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.     

Seasonal variation in root growth capacity during cultivation of container grown Pinus sylvestris seedlings

Root growth capacity (RGC) in Scots pine seedlings was studied from the time of sowing and during the following two growing seasons. The method used for measuring RGC is also described. In the first growing season root growth was intense during the period mid‐July to mid‐September with an earlier peak for early sowing dates. After a period of low growth activity during winter, RGC rose sharply in early spring. During periods of intensive shoot elongation in May and June root growth was depressed. After shoot elongation was completed, RGC rose again before declining during the autumn. During winter and the second growing season, higher RGC levels were obtained for seedlings sown in June compared to the ones sown in April. This result is discussed with regard to differences in cultivation regimes.     

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.     

Modelling incidence of root rot in picea abies plantations in Southern Sweden

An empirical model is presented to forecast the incidence of root rot at stand level. In addition, the impact of different thinning programmes on the incidence of root rot is evaluated. The model is based on data from 152 permanent forest research plots of pure Picea abies in southern Sweden, within which the incidence of root rot at stump height in thinned trees has been recorded after each thinning since 1950. In total, about 20 000 stumps have been studied. According to simulations with the model, areas previously used as fields or for grazing are particularly susceptible to root rot, while old hardwood sites are less susceptible. Furthermore, simulations with the model imply that the earlier, the harder or more often a stand is thinned, the faster will be the development of root rot.     

Tap root formation and early root/shoot ratio of Pinus Contorts and Pinus sylvestris

Scots pine and different provenances of lodgepole pine were compared in regard to root characteristics and the distribution of biomass within individual trees. Tap root formation was more frequent and the root/shoot ratio was higher on Scots pine than on lodgepole pine but significant variation of these traits was found between provenances of planted lodgepole pine. A non‐destructive method for estimation of root biomass was developed.     

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.     

Effects of different overwinter storage regimes on shoot growth and net photosynthetic capacity in Pinus Sylvestris Seedlings

Seedlings of Pinus sylvestris were cold‐stored for three or six months at ‐4°C or 2°C or overwintered outdoors. Dry weight development and net photosynthetic capacity were then measured during a 60‐day cultivation period in a controlled environment. In all storage regimes the longer storage period gave a faster growth initiation. Photosynthetic recovery was faster for seedlings stored at 2°C than at ‐4°C, due to better recovery of stomatal conductance. The results suggest that there is little difference in seedling development between storage temperatures of ‐4°C and 2°C whereas outdoor storage might cause certain negative effects on subsequent growth of seedlings. This result is discussed with regard to the present winter climate.     

Determination of mean diameter by basal area in stands of picea abies and pinus sylvestris in southeastern Norway by means of aerial photographs

Mean diameter by basal area (d_g) is an important stand variable for long‐term economic forecasts of forest holdings. In order to use stand‐by‐stand surveys based on aerial photo interpretation as the data basis for forecasts, d_g has to be determined. The objective was to develop and test a regression function for d_g in mature stands of Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.) applicable in southeastern Norway. A study of 700 plots was used to estimate a function for d_g . An additive model was found to be most suitable. The independent variables were potential yield capacity, Lorey's mean tree height, crown closure determined by ocular estimation by means of aerial photographs, and the product of potential yield capacity and crown closure. The R² value was 0.604 and the coefficient of variation was 10.8%. The regression fitted most parts of the calibration data quite well, but it may overestimate the mean diameter in pure spruce stands by 1–2%, and underestimate the diameter in pure pine stands by 3%. For mixed coniferous stands the regression seems satisfactory. Testing by means of an independent data set showed systematic errors of 3–23%. The systematic errors were due partly to calibration problems in connection with the ocular crown closure estimation.     

Planting performance potential of Pinus sylvestris seedlings as evaluated by root growth capacity and triphenyl tetrazolium chloride reduction methods

Root growth capacity (RGC) and root viability by the reduction of triphenyl tetrazolium chloride (TTC) of Scots pine (Pinus sylvestris L.) seedlings were measured in the winter and spring after freeze‐induced stress (0, ‐10, ‐15, ‐20°C). After freezing, seedlings were also stressed with different storage temperatures, and drying. Field performance during two years at two sites was used to determine the prediction abilities of the two test methods. RGC and TTC differed between the winter and spring tests. Coefficients of variation (CV) for RGC were larger in the winter than in the spring. The opposite applied for TTC. RGC was significantly lower in the winter than in the spring. TTC reduction was lower in the spring than in the winter. RGC‐ and TTC‐values decreased as freezing temperatures were lowered. Correlation coefficients between treatment means of RGC and TTC were low. Coefficients of determination (r ²) of linear regressions of RGC to the field performance attributes, first and second year diameter increment, height increment, and survival, were between 0.30 and 0.40. For TTC the revalues ranged from 0.50 to 0.80. In general, the TTC‐method seemed to be a better indicator of Scots pine seedling performance in the field.     

