Effects of browsing on recruitment and mortality of European aspen (Populus tremula L.)

Abstract

High browsing pressure may reduce or halt regeneration of browsing-prone tree species. In this study, exclosures were used to quantify the effects of browsing on vital demographic parameters in European aspen (Populus tremula L.) stands. The experiment focused on stands located in managed forest landscape with densities of 0.85 moose (Alces alces L.)km^?2. Other herbivores such as mountain hare (Lepus timidus L.) were present at densities common for this region and roe deer (Capreolus capreolus L.) at low densities. Recruitment and mortality of aspen ramets were recorded before and after the growing season, and over four consecutive years. Seven to 19% of the ramets within browsing reach were browsed annually. No differences were found in recruitment or mortality attributable to browsing per se. Instead, there were large among-stand variability in vital population parameters unrelated to browsing incidence. In general, aspen stands with high ramet density showed higher recruitment rates than stands with low ramet density. There was substantial variability in self-replacement capability unrelated to ecological factors. The results suggest that more attention should be given to finding and managing stands with high capacity for suckering, as these stands may have a disproportionate impact on aspen recruitment at the landscape scale.     

European aspen (Populus tremula L.): a review with particular reference to Scotland II. Values, silviculture and utilization

Interest in European aspen (Populus tremula L.) has increasedin recent years as a result of efforts to restore and extendnative woodland, and to diversify the range of species usedin forestry plantations. Aspen is a straightforward tree tomanage; it is of considerable ecological value and capable ofproducing utilizable timber. Planting of aspen in native woodlandis currently limited by a shortage of plants of Scottish origin.Propagation from seed, whilst possible, has not been practised,and recent efforts to produce stock have concentrated on vegetativemeans. The use of aspen as a component of plantation forestrycould be increased on appropriate sites; it is an excellentamenity species and its timber is currently acceptable as chipwoodand could complement the production of poplar timber in thelowlands. However, on the basis of limited data, growth ratesand stem form of aspen in Scotland appear to be rather poorerthan on the European continent where aspen is a significanttimber producing species. More would also need to be known aboutits susceptibility to disease before its timber production potentialcould be evaluated properly.     

European aspen (Populus tremula L.): a review with particular reference to Scotland I. Distribution, ecology and genetic variation

European aspen (Populus tremula L.) has attracted relativelylittle research attention in Britain. Interest in the specieshas increased in recent years because of efforts to restoreand extend native woodland, of which aspen is an infrequentbut important component species. This paper draws together informationfrom British, European and Russian literature on the ecologyand genetics of aspen. The species is distributed throughoutBritain in many different types of woodland. It is particularlycommon in the Scottish Highlands and Islands where it occurson a diverse range of sites from sea cliffs to near the treeline.It is associated with a distinct insect fauna, several speciesof which are considered endangered. Information is lacking onseveral fundamental aspects of the ecology and genetics of aspenin Scotland including seed production, growth rates, geneticvariation and its conservation value and status in native forestecosystems. Some useful information on these topics is presentedfrom studies of aspen elsewhere in Northern Europe, particularlyNorway.     

Application of percolation modelling on end-grain water absorption in aspen (Populus tremula L.)

Abstract

The aim of this study was to improve our understanding of the capillary uptake of water in aspen sapwood and heartwood, which may help to explain the variable performance of aspen used outdoors. The study uses a percolation model developed for softwoods to examine capillary liquid absorption in aspen and compares predicted results with ones obtained experimentally using computed tomographic scanning. The study shows that there is an equilibrium saturation level where new flow paths are being found at the same rate as old ones are being blocked. This is seen as a plateau where the water content maintains a relatively constant height within the material. This makes it possible to model the capillary behaviour of aspen sapwood. In heartwood, however, the uptake of moisture seems to be mainly restricted to bound water, except for a short region in the sample wet end. This absorption is thus governed mainly by bound water diffusion. This improved understanding of the capillary behaviour of aspen may contribute to a future market expansion where aspen may be utilized to a greater extent outdoors and above ground.     

