Relationship between height growth of Scots pine (Pinus sylvestris L.) and site factors in Great Britain

The rate of height growth in Scots pine stands throughout Great Britain was examined in relation to site factors. The site factors included measures of geographical variation, topography, soil chemical and physical variables at two levels, several measures of soil phosphorus status, foliar monoterpenes and estimates of mean values of climatic variables. Principal component analysis was used to make an initial selection of regressor variables from 88 site factors for use in multiple regression equations. A dependent variable was obtained by removing the effect of age on height. The dependence of this expression on site variables was then examined using step-up multiple regression, starting with the selection of regressor variables made by the principal component analysis (P.C.A.), then adding transformations and interaction terms until 69% of variation was significantly accounted for over Great Britain, and up to 99% for parts of the country. Simpler equations using only those regressor variables which could be read from maps were also calculated.Variations in growth over Great Britain are associated mainly with solar radiation, soil texture and soil moisture content. For separate parts of the country the solar radiation term disappears from regression equations.     

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.     

Susceptibility of four conifer species to Gremmeniella abietina

G. abietina causes severe dieback in pole-stage stands of Scots pine in Britain. The susceptibility of ll provenances of Scots pine and of three other conifer species was tested by inoculations. Significant differences in susceptibility were found amongst Scots pine provenances, with those originating from a native Scottish pinewood at Loch Maree showing the highest levels of disease. Corsican pine was shown to be the most susceptible species tested although sporulation was greatest on Scots pine. Norway spruce was also shown to be susceptible when inoculated late in the growing season. Infection in lodgepole pine was negligible. High levels of β-phellandrene were tentatively linked with resistance among trees in a single stand of Scots pine.     

Dendrochronological analysis of the growth and growth-climate relationships of conifers in the region of alkaline dust deposition

Relationships between climate and radial growth of Oxalis-Myrtillus-site type Norway spruce and Scots pine stands under different cement dust loads were investigated. Dendrochronological methods were used. Long-term alkaline (pH 13.2-12.7) dust pollution emitted over 40 years from a cement plant was the reason of alkalisation (pH 7.8-8.1) and high concentrations of K, Ca and Mg in soil of affected territories. Two study sites, Kunda and Malla (2.5 and 5.0 km E from the emission source), were influenced by the dust emissions of the cement plant. Two other study sites, Eru (38 km W) and Revoja (34 km W), were situated on a relatively unpolluted area and served as control sites. The relationships between the radial growth and climate were almost similar on the control and dust-polluted sites. The climatic variables that had a significant effect on the radial growth of Norway spruce at both control and polluted sites were the temperature and precipitation of summer months of the current and preceding years. Specific to the Scots pine stands growing under dust pollution was a significant positive impact of the current spring temperature on the radial growth. A positive effect of precipitation during winter months on the radial growth was found at all sites. A significant negative effect of cement dust on the radial growth of Scots pine stands during the period of large amounts of dust emission (1966-1991) was detected. The effect of cement dust emissions on the radial growth of Norway spruce stands was also negative but weak. The decrease in the dust emissions since 1992 improved the growth conditions at pine stand sites.     

Seasonal acclimation of photosystem II in Pinus sylvestris. II. Using the rate constants of sustained thermal energy dissipation and photochemistry to study the effect of the light environment

Photosynthesis in evergreen conifers is characterized by down-regulation in autumn and rapid up-regulation in spring. This seasonal pattern is largely driven by temperature, but the light environment also plays a role. In overwintering Scots pine (Pinus sylvestris L.) trees, PSII is less down-regulated and recovers faster from winter stress in shaded needles than in needles exposed to full sunlight. Because the effect of light on the seasonal acclimation of PSII has not been quantitatively studied under field conditions, we used the rate constants for sustained thermal energy dissipation and photochemistry to investigate the dynamics and kinetics of the seasonal acclimation of PSII in needles exposed to different light environments. We monitored chlorophyll fluorescence and needle pigment concentration during the winter and spring in Scots pine seedlings growing in the field in different shading treatments, and within the crowns of mature trees. The results indicated that differences in acclimation of PSII in overwintering Scots pine among needles exposed to different light environments can be chiefly attributed to sustained thermal dissipation. We also present field evidence that zeaxanthin-facilitated thermal dissipation and aggregation of thylakoid membrane proteins are key mechanisms in the regulation of sustained thermal dissipation in Scots pine trees in the field.     