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.     

The occurrence and pathogenicity of uni- and binucleate Rhizoctonia and Pythiaceae fungi among conifer seedlings in Finnish forest nurseries

Uni- and binucleate Rhizoctonia and Pythiaceae fungi were isolated from roots of dieback seedlings. Only binucleate Rhizoctonia strains were present in the roots of healthy seedlings. In pathogenicity tests, uninucleate Rhizoctonia sp. were pathogenic and binucleate Rhizoctonia spp. non-pathogenic. Pythium ultimum var. ultimum and Phytophthora undulata were the only Pythiaceae fungi that killed 12-week-old seedlings.     

高浓度CO2下红松幼苗根系呼吸对土壤呼吸的贡献

本文于2003年5月至10月在长白山森林生态系统定位站内研究了高浓度CO2(500和700靘olmol-1)对红松幼苗土壤呼吸以及根系呼吸对土壤呼吸的贡献。经过4个生长季高浓度CO2的处理,利用LI-6400-09土壤呼吸室对红松幼苗土壤总呼吸和根系呼吸进行了测定。为了区分根系呼吸对土壤总呼吸的贡献,本文采用了PVC管断根法,即每种处理下将三根PVC管插入土壤中30cm以切断根系,从而终止了植物冠层对根系碳水化合物的供应。分别于6月16日、8月20日和10月8日对管内外土壤呼吸进行了测定。结果表明大气和土壤5cm温度都存在明显的日变化,但不同处理之间没有显著差别(P>0.05)。土壤总呼吸和断根土壤呼吸也有明显的日变化和季节变化。不同处理之间土壤总呼吸和断根土壤呼吸差异显著(P<0.01)。6月16日、8月20日和10月8日不同处理下土壤总呼吸和根系呼吸的贡献的平均值分别为3.26、4.78和1.47靘olm-2s-1以及11.5%、43.1%和27.9%。图5表1参38。     

Net photosynthetic response to light intensity of shoots from different crown positions and age in picea abies (L.) karst

Carbon dioxide uptake response to light intensity of detached shoots of open‐ and forest‐grown Norway spruces was investigated in controlled standard conditions. The initial slope of the CO₂ uptake light response curve, calculated on leaf area basis and the internal conductance, calculated on leaf weight basis were almost constant for shoots from different positions in the forest canopy. In open‐grown trees the maximum photosynthesis and internal conductance (on leaf weight basis) decreased in relation to the tree age, so that in the 8‐year‐old tree it was about two times of that in the 66‐year‐old tree. These results suggest that 1) the enhanced efficiency of “shade” shoots is caused by morphologic adaptation, i.e. the more sparsely packed photosynthetic apparatus in needles and shoots, and 2) adaptational possibilities of Norway spruce photosynthetic apparatus, its ecological plasticity diminish during tree ontogenesis.     

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.     

Frequencies and patterns of browsing by large herbivores on conifer seedlings

Browsing by large herbivores on planted and naturally regenerated conifer seedlings (Picea abies and Pinus sylvestris) was recorded in 104 clear‐cuts in east‐central Sweden during 1990 and in 47 clear‐cuts in 1991. The number of seedlings browsed and the browsing patterns were analysed in relation to seedling type. Browsing frequencies were also compared between forest stands with different site productivities and subjected to different management practices. The variation in the number of seedlings browsed in 1990 was explained mainly by seedling category. Among both planted and naturally regenerated seedlings, pine was browsed more than spruce. Two‐year‐old containerized seedlings of spruce was browsed more than 4‐yr‐old bare‐rooted spruce. In 1991, browsing was more equal among species and seedling types. Number of seedlings with their leader browsed and the amount of biomass left after browsing differed significantly between seedling types. Seedlings that had been browsed in 1990 experienced significantly higher browsing frequencies in 1991 when compared with unbrowsed seedlings. The effects of stand characteristics were not found to be significant.     

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.     