Killing effect and translocation of picloram in the stem of aspen,Populus tremula

Picloram was injected to different depths in the stems of aspen trees during spring and summer, and the killing effect on the foliage was studied. The efficacy of the applied dosage was correlated better to the surface area and volume of the stem than to the diameter. The young foliage was affected first by the herbicide. Both the xylem and the phloem were involved in the effects that resulted from translocation of the herbicide.     

On the reproduction of aspen (Populus tremula L.) with emphasis on its suckering ability

Numerical data are presented that support the proposal mentioned as early as the mid‐19th century that aspen roots may remain alive for a long period after the disappearance of parent trees. The mechanism that makes this possible is the ability of this species to form suckers continuously. In stands the bulk of the suckers live a few years and are then replaced by new ones. If conditions allow some suckers may develop into big trees even in old stands. The conclusions are based on material obtained in experiments comprising chemical control of aspen, including different application methods, herbicides, doses and in some cases control treatment (cleaning).     

Characterization of BspA, a major boiling-stable, water-stress-responsive protein in aspen (Populus tremula)

We identified a novel 66 kDa boiling-stable protein (BspA) in cultured shoots of aspen (Populus tremula L.) which was highly expressed in response to gradual water stress. The BspA protein, which was highly expressed as early as 1 h after initiation of a drought treatment, accumulated during progressive water stress, decreased on rehydration, and was expressed in response to abscisic acid (ABA) application, as detected by SDS-PAGE protein analysis and Western blotting. Anti-BspA antibodies also cross-reacted with a 119 kDa protein. The 119 kDa protein was also induced by water stress, but it was detected only in the total protein fraction and not in the heat-stable fraction. The BspA protein cross-reacted with antibodies raised against a water-stress-responsive protein isolated from the African resurrection plant Craterostigma plantagineum Hochst. The N-terminal amino acid sequence of BspA was determined and exhibited high homology with the wheat germins GF-2.8 and GF-3.8. The BspA protein was the only major, water-stress-responsive boiling-stable protein detected in aspen.     

Developmental factors responsible for heterosis in aspen hybrids (Populus tremuloides x P. tremula

Juvenile growth and bud set phenology were analyzed to study the biological basis of heterosis (hybrid vigor) in interspecific hybrids of Populus tremuloides Michx. (T) and P. tremula L. (Ta). Growth, measured as seedling volume index, was significantly higher for each of the two reciprocal interspecific crosses, T x Ta and Ta x T, than for the T x T intraspecific cross. Broad-sense heritabilities were 2-6 times larger than narrow-sense heritabilities for growth and shoot components in the T x T intraspecific cross, suggesting an important role for dominance or overdominance in aspen growth. Previous genetic analyses have indicated that hybrid vigor may be the result of overdominance at several key loci each with an allele inherited from each of parental species. Internode length and leaf number contributed substantially to the heterosis of stem volume, but their effects on heterosis differed between the T x Ta and Ta x T hybrids. In T x Ta seedlings, heterosis of stem volume was attributed to a high diameter growth rate, whereas in Ta x T seedlings heterosis of stem volume was probably the result of delayed bud set resulting in a longer duration of height growth. In addition to internode number and length and leaf number, other morphological or physiological components might affect heterosis, for example, extended leaf retention.     

Productivity and thinning effects in hybrid aspen (Populus tremula L. x P. tremuloides Michx.) stands in southern Sweden

The productivity of hybrid aspen stands in southern Sweden,originating from clone selections performed during the 1980s,was investigated. Thirteen former research and demonstrationsites were included. Repeated measurements of tree diametersand heights were taken, from which yields were estimated. Theresults indicate that the mean annual increment, obtained withoutany artificial addition of fertilizers or irrigation, will exceed20 m³ of stem wood ha^–1 a^–1 during a 20–25-yearrotation period. This corresponds to a total average woody biomassproduction, including branches, of over 8 tonnes dry matterha^–1 a^–1. These production levels show that hybridaspen is a competitive alternative for short rotation forestryin Sweden, and that the clone selection programme has greatlyenhanced productivity, compared with published growth ratesof the material previously used. For comparison, two standsthat regenerated via root suckers from material used in thebreeding programme up to the 1960s were included. They showedhigher growth than the 13 planted stands, mainly due to higherstem density and faster height development. This indicates thateven further increases in hybrid aspen yield are possible instands derived from suckers of the most recently selected clones.Thinning studies, including three different weights of thinning,were carried out on five of the sites. Five years after thestart of the treatments, stands given no thinning showed significantlyhigher current annual increment, but significantly smaller meanstem diameter than stands in which thinning was applied. However,the faster diameter development (and thus higher volumes ofvaluable wood assortments) associated with heavier thinningsmay compensate economically under current market conditionsfor the loss in total volume production.     