The effects of species mixture on the growth and yield of mid-rotation mixed stands of Scots pine and silver birch

The effect of tree species mixture on stand volume yield and on tree-species-specific diameter and height growth rates were analysed in managed mixed stands of Scots pine (Pinus sylvestris L.) and silver birch (Betula pendula Ehrn.).Data were obtained from 14 repeatedly measured stands located in Southern Finland on mineral soil sites with varying admixture of Scots pine and silver birch. Statistical analysis was carried out for studying the effect of species mixture on the development of stand characteristics. For the analysis, the plots were categorised into three groups (plot types) according to the species dominance. In order to analyse species-specific growth rates, individual-tree mixed linear growth models for tree diameter and height growth were developed for both tree species.The results clearly show that the yield of the managed mid-rotation, mixed stands was greater for stands dominated by Scots pine than for stands dominated by birch, and the stand volume increment decreased with an increasing proportion of silver birch. Analysis of diameter and height growth by tree species revealed that the main reason for this pattern is the negative impact of birch competition on the growth of pine trees. The increase in diameter of pine was clearly hampered if the proportion of birch was high. An abundance of birch also slightly decreased the growth in height of Scots pine, although the effect was less than on diameter growth. Species mixture did not affect the diameter growth of birch but did have a significant effect on height development. Height growth of birch was considerably greater in pine-dominated stands than in birch-dominated stands. In pine-dominated mixed stands, the height growth of birch was quite close to that of dominant pine trees, and birches can endure in competition with pines for light.The results apply for even-aged and single-storey managed stands, where stocking density and structure are controlled with pre-commercial and commercial thinnings. The results are not applicable to unmanaged mixed stands undergoing self-thinning. This study provides new information on mixed stands from a silvicultural perspective, which can be applied in decisions involving the management of mixed stands.     

Structure and abundance of arachnid communities in Scots and lodgepole pine plantations

Between April and September 1994, six plots within Aberfoyle Forest in Scotland, three of Scots (Pinus sylvestris) and three of lodgepole pine (Pinus contorta), were surveyed for spiders and harvestmen, as these are predators of several forest invertebrate pests.A total of 64 species of spider were trapped in Scots and lodgepole pine; 56 in Scots pine and 32 in Lodgepole pine. The most abundant spiders, both in Scots and Lodgepole pine, were Linyphiidae.Scots pine plots were floristically more diverse and structurally more complex than lodgepole pine plots. More species of spiders and harvestmen (midsummer only) were found at ground level in Scots pine than lodgepole pine plots. More species and a greater numbers of spiders were found in the canopy of Scots pine than in that of lodgepole pine. However, no differences in the abundance of harvestmen were found between the canopies of Scots and lodgepole pine. Spider species diversity was greatest in Scots pine.     

Browsing by large herbivores on Scots pine (Pinus sylvestris) seedlings in mixture with ash (Fraxinus excelsior) or silver birch (Betula pendula)

Abstract

Food selection by large herbivores occurs at a hierarchy of scales, for example landscape, patch or plant. Several hypotheses regarding food plant selection on patch or plant level have been developed. In this cafeteria-type design field experiment, conducted during one winter immediately after planting, we tested the effect of species mixture on browsing by large herbivores (mainly roe deer) on Scots pine seedlings in mixture with seedlings of ash (highly preferred) or silver birch (less preferred). Browsing on Scots pine was not affected by species mixture, neither in terms of the number of browsed pines nor browsing intensity. Instead, browsed biomass was positively and significantly correlated to the total biomass available for browsing. Also, there were differences due to species, with ash being most browsed (44.6%), followed by Scots pine (18.9%) and silver birch (11.6%). Browsed biomass per browsed seedling, however, was largest for Scots pine. In addition, browsed seedlings were initially taller compared to unbrowsed seedlings for all species. The main management implication in this study is that the species mixture did not influence large herbivore browsing on Scots pine seedlings. Hence, removing or discouraging more (or less) attractive browse species in early stages of pine regeneration activities seems unnecessary from the point of large herbivore browsing.     

