Interactive effects of elevated ozone and temperature on carbon allocation of silver birch (Betula pendula) genotypes in an open-air field exposure

In the present experiment, the single and combined effects of elevated temperature and ozone (O(3)) on four silver birch genotypes (gt12, gt14, gt15 and gt25) were studied in an open-air field exposure design. Above- and below-ground biomass accumulation, stem growth and soil respiration were measured in 2008. In addition, a (13)C-labelling experiment was conducted with gt15 trees. After the second exposure season, elevated temperature increased silver birch above- and below-ground growth and soil respiration rates. However, some of these variables showed that the temperature effect was modified by tree genotype and prevailing O(3) level. For instance, in gt14 soil respiration was increased in elevated temperature alone (T) and in elevated O(3) and elevated temperature in combination (O(3) + T) treatments, but in other genotypes O(3) either partly (gt12) or totally nullified (gt25) temperature effects on soil respiration, or acted synergistically with temperature (gt15). Before leaf abscission, all genotypes had the largest leaf biomass in T and O(3) + T treatments, whereas at the end of the season temperature effects on leaf biomass depended on the prevailing O(3) level. Temperature increase thus delayed and O(3) accelerated leaf senescence, and in combination treatment O(3) reduced the temperature effect. Photosynthetic : non-photosynthetic tissue ratios (P : nP ratios) showed that elevated temperature increased foliage biomass relative to woody mass, particularly in gt14 and gt12, whereas O(3) and O(3) + T decreased it most clearly in gt25. O(3)-caused stem growth reductions were clearest in the fastest-growing gt14 and gt25, whereas mycorrhizal root growth and sporocarp production increased under O(3) in all genotypes. A labelling experiment showed that temperature increased tree total biomass and hence (13)C fixation in the foliage and roots and also label return was highest under elevated temperature. Ozone seemed to change tree (13)C allocation, as it decreased foliar (13)C excess amount, simultaneously increasing (13)C excess obtained from the soil. The present results suggest that warming has potential to increase silver birch growth and hence carbon (C) accumulation in tree biomass, but the final magnitude of this C sink strength is partly counteracted by temperature-induced increase in soil respiration rates and simultaneous O(3) stress. Silver birch populations' response to climate change will also largely depend on their genotype composition.     

Effects of elevated concentrations of ozone and carbon dioxide on the electrical impedance of leaves of silver birch (Betula pendula) clones

Effects of elevated concentrations of tropospheric ozone ([O3]) and carbon dioxide ([CO2]) on leaves of two silver birch (Betula pendula Roth) clones were monitored for three growing seasons (1998, 1999, 2000) by means of electrical impedance spectroscopy (EIS). The field trial with open-top chambers (OTCs) was conducted on two clones (Clone 4 and Clone 80) with five treatments and four independent replicates. Treatments were: (1) outside control, (2) chamber control, (3) 2x ambient [O3], (4) 2x ambient [CO2] and (5) 2x ambient [CO2] + 2x ambient [O3]. Fumigations started in 1999 and continued in 2000. Measurements were made in 1998 before the fumigations and thereafter EIS was carried out four times in each season. The impedance spectra of about 10 leaves from each tree at each time were measured at 42 frequencies between 80 and 1 MHz. Leaf spectra were modeled by a distributed circuit element model (DCE) (one DCE in series with a resistor), which yields the extracellular and intracellular resistances, the relaxation time and the distribution coefficient of the relaxation time. The EIS properties of the leaves changed significantly during the growing season when new leaves were expanding. The clones differed in their EIS properties. Clone 4 had a significantly higher extracellular resistance and distribution coefficient than Clone 80. The clones responded similarly to the fumigation treatments. Differences between treatments emerged especially during the second fumigation season in 2000. Elevated [O3] reduced both the relaxation time and the extracellular resistance, indicating cell membrane damage. Elevated [CO2] increased the intracellular resistance, indicating changes in symplastic composition. The biological interpretation of the EIS parameters in birch leaves is discussed.     