Parasitoids of Saperda populnea (L.) (Coleoptera: Cerambycidae) on aspen (Populus tremula L.) in Bulgaria

The parasitoids of Saperda populnea (L.) (Col.: Cerambycidae) were studied in Bulgaria during the period 1997?–?2001. Galls with pest larvae were collected from young aspen trees at eight locations (Sofia, Kokaliane, Plana, Churek, Gorni Lom, Gintzi, Dolno Kamartzi and Klisura) and examined under laboratory conditions. Four species were reared from 1118 galls containing overwintering pest larvae: Iphiaulax impostor (Scop.) (Hym.: Braconidae), Dolichomitus populneus (Ratz.), Schreineria populnea (Gir.) (Hym.: Ichneumonidae) and Billaea irrorata (Meig.) (Dipt.: Tachinidae). All of the parasitoids were solitary. Schreineria populnea was found as a new parasitoid of the host in Bulgaria. B. irrorata emerged mainly from late-stage S. populnea larvae; the remaining species from both early- and mid-stage larvae. In the parasitoid complex of S. populnea, the most numerous were B. irrorata (59.8?%) and D. populneus (29.4?%). The percentage of host larval parasitism varied from 2.4 to 33.3, with an average of 9.7?%. B. irrorata was the most important in reducing the pest number. The average mortality of S. populnea caused by the tachinid was 5.8?%, and the highest mortality observed in individual study was 20.2?%.     

Chemical characterization of Pinus halepensis sapwood and heartwood

ABSTRACT

This paper describes the chemical composition of sapwood (SW) and heartwood (HW) of Pinus halepensis Mill stem. Extractives were first isolated by accelerated solvent extraction and then analysed by gas chromatography-mass spectrometry (GC-MS). The cellulosic polysaccharide content present in the pre-extracted wood samples was determined with acid hydrolysis and GC. The hemicelluloses content was determined with acid methanolysis and GC. Free monomers were additionally analysed by GC. The amount of lignin was determined gravimetrically by the Klason lignin method and the acid-soluble lignin was determined by a UV method. Formic and acetic acids in wood were determined after alkaline hydrolysis and analysed by HP-SEC. It was found that lipophilic and hydrophilic extractives were more abundant in heartwood (1.6% and 2.5%) than in sapwood (1.1% and 1.8%). Celluloses content was higher in sapwood (42.5%) than in heartwood (39.7%), whereas lignin, hemicelluloses and sugar monomer contents were more abundant in heartwood (28.9%, 26.8% and 0.3%) than in sapwood (28.0%, 24.5% and 0.2%). The variation in acetic and formic acids and ash contents between sapwood (0.7%, 0.2% and 0.5%) and heartwood (0.6%, 0.1% and 0.4%) was small. The acetylation degrees were found to be slightly similar in sapwood (0.4) and heartwood (0.3).     

Sprouting of Populus tremula L. in spruce regeneration areas following alternative treatments

We investigated the efficiency of three different methods to restrict sprouting in European aspen (Populus tremula L.) in regeneration areas of spruce in Finland. The methods compared were (1) traditional method (cutting at 10?C15 cm height) (2) cutting at 1 m height, and (3) fungus treatment (cutting at 10?C15 cm height, and mycelium of a white-rot fungus, Chondrostereum purpureum [Pers. Ex Fr.] Pouzar was applied to freshly cut stumps). The results indicated that the fungus treatment can restrict the emergence and growth of stump sprouts better than other methods, whereas cutting at 1 m seemed to yield the lowest numbers of root suckers. However, differences between the treatments were not significantly different (P values were > 0.05). For future experimentation, we suggest that the cutting of higher stumps to control the number of root suckers should be combined with application of C. purpureum to restrict the emergence and growth of stump sprouts. We also suggest developing more efficient C. purpureum isolates to control European aspen stump sprouting.     

