.s.l., whereas at 290–350 m a.s.l., 0-46%, 39-100%, and 0–33% of the spruce, pine and birch trees, respectively, had broken tops. Birch appeared to be the most resistant and pine the most susceptible to snow breakage.     

Effects of growth regulators on germination of picea abies and Pinus sylvestris seeds

Nine seed lots each of Norway spruce (Picea abies L.), and Scots pine (Pinus sylvestris L.) were treated with combinations of the growth regulators: Indole‐3‐acetic acid, indole‐3‐butyric acid, etephone, kinetin, fusicoccin, and the gibberellins A₁ A₃ A_4/7, and A₉. GA₉ GA_4/7 and fusicoccin significantly promoted the germination of five Norway spruce, and three Scots pine, seed batches. Independent of species, the germinability of seeds treated with auxins, etephone, kinetin, GA₁ and GA₃ were unaffected or reduced. The rate of germination was stimulated for both species when treated with GA₉ or GA_4/7. The germination percentage for Scots pine seeds was enhanced by GA₉ but not by GA_4/7. Neither of the two GAs enhanced the germination percentage of Norway spruce seeds.     

Distribution and yield of biomass from young Pinus sylvestris and Picea abies stands on drained and fertilized peatland

Above‐ground biomass has been measured on fertilized and control plots up to stand age 31 years. Each biomass fraction was estimated by two statistically coupled linear models with squared diameter at breast height as independent variable. All fractions except reproductive structures of pine and dead branches of spruce were estimated at high precision levels, R. between 0.93 and 0.99. The above‐ground biomass fractions per hectare could be precisely described by stem volume with bark. Application of the models for prognostic purposes is discussed. Annual above‐ground net biomass production in pine increased from 2.05 to 4.34 MT d.m./ha‐yr on control plots over 11 years, while plots given complete fertilizer mixture ranged from 6.75 to 9.09. Spruce stands with a nearly optimum water relationship and fertilizer programmes yielded from 9.50 to 11.86 MT d.m./ha‐yr. The combination of energy and timber production in highly productive peatland stands is discussed.     

Effects of exogenous gibberellin and auxin on shoot elongation and vegetative bud development in seedlings of Pinus sylvestris and Picea glauca

The hormonal control of stem unit (foliar appendage and axillary structure, if present, plus subtending internode) number and length was investigated in shoots of Scots pine (Pinus sylvestris L.) and white spruce (Picea glauca (Moench) Voss). Seedlings were treated with six gibberellins (GA1, GA3, GA4, GA5, GA9 and GA20) and two auxins (indole-3-acetic acid (IAA) and naphthaleneacetic acid (NAA)) when either neoformed growth was occurring or the terminal vegetative bud was developing. Hormones were applied by drenching the shoot tip, injecting the stem or spraying the foliage. Combined results for all three application methods indicated that shoot elongation in first-year seedlings (i.e., neoformed growth) was promoted in both species by GA1, GA3, GA4 and, less obviously, by GA9. This promotion was attributable to an increase in length, rather than number, of stem units. However, the number of stem units formed during terminal bud development, as reflected in the number of needles (white spruce) or cataphylls (Scots pine) present on the shoot resulting from the terminal bud, was stimulated by GA1, GA3 and GA4 in both species and by GA9 in Scots pine. The GA-induced increase in the number of preformed stem units was associated with increased bud width in white spruce and increased bud length and resulting shoot length in Scots pine. In contrast, application of IAA or NAA either did not affect or inhibited both neoformed growth and terminal bud stem unit number, depending on the application method and concentration. We conclude that, in the Pinaceae, (1) GA stimulates the activity of both the subapical meristem during neoformed growth and the apical meristem during vegetative bud development, and (2) the early non-hydroxylation pathway, via GA9, is the major route of GA biosynthesis. The role of auxin in the control of stem unit number and length remains to be resolved.     

Application of growth regulators in aqueous media and organic solvents to seeds of Picea abies and Pinus sylvestris

Incubation with acetone and dichloromethane had little effect on the germination of different lots of Norway spruce seeds. This was not the case with Scots pine where almost half the seed lots tested were adversely affected by acetone and dichloromethane. The auxins indole‐3‐acetic acid (IAA) and indole‐3‐butyric acid (IBA) were applied to Scots pine seeds in acetone, dichloromethane and phosphate‐citrate buffer. The uptake of the two compounds was most rapid from acetone. Although the uptake from dichloromethane was slower than from acetone it reached the same level after 6 h. Uptake of the growth regulators from buffer was significantly lower than from the organic solvents. The accumulation of IAA and IBA into the seeds from both acetone and dichloromethane was linear between 0.5 and 10 mM, and between 0.5 and 15 mM from buffer. IAA and IBA levels in infused seeds declined over a 5‐day germination period, the former more rapidly than the latter. The metabolic activity of the IAA degradation system was as effective in seeds after immersion in the organic solvents as seeds immersed in buffer.     

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.     

Influence of soil scarification on reindeer foraging and damage to planted Pinus sylvestris seedlings

Abstract

The effects of soil scarification on reindeer lichen cover and re-establishment, reindeer foraging and damage possibly caused by reindeer to planted Scots pine seedlings were studied during six growing periods in a field experiment established in a reindeer lichen site that was grazed by a herd of 500–600 reindeer each winter. Seedlings (at a density equivalent to 2000 ha^?1) were planted in mounds with mineral soil on top, in tracks with exposed mineral soil, in tracks with mixed organic material and mineral soil and in intact lichen mat (control) with no soil disturbance. The disturbed area varied from 0 to 28%. After six growing periods, the reindeer lichen cover and volume were 10–20% lower in the scarified plots than in the control plots. There was no clear evidence that the reindeer avoided foraging even in plots with the highest levels of soil disturbance. However, the behaviour of the reindeer during winter grazing seemed to be affected by both coverage of reindeer lichen (positively) and the proportion of exposed mineral soil (negatively). Damage possibly caused by reindeer trampling affected 8.2% of the living seedlings each year and approximately 50% of these seedlings were subsequently infected by fungal diseases (compared with 20% of viable undamaged seedlings). Scarification treatments that exposed mineral soil mildly resulted in higher survival and growth rates than the other treatments, especially the control.