Dependence of dormancy release on temperature in different origins of Pinus sylvestris and Betula pendula seedlings

The chilling requirement of rest completion and the high temperature requirement of growth initiation were determined in three origins of silver birch (Betula pendula Roth.) seedlings and five origins of Scots pine (Pinus sylvestris L.) seedlings. In both the pine and birch the chilling requirement was highest in maritime Scottish origins and lowest in the most continental Finnish and Russian origins. The requirement for southern mountainous Spanish and Bulgarian pine origins was in between. In terms of the high temperature requirement, there were no clear differences between origins. These results suggest that owing to their high chilling requirement, which prevents the beginning of growth and the loss of frost hardiness during the frost‐exposed season, origins from a maritime climate could be the most tolerant under climatic warming.     

三倍体欧洲山杨组培技术的研究

通过对三倍体欧洲山杨组培技术的研究试验,采用增殖与生根同步进行的方法进行快繁,其年增殖次数可达8次,平均每次增殖培数为4．4倍,可以达到工厂化生产的指标要求。培养出的组培苗质量好,移栽成活率高,而且可以降低苗木的成本。     

Biochemical acclimation patterns of Betula pendula and Pinus sylvestris seedlings to elevated carbon dioxide concentrations

Acclimation of photosynthesis to increasing atmospheric carbon dioxide concentration ([CO2]; 350 to 2,000 micromol mol-1) was followed in silver birch (Betula pendula Roth.) and Scots pine (Pinus sylvestris L.) seedlings for two years. Chlorophyll fluorescence and concentrations of Rubisco, chlorophyll, total soluble protein and nitrogen were monitored together with steady-state gas exchange at three CO2 concentrations (ambient [CO2] (345 +/- 20 micromol mol-1), the growth [CO2] and 1950 +/- 55 micromol mol-1). Rubisco and chlorophyll concentrations decreased in birch and Scots pine with increasing growth [CO2]. A nonlinear response was recorded for Rubisco and chlorophyll concentrations in birch, which was correlated with a significant decrease in specific leaf area. Nitrogen concentration decreased in birch leaves, but was unchanged in Scots pine needles. The species differed substantially in their steady-state CO2 exchange response to increasing growth [CO2]. The principal effect in birch was a significant nonlinear decrease in the steady-state gas exchange rate at the ambient [CO2], whereas in Scots pine the main effect was a significant increase in the steady-state gas exchange rate at the growth [CO2].     

Differential responses of silver birch (Betula pendula) ecotypes to short-day photoperiod and low temperature

We investigated interrelations of dormancy and freezing tolerance and the role of endogenous abscisic acid (ABA) in the development of silver birch (Betula pendula Roth) ecotypes in controlled environments. Short-day treatment induced growth cessation, bud set and dormancy development, as well as initiation of cold acclimation and an increase in freezing tolerance. Subsequent low temperature and short days (12-h photoperiod) resulted in a significant increase in freezing tolerance, whereas bud dormancy was gradually released. The concentration of ABA increased in response to short days and then remained high, but ABA concentrations fluctuated irregularly when the dormant plants were subsequently exposed to low temperature during short days. Although there was a parallel development of freezing tolerance and bud dormancy in response to short days, subsequent exposure to low temperature had opposite effects on these processes, enhancing freezing tolerance and releasing dormancy. Compared with the southern ecotype, the northern ecotype was more responsive to short days and low temperature, exhibiting earlier initiation of cold acclimation, growth cessation and an increase in ABA concentrations in short days, and higher freezing tolerance, faster dormancy release and greater alteration in ABA concentrations when subsequently exposed to low temperature during short days. The rates and extent of the increases in ABA concentration may be related to increases in freezing tolerance and dormancy development during short days, whereas the extent of the fluctuations in ABA concentration may play an important role in enhancing freezing tolerance and releasing dormancy during a subsequent exposure to low temperature during short days.     

Micropropagation of a mature colchicine-polyploid and irradiation-mutant of Betula pendula Roth

Tetraploid plantlets were regenerated from cultured apical and axillary buds of a 23-year-old colchicine-polyploid and irradiation-mutant Betula pendula Roth tree. Bud explants were grown on modified Murashige and Skoog medium supplemented with 2.0 mg l(-1) benzylaminopurine (BAP) and 0.01 mg l(-1) 1-naphthaleneacetic acid (NAA). The medium allowed both induction of adventitious buds and development of shoots. The cut ends of new shoots produced new buds and shoots during a 4-week culture period. The micropropagated shoots were rooted on modified Murashige and Skoog medium containing 0.1 mg l(-1) NAA as the sole growth regulator. Plantlets were transferred to a peat/soil mixture (1:1) in the greenhouse, acclimated and then transplanted to a cold frame. The regenerated plantlets had a tetraploid chromosome set (4n = 56) and an altered leaf morphology typical of colchicine-polyploid birches. The leaves were hypertrophied and asymmetrical, with curly leaf margins. The mutant nature of the parent tree was also evident in the light-green color of the leaves of the plantlets.     