Impact of solar activity on the growth of pine trees: case study

In this work, we focus on the analysis of data for the ring width of two pine tree species: samples of six maritime pines (Pinus pinaster), grown in the north of Portugal, and one sample of Scots pine (Pinus sylvestris), grown in the south of Slovakia. A negative impact of solar activity (SA) was observed on the growth of these pine trees with particularly strong impact in the case of the maritime pines. The width of the annual rings was generally smaller in the years of maximum SA; furthermore, it was found that it is latewood width that is affected whereas earlywood width is not affected; as a corollary, the percentage of latewood also shows a significative negative correlation with SA.     

Relationship between height growth of stand and open-grown single trees of Scots pine (Pinus sylvestris L.) and site factors in Great Britain

The relationship between height growth of Scots pine and site factors was examined for single trees growing in plantations and open situations, in order to examine the complete range of variation in competition, throughout Great Britain. As in the preceding paper (White, 1982) reporting a study of stand growth, a wide range of variables was recorded expressing geographical, topographical, soil chemical and physical variables and climatological change, as well as differences in genotypes expressed by levels of monoterpenes. Additional variables expressing competition were added in this study. A principal component analysis was used to determine the main sources of variation in the site variables, and to make an initial selection of regressor variables for multiple regression. Multiple regressions were then calculated of the rate of height growth on the selected variables plus transformations and interaction terms. An equation accounting for 63% of the variation shows height growth rate to depend on competition effects, rainfall in April to June, and soil water holding capacity at 0–5 cm. This result is considerably different from equations in the preceding paper. A simpler equation which can be used for prediction without site visits, but which accounts for only 46% of variations is also presented.     

Using cattle for conservation objectives in a Scots pine <Emphasis Type="Italic">Pinus sylvestris</Emphasis> forest: results of two trials

It has been argued that large ungulates play a key role in natural forest dynamics, but in Britain, the largest native ungulates (aurochs and elk) are extinct. Cattle could have some similar effects, and are widely used, but rarely tested, for nature conservation management. Here, we test conservation management with cattle at a native Scots pine Pinus sylvestris forest in Scotland. Our hypotheses were that cattle impacts would (a) increase the abundance of an understorey shrub of conservation importance, bilberry Vaccinium myrtillus; and (b) increase Scots pine seedling establishment. Two cattle trials were established, one for each objective, based on a 14-ha plot within the forest, and a 6.1-ha plot at the forest edge. Cattle were introduced for 2 months to both plots, giving an approximately 100- to 150-fold increase in ungulate biomass per hectare when compared to background deer abundance. The ground layer vegetation in both treatment and matched control plots was monitored before and after treatment, and subsequently over a 4-year period. At the within-forest trial, bilberry percent-cover, 4 years after treatment, was 1.9 times higher in the cattle area than in the control (95% confidence limits 1.6–2.3). Bilberry percent-cover increases were directly related to the degree of trampling impact on heather Calluna vulgaris, as recorded immediately after the trial. At the forest-edge trial, there were vegetation changes in the cattle plot that were considered favourable to pine seedling establishment: reduced moss/litter depth and vegetation percent-cover, and increased ground-level light incidence. However, too few new pine seedlings were found for formal analysis, partly due to unexpectedly low seed-fall. Nevertheless, the potential for cattle to create conditions that would subsequently promote pine regeneration was illustrated by the observation of large numbers of new, young pine seedlings at the within-forest cattle plot. We recommend that managers and researchers collaborate to develop further trials like the ones reported here, to give an improved understanding of the conditions under which cattle can yield nature conservation benefits in woodlands.     

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.     

Annual and spatial variation in coning shown by the Forest Condition Monitoring programme data for Norway spruce, Sitka spruce and Scots pine in Britain

Conifer seed is a valuable forest resource; as well as beingreproductive material, it is an important foodstuff for manywoodland species. Information on cone and therefore seed productionof the main forest species are useful in the fields of bothforest management and forest ecology. Coning has been recordedsince 1989 in plots of Norway spruce, Sitka spruce and Scotspine across Britain as part of the Forestry Commission's ForestCondition Monitoring programme. An 11-year period of these recordswas analysed and a summary of annual cone production by speciesand coning synchrony is presented. Both Norway spruce and Sitkaspruce coned sporadically, with a 4-year period between mastcrops during which cone density was very low or cone productionabsent, whereas Scots pine produced a good cone crop in mostyears throughout Britain. Coning was synchronous within andbetween Norway and Sitka spruce across the whole of Britain(at distances >600 km) but coning within Scots pine was insynchrony only over distances of up to 200 km. Synchrony ofcone production was not evident between the spruce species andScots pine. These findings have implications when planning andinterpreting monitoring used to predict cone crops and in thedesign of forests for wildlife conservation.     