Shoot and root sensitivity of containerized black spruce,white spruce and jack pine seedlings to late fall freezing

Damage to containerized forest seedlings due to freezing can occur in the fall or early winter in Canadian forest nurseries. The following spring, damage to shoots and impairment of growth is observed. The objectives of this experiment were to measure the impact of late fall low temperatures (0° to --30°C) on whole seedlings of the three most common species used for reforestation in Quebec: black spruce (BS), white spruce (WS) and jack pine (JP). Impacts of freezing temperatures on (i) whole seedling and apical bud mortality, (ii) shoot growth and root mortality, (iii) stem electrical resistance, (iv) shoot and root water relations, (v) concentrations of N, P, K, Ca, Mg, and total sugars in shoots were assessed. JP showed the highest rate of whole seedling mortality while WS showed the highest rate of apical bud mortality. JP was the most severely affected: destruction of the root system at low temperatures as well as a reduction of shoot growth and stem diameter and a decrease (more negative) in shoot and root water potential. WS showed a reduction of shoot growth despite no apparent damage to the root system at low temperatures. BS was not affected by temperatures as low as --30°C. Nutrient and sugar concentrations were not affected by low temperature treatments.     

Field performance of containerized black spruce seedlings with root systems damaged by freezing or pruning

Roots of 2-year-old black spruce seedlings (Picea mariana [Mill.] B.S.P.) were exposed to freezing temperatures to destroy 20, 40, 60, and 80% of the root systems. For comparison, other root systems were pruned to eliminate the same proportions of roots. Treated and control seedlings were planted in spring 1992 at Forêt Montmorency (the Université Laval forest research station, 50 km north of Quebec City). From 1992 to 1995, survival, shoot height and stem diameter were measured. Mortality mainly occurred in 1993, the year after planting; pruned seedlings showed practically no mortality while seedlings with 20, 40, 60, and 80% of their root systems affected by frost showed mortality rates of 0, 6, 17, and 24%, respectively. Shoot height and stem diameter decreased with increasing root damage when compared to controls. After three years on the planting site, shoot height was reduced by 2, 8, 11, and 18% while stem diameter was reduced by 4, 21, 25, and 24% for 20, 40, 60, and 80% frost damage, respectively. For pruned seedlings, shoot height was increased by 3% at 20% damage and was decreased by 1, 3, and 13% for 40, 60, and 80% root damage while stem diameter was reduced by 1, 4, 8, 19% for 20, 40, 60, and 80% respectively. Use of damaged seedlings on the planting sites should be limited in order to reduce the cost of replacement planting.     

Involvement of Cylindrocarpon destructans in root death of Pinus sylvestris seedlings: Pathogenic behaviour and predisposing factors

The common rhizospheric fungus Cylindrocarpon destructans was investigated in relation to its role in root death of Pinus sylvestris in Nordic nurseries and plantations. Laboratory methods were developed for studying similar root problems as well as the early effects of phytotoxicity and fungal infection. Seedlings grown under standardized optimal conditions were exposed to controlled stress (known to occur in nurseries), with or without C. destructans in the rhizosphere. Low light conditions, anaerobic root environment, and fungicide treatment were each found to predispose pine seedlings to invasion by the pathogen. The pathogen was very sensitive to competition as well as antagonism on the root, and fungicide‐induced inhibition of antagonists such as Trichoderma spp. also increased the severity of attack by the pathogen. To compete successfully, the pathogen would have to invade and dominate weakened roots prior to the arrival of saprophytes. Toxic metabolites produced by the pathogen weakened or killed nearby root tissues, and pathogen metabolites seemed to prevent saprophytes from taking hold in the infected roots. Such heavily infected, dead roots may act as inoculum sources allowing the pathogen to invade adjacent living roots, even healthy ones. Dead roots left in nursery soil after earlier harvests may act as reservoirs of inoculum for long periods and pose a threat to new plants.     

Frost hardiness gradients in shoots and roots of picea mariana seedlings

Frost hardiness of tissues along the length of the stem and the root was investigated in first‐year black spruce (Picea mariana (Mill.) B.S.P.) seedlings. Frost hardiness of 1 cm long stem and root segments was evaluated based on Index of Injury, calculated from post‐freezing electrolyte leakage. Frost hardiness was tested approximately weekly beginning seven weeks after seedlings were transferred from an 18 to a 10 h photoperiod, both at day/night temperatures of 26°C/16°C. Trees were transferred to temperatures of 10°C day and 5°C night at a 10 h photoperiod after a further 18 days. Frost hardiness was greater at the terminal bud and least at the root tips. Although shoots were generally more frost hardy than roots, differences in hardiness along the stem and root axes were gradual, rather than abruptly differing at the shoot‐root interface. All tissues, including root tips, increased in frost hardiness after conditioning for 18 days under short photoperiods (10 h) and warm temperatures (26?C/16°C, day/night). Under cold temperatures (10°C/5°C, day/night) all tissues, excepting the root tips, tolerated — 16°C with little subsequent electrolyte leakage.