Peroxidase activity, isoenzymes and histological localisation in sapwood and heartwood of Scots pine (Pinus sylvestris L.)

Peroxidase activity and isoenzymes of fresh wood samples of the third shoot of 12-year old trees and from the sapwood, transition zone and heartwood of c. 60-year old stems of Scots pine (Pinus sylvestris L.) were investigated. Wood samples were ground at −30°C, extracted, and the extracts concentrated c. 20-fold for peroxidase activity assays (guaiacol method) and for IEF-PAGE. At least 11 major isoenzymes could be found in the gels. Even the heartwood contained some peroxidase isoenzymes. Isoenzyme patterns of the juvenile wood did not change with the season. However, juvenile wood showed the highest peroxidase activity at the end of the growing season. Peroxidase activity decreased from the outer sapwood towards the heartwood. Thin sections of different wood zones stained for peroxidase revealed activity in ray parenchyma and resin canal epithelial cells. Intensive staining was localised in the bordered pits of vertical and ray tracheids, and in the end walls of ray parenchyma cells.     

The decay pattern of Phellinus tremulae (Bond.) Bond,et Borisov in Populus tremula L.

The distribution of Phellinus tremulae in stems of Populus tremula was studied visually, macroscopically as well as microscopically, and by means of isolation of the fungus in pure culture. Evidence suggests that P. tremulae occurs alone in most of the tree. Microorganisms associated with P. tremulae in the wood are suggested to be independent invaders or secondary invaders. A dark reaction zone was formed in certain parts of the wood and consisted microscopically of melanized hyphes and brown material in the ray cells. An inner wood, formed as a result of the death of nearby brandies, is apparently the main part infected by P. tremulae, and its outer border may basically limit the spread of the fungus in the trunk. The formation of “heartwood” in aspen is discussed, and a suggested outline shown for the infection and decay processes caused by the fungus.     

Sexual Reproduction of European Aspen (Populus tremula L.) at Prescribed Burned Site: The Effects of Moisture Conditions

Aspen (Populus tremula) is capable of reproducing both sexually and asexually from root suckers, the latter being more common. Seedlings are rare even though potential seed production is enormous. The aim of this study was to find out how different moisture conditions affect emergence and survival of aspen seedlings on burned soil. We set up a sowing experiment on mineral soil and on humus. The sowing was done at 240 microsites (θ7 cm) in 15 blocks (1×1 m). Treatments were sowing time, watering and sowing shelter. A total of 1463 seedlings emerged on mineral soil and 26 on humus. Watering, covering, their interaction, block and sowing time had significant effects on the number of seedlings. The largest numbers of seedlings were recorded at microsites that were watered and covered. The survival of seedlings was 20% during the growing season and 45% over winter. We conclude that seedbed conditions and moisture are the most essential factors for sexual reproduction of aspen. Short-term weather conditions have an important effect on seedling emergence and are likely to greatly affect field studies.     

Studies on the epiphytic bacteria on Populus tremula L. and Populus alba L. at the time of optimal and minimal cambium activity and infection by Venturia

The participation of epiphytic bacteria in the groups of bacteria dominating on the shoots and root collars of two year aspen and white poplar differs both in quantity and quality and depends on the chemical composition of the diffusion exudates related to cambium activity. This indicates that epiphytic bacteria can utilize carbon sources in the exudates and transform lignin precursors.     

Genotypic variation in growth and physiological responses of Finnish hybrid aspen (Populus tremuloides x P. tremula) to elevated tropospheric ozone concentration