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.     

CLIMATE AND SILVICULTURAL IMPLICATIONS IN MODIFYING STAND COMPOSITION IN MIXED FIR-PINE STANDS

ABSTRACT

A growing concern exists over the decreasing proportion of Scots pine (Pinus sylvestris L.) trees in mixed Trojan fir (Abies nordmanniana subsp. equi-trojani [Asch. & Sint. Ex Boiss] Coode & Cullen)-Scots pine forests in northern Turkey. Given the changes in mean annual temperature and annual precipitation for the last 40 years in the region, understanding of the sensitivity of Trojan fir and Scots pine trees to changing climate seems to be critical so that better management strategies can be developed against the potential climate extremes of the future in the region. Moreover, our knowledge on the influence of current silvicultural methods on seedling density and growth in these forests is limited. Therefore, in this study, the growth response (i.e., radial growth) of Trojan fir and Scots pine trees to the changing climate were examined. In addition, the influence of current silvicultural practices on seedling growth and density in these forests was also monitored. Trojan fir appeared more sensitive to the climatic variables (i.e., temperature and precipitation) than Scots pine in the region. It was also found that the current silvicultural practices might favor Trojan fir over Scots pine in these forests. Natural dynamics and current silvicultural implications would benefit Trojan fir, however, given the species’ potential sensitivity to the changing climate, current silvicultural approaches can be modified to favor Scots pine, and to increase the resilience of these forests against climate extremes expected in the region’s future.     

Differences in delta13C and diameter growth among remnant Scots pine populations in Scotland

Published data suggest that differences in wood cellulose carbon isotope composition (delta13C) and xylem ring width among natural populations of Scots pine in Scotland (Pinus sylvestris L.) are attributable to the persistence of palaeotypes of various post-glacial migratory origins. We assessed differences in wood cellulose delta13C and ring width among Scottish Scots pine populations grown in a clone bank and in natural stands at various locations in northern and central Scotland. Ring width and wood cellulose delta13C varied significantly among natural stands. Potential water deficit was positively correlated with wood cellulose delta13C and xylem ring width in the natural stands. Neither wood cellulose delta13C nor xylem ring width of clone bank trees correlated with any climate variables at the sites from which the trees originated, indicating little adaptation to climate for these traits. Xylem ring width showed a site x population interaction for the growth sites (i.e., natural stands versus clone bank), but wood cellulose delta13C did not. These results suggest that climate variation in Scotland has not resulted in significant genetic variation in wood cellulose delta13C or xylem ring width in post-glacial populations.     

The impact of cronartium flaccidum on the growth of pinus sylvestris

The blister rust of two‐needle pines in Europe is caused by the rust fungus, Cronartium flaccidum (Alb. Schw.) Wint. There are two races of the fungus: One host‐alternating and another pine‐to‐pine race. The latter race is considered to be more common in northern Sweden. The impact of this rust on growth of Scots pine (Pinus sylvestris L.) was investigated in three selected stands in northern Sweden. Radial stem increment was reduced 40–70% by severe attacks and 20–40% by minor attacks. The reduction of stem volume growth was probably greater. The faster growing trees seemd to be more severely attacked. Trees growing on poor soils appeared to lose almost as much growth capacity from a minor attack as from a severe attack.     

