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.     

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.     

Fungal decomposition of sapwood and heartwood of European aspen,Populus tremula L.

In contrast to the woodrotting fungi which can only attack Populus tremula after felling, Phellinus tremulae successfully invades the central part of the living tree. Possible causes for this difference were investigated. It was found that sterile wood samples were decomposed more rapidly by saprophytic fungi than by parasitic fungi. With both types of fungi, heartwood was more resistant to decay than sapwood. It is postulated that living sapwood inhibits infection by most woodrotting fungi, but that P. tremulae can successfully overcome this barrier. This postulate cannot be confirmed by experiments using sterilized wood samples.     

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?%.     

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).     

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.     

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.     

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.     

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.     

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.     

The occurrence of Phellinus tremulae (Bond.) Bond. and Borisov as a primary parasite in Populus tremula L.

Phellinus tremulae was inoculated on Populus tremula. It spread in the sapwood and inner wood of a living tree without other organisms being present. Plating experiments with wood meal from different parts of a sound tree showed that there were very few, if any, propagules of micro organisms in the sound wood, neither were there any organisms found in front of the decay of P. tremulae at the top of the decay column. These results support the theory that P. tremulae is a primary parasite in aspen. Decay fungi grown on fresh and autoclaved wood of aspen had very different growth rates. Most of the fungi tested grew best on autoclaved sapvvood. The difference in growth rate on fresh and autoclaved heartwood was lcss significant, suggesting that there is a barrier in the living wood which is destroyed by autoclaving. P. tremulae was thc only fungus that grew best on fresh inner wood.     

Ecology, history and silviculture of Scots pine (Pinus sylvestris L.) in western Norway - a literature review

Results from a literature review on pinewood ecology, silviculture,genetics, aspects of history and forest resources of Scots pine(Pinus sylvestris L.) in western Norway are presented. The pinewoodscover 40 per cent of the forested land, 0.31 million ha. Duringthe last 75 years, the area has increased by 17 per cent andthe growing stock has risen from 10 to 34 million m³. The impactof man in previous times was very marked, and has had a significantinfluence on the present forest conditions. The pronounced climaticgradients mixed with the topographic variation – fromthe coastal plains via the fjord systems to the high mountains– is reflected in rather steep gradients in the pine forestvegetation. Various floristic elements can be distinguished,from oceanic via the suboceanic in the outer islands to thethermophytic, boreonemoral and boreal elements in the innerfjord districts and valleys. The introduction of spruce (Piceaspp.) plantations on 10–15 per cent of former native pineforests has not negatively affected the bird fauna at the landscapescale. Although not particular species rich, the pine forestsharbour species usually not found in other forest types. Sofar, most work in the field of silviculture and forest ecologyin the pinewoods of West Norway has been in the form of casestudies. Implications of the results for forestry in the regionare briefly discussed.     

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.     

Understanding the role of the cytoskeleton in wood formation in angiosperm trees: hybrid aspen (Populus tremula x P. tremuloides) as the model species

The involvement of microfilaments and microtubules in the development of the radial and axial components of secondary xylem (wood) in hybrid aspen (Populus tremula L. x P. tremuloides Michx.) was studied by indirect immunofluorescent localization techniques. In addition to cambial cells, the differentiated cell types considered were early- and late-wood vessel elements, axial parenchyma, normal-wood fibers and gelatinous fibers, and contact and isolation ray cells. Microfilaments were rare in ray cambial cells, but were abundant and axially arranged in their derivatives once cell elongation had begun, and persisted in that orientation in mature ray cells. Microfilaments were axially arranged in fusiform cambial cells and persisted in that orientation in all xylem derivatives of those cells. Microtubules were randomly oriented in ray and fusiform cells of the cambial zone. Dense arrays of parallel-aligned microtubules were oriented near axially in the developing gelatinous fibers, but at a wide range of angles in normal-wood fibers. Ellipses of microfilaments were associated with pit development in fiber cells and isolation ray cells. Rings of co-localized microtubules and microfilaments were associated with developing inter-vessel bordered pits and vessel-contact ray cell contact pits, and, in the case of bordered pits, these rings decreased in diameter as the over-arching pit border increased in size. Although only microtubules were seen at the periphery of the perforation plate of vessel elements, a prominent meshwork of microfilaments overlaid the perforation plate itself. A consensus view of the roles of the cytoskeleton during wood formation in angiosperm trees is presented.     