Saplings of six Finnish hybrid aspen (Populus tremuloides Michx. x P. tremula L.) clones were exposed to 0, 50, 100 and 150 ppb ozone (O3) for 32 days in a chamber experiment to determine differences in O3 sensitivity among genotypes. Based on the chamber experiment, three clones with intermediate sensitivity to O3 were selected for a free-air O3 enrichment experiment in which plants were exposed for 2 months to either ambient air (control) or air containing 1.3 x the ambient O3 concentration. We measured stem height and radial growth, number of leaves, dry mass and relative growth rate of leaves, stem and roots, visible leaf injuries, net photosynthesis and stomatal conductance of the clones. There was high clonal variation in susceptibility to O3 in the chamber experiment, indicated by foliar injuries and differential reductions in growth and net photosynthesis. In the free-air O3 enrichment experiment, ozone caused a shift in resource allocation toward stem height growth, thereby altering the shoot to root balance. In both experiments, low O3 concentrations tended to stimulate growth of most clones, whereas 100 and 150 ppb O3 in the chamber experiment impaired growth of most clones. However, growth of the most O3-tolerant clone was not significantly affected by any O3 treatment.     

Variation of extractives content in heartwood and sapwood of Eucalyptus globulus trees

The variation in extractives content in sapwood and heartwood was investigated among 12 trees in each of four commercial plantations of Eucalyptus globulus in central Portugal. The study was carried out at the 15% height level and extractions used successively dichloromethane, ethanol and water. At all sites, heartwood had significantly more extractives than sapwood, on average 3.8 and 2.4%, respectively. Most extractives consisted of ethanol soluble material (on average 52% of total extractives). Among the sites, there was a statistically significant difference in the content of extractives but the most important source of variation was the within-tree variation between sapwood and heartwood. Differences in the content of extractives were also observed among trees. A strong relation between extractives content and heartwood proportion was found. The potential loss of pulp yield and problems associated with accumulation of extractives are directly related to the heartwood proportion in the eucalypt stems. Forest management should take into account heartwood development and selection for minimising heartwood extractives.     

Characterization of Douglas-fir grown in Portugal: heartwood,sapwood, bark,ring width and taper

Douglas-fir (Pseudotsuga menziesii) is one of the best timber conifers providing long sawnwood components. Original from North America, it has been planted in Europe on approximately 550 thousand ha. Twenty Douglas-fir trees growing in two sites in Portugal were studied regarding ring analysis, heartwood, sapwood and bark development, and taper. The radial growth rate was 7.1 and 6.6 mm year⁻¹ at stem base for 45- and 50-year-old trees, respectively, in the two sites. Initial growth rate was slower, increasing until about 20 years and decreasing afterwards. Heartwood proportion represented on average 49% of the cross section in the lower part of the stem and decreased upwards. Heartwood formation was estimated to start at a cambial age of 8–9 years and increasing by 0.7–0.9 rings year⁻¹. Sapwood width was on average 75 mm at stem base, decreasing upwards. Bark was 26–27 mm thick at stem base, where it represented 15% of the cross-sectional area and decreased to 3–5 mm at the top. Stemwood and heartwood tapers were on average 15 mm m⁻¹ in the lower stem part and 21 and 18 mm m⁻¹, respectively, in the upper part. Douglas-fir showed a good potential for the mountain areas of Portugal, and under the silvicultural conditions of both stands the trees presented ring homogeneity, small conicity and low taper suitable for long wood components.

     

Chemical composition of earlywood and latewood in Norway spruce heartwood,sapwood and transition zone wood

This study focused on the distribution of wood components along a cross section of a spruce stem. Thin samples of earlywood and latewood were analysed by special micro-scale analytical techniques. Heartwood contained significantly more lignin and less cellulose than sapwood. The total content of hemicelluloses was the same along the radial direction, but the distribution of sugar units differed. The amounts of arabinoglucuronoxylan and pectins were larger in the heartwood. The transition zone between heartwood and sapwood had a specific composition, with less lignin and lipophilic extractives than heartwood and sapwood. For earlywood and latewood, significant differences were found in the distribution of sugar units in hemicelluloses. Latewood contained clearly more galactoglucomannan than earlywood, and conversely less pectins. The lipophilic extractives were also less concentrated in the latewood.Abbreviations EW or E earlywood - LW or L latewood - HW heartwood - SW sapwood - TZ transition zone wood - A.R. annual ring - AcBr Acetyl bromide - Ara arabinose - Xyl xylose - Gal galactose - Glc glucose - Man mannose - Rha rhamnose - GlcA glucuronic acid - MGlcA 4-O-methyl-glucuronic acid - GalA galacturonic acid - o.d. oven dry