欧洲白桦早期生长性状表现与优良家系选择

在测定分析欧洲白桦13个家系子代生长节律、适应性及早期性状表现的基础上,系统研究生长性状变异规律,并结合优树、子代测定林和子代苗期的生长表现,进一步开展优良家系选择。结果表明：各家系子代生长高峰期均集中在7月上旬至8月初,树高和地径性状均呈正态分布,且家系间和家系内差异显著,家系间抗寒适应性差异较小。综合评价各项指标,筛选出5个生长稳定、遗传分化较小的优良家系,即：6#、9#、1#、10#及13#家系。     

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.     

Root Egress and Field Performance of Actively Growing Betula pendula Container Seedlings

The objective of this study was to evaluate the possibility of enlarging the planting window of container-grown silver birch seedlings from spring and autumn to summer by planting young, actively growing seedlings. For over 3 yrs the study investigated the growth, survival and damage of silver birch seedlings grown in containers and planted at different times during the growing season on 18 sites in fields and forest sites in central Finland. In a pot experiment in the greenhouse, the root-egress ability of seedlings planted at different times during the growing season was monitored. Root egress of seedlings was rapid from the beginning of July to the middle of August. Actively growing seedlings planted in summer on a site suitable for silver birch grew and survived at least as well as seedlings planted in spring during the dormant stage. The risk of drought stress increased when seedlings were planted on easily drying, coarse sandy soils. The risk was also high in soils with high levels of fine soil fractions that become hard when dry and when the dry period continued for several weeks before planting. In conclusion, it is possible to enlarge the planting window of birch seedlings to July and the beginning of August by using actively growing container seedlings.     

Growth and nutrition of Betula pendula at different relative supply rates of zinc

Small birch plants (Betula pendula Roth) were cultivated in a hydroponic spray solution where the relative growth rate (R(G); day(-1)) was controlled by the relative supply rate of zinc (R(Zn); day(-1)). After an adjustment phase to steady-state growth, R(G) equaled R(Zn). The R(Zn) treatments were 0.05, 0.125 and 0.20 day(-1) with free access to all other nutrients. In an additional treatment, there was free access to all nutrients, including zinc (FA treatment). The pH of the nutrient solution was approximately 4.5 and conductivity was 100 &mgr;S cm(-1). The duration of each treatment depended on R(Zn) and ranged from 4 (FA treatment) to 10 weeks (at R(Zn) = 0.05 day(-1)). The plants showed persistent and typical zinc-deficiency symptoms at steady-state growth when R(G) was 0.05 and 0.125 day(-1), whereas there were few symptoms when R(G) was 0.2 day(-1). The Zn concentration of the plants ranged from 8 (at R(Zn) = 0.05 day(-1)) to 21 &mgr;g g(DM) (-1) (DM = dry mass) (at R(Zn) = 0.2 day(-1)) and was approximately 42 &mgr;g g(DM) (-1) in the FA treatment. Uptake rates of Zn, calculated per root growth rate (&mgr;mol g(DM, root) (-1)), were about 2.8 times higher at R(Zn) = 0.20 day(-1) than at R(Zn)= 0.05 day(-1). The root and stem biomass fractions were approximately constant at all supply rates of Zn, whereas the leaf biomass fraction tended to increase with increasing supply rate of Zn. Net assimilation rate was constant from FA to an R(Zn) of 0.125 day(-1) but decreased by a factor of about 2 at R(Zn) = 0.05 day(-1). Leaf area ratio and specific leaf area were smaller at low supply rates of zinc than at high supply rates.     

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.     