The effects of mechanical site preparation and subsequent wildfire on trembling aspen (Populus tremuloides Michx.) regeneration in central Alberta,Canada

The objective of this study was to assess the regeneration response oftrembling aspen (Populus tremuloides Michx.) to differentmechanical site preparation (MSP) techniques commonly used in operationalforestry (disc trenching, drag scarifying and blading) and the specificmicrosites created by each treatment. This study was designed to measureregeneration after at least the first two growing seasons, however a largewildfire burned 80% of the study sites at the beginning of the second growingseason. Consequently, only limited second year data were presented, butregeneration from the first growing season following the fire was alsoassessed.Results indicated that microsites where the forest floor was disturbed and theparent root system was only lightly injured were more conducive to suckeringthan undisturbed microsites or where the root system was severely injured.Also,the fire disturbance after the first growing season resulted in increasedsuckering relative to the untreated controls in the first year. These resultssuggest that aspen sites with thick organic layers or vigorous competition fromother species can benefit from MSP when applied before the first growingseason.In addition, if first year suckering is inadequate, subsequent disturbancessuchas prescribed fire have the potential to improve suckering provided the parentroot system remains intact.     

An outbreak of Vapourer Moth (Orgyiaantiqua L.: Lepidoptera Lymantridae) on Sitka spruce (Picea sitchensis (Bong.) Carr.) in Central Scotland

Highly significant differences were observed in the intensityof defoliation on Sitka spruce of different heights at CarronValley Forest, Central Region. Most severe defoliation occurredon the smallest trees. Significant differences also occurredbetween degree of defoliation and foliar potassium and phosphorusconcentrations. Winter and summer rainfall patterns immediatelyprior to the outbreak showed higher than average winter andlower than average summer rainfall compared with the 30-yearmean. Experience with this pest suggests that it will continueto have local nuisance value.     

Browsing of sallow (Salix caprea L.) and rowan (Sorbus aucuparia L.) in the context of life history strategies: a literature review

Sallow (Salix caprea L.) and rowan (Sorbus aucuparia L.) constitute small proportions of the deciduous tree volume in Scandinavia, but are highly preferred winter forage for moose and red deer, which occur at historically high densities. Thus, a possible decline of these tree species has been indicated. Against this background, we have reviewed the life histories of relevance for browsing, as well as the basic biology and genetics of sallow and rowan. The species show similarities with respect to short lifespan, small size and sympodial growth pattern, which are risk factors in a browsing context. They also have high juvenile growth rate, important for growing quickly out of reach of browsers. Sallow depends strongly on disturbance for establishment and is more demanding with respect to soil and light conditions than rowan, possibly important for the substantially lower abundance of sallow on the Norwegian Forest Inventory plots. Similarly, the relative recruitment of small size classes of sallow is less than for rowan. Although recruitment is reported to be hampered in wintering areas with high moose or red deer densities, the inventory data, however, dating only back to 1994, do not suggest a general decrease in any of the species. Sallow and rowan saplings show low mortality in moose and deer dominated areas and the species can be characterised as rather resilient to browsing. Of more concern is that browsing can constrain the development of mature rowan and sallow trees locally, with possible consequences for associated epiphytic biodiversity.     

辐射松引种栽培综述

辐射松(PinusradiataD.Don)原产于加利弗尼亚海岸,被引种到世界各地,现在是南半球主要的人工林造林树种。1990年以来辐射松被引种到长江上游四川省阿坝州的干旱河谷地区,幼树的成活和早期生长表现良好。本文的主要内容包括:辐射松在原产地和引种地的生长情况,种源和遗传上的差异,环境限制因子和辐射松的气候生境,病虫害,引种不成功的经验教训以及辐射松在生态恢复中的应用。文中还简述了辐射松在阿坝干旱河谷地区的早期生长情况,列举了在长江上游干旱河谷地区引种辐射松可能遇到的问题以及所需开展的研究。图4表5参90。     

Productivity of mixed versus pure stands of oak (Quercus petraea (Matt.) Liebl. and Quercus robur L.) and European beech (Fagus sylvatica L.) along an ecological gradient

The mixture of beech (Fagus sylvatica L.) and oak (sessile oak, Quercus petraea (Matt.) Liebl., and pedunculate oak, Q. robur L.) is of considerable importance in Europe and will probably become even more important under climate change. Therefore, the performance of oak and beech in mixture was compared with the species’ growth in pure stands. Data from 37 long-term mixing experiments in Poland, Germany and Switzerland were pooled for analysis of mixing effects on stand productivity and possible interrelationships with mixing portions or site conditions. We found that on average, mixed stands of oak and beech exceeded biomass productivity in pure stands by 30 % or 1.7 t ha^?1 year^?1, as the growth of both species was benefitted by the mixture. However, that the interaction actually ranged from facilitation and overyielding on poor sites to underyielding on fertile sites triggered by competition. An empirically derived interaction model showed volume and dry mass growth changing in mixed stands from gains of 50 % to losses of 10 % depending on site conditions. It is concluded that the analysed mixture grows in accordance with the stress-gradient hypothesis and that our results suggest a site-specific relationship between species mixture and biomass productivity. As a consequence, an adequate species mix should result in increased productivity under steady state as well as climate change.