Evidences of drought stress as a predisposing factor to Scots pine decline in Valle d’Aosta (Italy)

Scots pine (Pinus sylvestris L.) forests of many inner Alpine valleys have recently displayed a quick loss of vitality. A decline disease has been suggested as the cause, with drought as the main predisposing factor and the additional contribution of biotic agents inciting tree dieback. This study is focused on Valle d’Aosta, a dry, inner-Alpine region in NW Italy. We inferred vitality changes between years 2000 and 2007 by computing reductions in enhanced vegetation index (EVI). Image differencing was carried out on pre-processed Moderate Resolution Imaging Spectroradiometer (MODIS) imagery taken in late springtime and validated against ancillary ground truth. We: (1) tested whether EVI reductions in Scots pine forests were significantly higher than those of a control species and of a wetter region for the same species, (2) analyzed decline incidence as a function of site and topographic variables, and (3) assessed the relative influence of site and stand structure on decline probability by means of path analysis. Mean EVI in the study area increased due to an early onset of the 2007 growing season. Nevertheless, the incidence of decline was 6.3% and significantly greater for Scots pine than the control species and site. Low-elevation, northerly exposed sites exhibited the highest incidence of decline. Path analysis suggested that the most important determinants of decline probability were slope, solar radiation, and stand sparseness.     

The response of Pinus sylvestris to drought: stomatal control of transpiration and hydraulic conductance

We investigated the impact of drought on the physiology of 41-year-old Scots pine (Pinus sylvestris L.) in central Scotland. Measurements were made of the seasonal course of transpiration, canopy stomatal conductance, needle water potential, xylem water content, soil-to-needle hydraulic resistance, and growth. Comparison was made between drought-treated plots and those receiving average precipitation. In response to drought, transpiration rate declined once volumetric water content (VWC) over the top 20 cm of soil reached a threshold value of 12%. Thereafter, transpiration was a near linear function of soil water content. As the soil water deficit developed, the hydraulic resistance between soil and needles increased by a factor of three as predawn needle water potential declined from -0.54 to -0.71 MPa. A small but significant increase in xylem embolism was detected in 1-year-old shoots. Stomatal control of transpiration prevented needle water potential from declining below -1.5 MPa. Basal area, and shoot and needle growth were significantly reduced in the drought treatment. In the year following the drought, canopy stomatal conductance and soil-to-needle hydraulic resistance recovered. Current-year needle extension recovered, but a significant reduction in basal area increment was evident one year after the drought. The results suggest that, in response to soil water deficit, mature Scots pine closes its stomata sufficiently to prevent the development of substantial xylem embolism. Reduced growth in the year after a severe soil water deficit is most likely to be the result of reduced assimilation in the year of the drought, rather than to any residual embolism carried over from one year to the next.     

Photosynthetic utilization efficiency of absorbed photosynthetically active radiation by Scots pine and birch forest stands in the southern Taiga

Absorption and utilization of photosynthetically active radiation (PAR) were investigated in Scots pine (Pinus sylvestris L.) and birch (Betula pendula Roth.) stands that were 41 years old at the end of the experimental period. Canopy depth of the Scots pine stand was about half that of the birch stand (6.5 versus 11.0 m), but absorption of PAR was similar in the two stands. The Scots pine forest canopy, with a leaf area index of 8.9, absorbed 90% of the incoming PAR (APAR), whereas the birch forest canopy, with a leaf area index of 5.9, absorbed 92% of APAR. During maximum foliage development, the upper Scots pine canopy absorbed more PAR than the upper birch canopy (75 versus 66%). The upper, middle and lower layers of the Scots pine canopy contained 37, 48 and 15% of the total needle surface area, respectively. The corresponding distribution of foliage surface area in the three layers of the birch canopy was 50, 30 and 20%, respectively. Measurements of photosynthetic rate were combined with estimates of leaf area index and stand phytomass to determine rates of primary production on a sunny day, a cloudy day, and on an annual basis. The energy equivalents of short- and long-term carbon gain were used with determinations of APAR to calculate photosynthetic utilization efficiency. Throughout the growing season, photosynthetic utilization efficiency of APAR in the upper canopy layer of the Scots pine forest was almost twice that in the lower canopy layer. In the birch forest, photosynthetic utilization efficiency was greater in the lower canopy layer than in the upper canopy layer. In all cases, utilization efficiency was higher in the birch stand than in the Scots pine stand (52 versus 29 J kJ(-1)). Taking account of respiration of the non-photosynthetic parts of each stand (night respiration of needles or leaves; respiration of branches, trunk and roots), estimated utilization efficiency of APAR for net primary production was 11 J kJ(-1) for Scots pine and 19 J kJ(-1) for birch. Solar conversion ratios, expressed as whole-plant net primary productivity per unit of APAR for the growing season, were 0.81 g MJ(-1) for Scots pine and 0.93 g MJ(-1) for birch.