Growth responses and related biochemical and ultrastructural changes of the photosynthetic apparatus in birch (Betula pendula) saplings exposed to low concentrations of ozone

Saplings of ozone-sensitive and ozone-tolerant birch (Betula pendula Roth.), clones B and C, respectively, were exposed to ozone concentrations that were 1.7-fold higher than ambient for one growing season under open-field conditions. Ambient air was used as the control treatment. In the ozone-sensitive clone B, there was an initial stimulation of leaf area growth in response to the ozone treatment, but further ozone exposure caused reductions in leaf and stem biomass growth, Rubisco and chlorophyll a contents, net photosynthesis, water use efficiency and chloroplast size. It also caused an alteration in chloroplast shape and injury to thylakoid membranes. In the ozone-tolerant clone C, ozone fumigation did not affect growth rate, and there were no consistent changes in chlorophyll content, photosynthesis or water use efficiency. There were also fewer ultrastructural abnormalities in the chloroplasts of clone C than of clone B. Based on the observed biochemical, physiological and structural changes in chloroplasts of clone B in response to low concentrations of ozone, we conclude that the increasing concentration of tropospheric ozone represents a risk to natural birch populations.     

Improvement of Betula pendula by clonal and progeny testing of phenotypically selected trees

Abstract

Field tests, each including 39–83 Swedish plus-tree clones of silver birch (Betula pendula Roth) planted on 10 different sites in the southern part of Sweden, were evaluated at up to 10 years of age. The main conclusions can be summarized as follows. Growth traits were mostly under strong genetic influence, and showed substantial genetic variation and high potential genetic gain. Survival did not seem to be a trait of genetic importance. The genetic correlation between growth and growth cessation was weak, suggesting that it should be possible to find clones combining high growth and early autumn growth. Non-significant transfer effects for clones of different origin and weak G×E interaction were found, indicating that southern Sweden can be treated as a single utilization zone and that few sites are needed for genetic tests. Strong genetic age×age correlations were detected, suggesting that short test periods could be used.     

The impact of tropospheric ozone on landscape-level merchantable biomass and ecosystem carbon in Canadian forests

Studies have shown that tropospheric ozone (O₃) impacts trees in various ways, including growth reductions. To date, the landscape-level response of Canadian forests carbon (C) to O₃ exposure has not been quantified. We used a modified version of the Carbon Budget Model of the Canadian Forest Sector and data from Aspen FACE to quantify the landscape-level impacts of different O₃ exposure modelling experiments. The main strengths of our approach consisted of using the most complete empirical data available to estimate the amount and location of forest C across Canada, as well as explicitly simulating the consequences of fire, insect, and harvest disturbances on forest C dynamics. These disturbances lead to younger forests and, considering trees sensitivity to O₃ exposure to decrease with age, thus result in higher landscape-level modelled impacts for the same O₃ levels. Despite various sources of uncertainty, our results indicate that even under a modelling experiment where O₃ increases continuously over four decades, the landscape-level impacts on the merchantable biomass and ecosystem C remain limited. Our results also suggest that the current direct impacts of O₃ on Canadian forests are likely below detection at the landscape level.     

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.     

Stem lesions on Betula pendula seedlings in Finnish forest nurseries and the pathogenicity of Phytophthora cactorum

Phytophthora cactorum was isolated from necrotic stem lesions on 20–80% of diseased Betula pendula seedlings sampled from three nurseries in Finland. Fusarium avenaceum and Godronia sp. were also found in all nurseries. Inoculation of P. cactorum on B. pendula seedlings caused necrotic lesions and the spread of these lesions around the stem was detrimental. Neither the timing of N application nor two different PK-levels applied to the seedlings the previous year significantly affected the size of lesions caused by P. cactorum.     

欧洲山杨叶片衰老与光周期候选基因SNPs的关联分析

利用自然更新的欧洲山杨群体,以光周期途径基因为候选基因,通过连锁不平衡作图,对候选基因中能够引起氨基酸改变的单核苷酸多态性(SNP)与欧洲山杨秋季叶片衰老相关指标进行关联分析.结果表明:光敏色素A中第291个核苷酸A和G的转换,导致了3个基因型叶片衰老指数有显著差异(P=0.018,经Bonferroni修正后,P=0.04);光周期候选基因中的SNP、叶片衰老持续时间基因型值间无显著差异,SNP与叶片衰老持续时间之间不存在线性相关;同样是光敏色素A中位于第291个核苷酸位点的3种基因型,其叶片衰老起始时间的基因型值差异显著,但是经Bonferroni修正后的显著水平P>0.05,差异又不显著,反映了欧洲山杨秋季叶片衰老起始时间的复杂性.