Analysis of EST-SSRs in silver birch(Betula pendula Roth.)

Simple sequence repeats(SSRs) defined as sequence repeat units between 1 and 6 bp occur abundantly in both coding and non-coding regions in eukaryotic genomes and these repeats can affect gene expression. In this study, ESTs(expressed sequence tags) of Betula pendula(silver birch) were analyzed for in silico mining of ESTSSRs, protein annotation, open reading frames(ORFs),designing primers, and identifying codon repetitions. In B.pendula, the frequency of ESTs containing SSRs was 7.8 %with an average of 1SSR/4. 78 kb of EST sequences. A total of 188 SSRs was identified by using MISA software and dinucleotide SSR motifs(65.9 %) were found to be the most abundant type of repeat motif followed by tri-(27.1 %),tetra-(4.8 %), and penta-(2.2 %) motifs. Based on ORF analysis, 175 of 178 sequences were predicted as ORFs and the most frequent SSRs were detected in 50 UTR(58.43 %),followed by in ORF(31.46 %) and in 30UTR(8.43 %). 102 of 178 ESTs were annotated as ribosomal protein, transport protein, membrane protein, carrier protein, binding protein,and transferase protein. For a total of 102 SSRs(57.3 %)with significant matches, a set of 102 primers(100 %) with forward and reverse strands was designed by using Primer 3 software. Serine(Ser, 19.6 %) was predominant in putative encoded amino acids and most of amino acids showed nonpolar(35.3 %) nature. Our data provide resources for B.pendula and can be useful for in silico comparative analyses of Betulaceae species, including SSR mining.     

Origin of the acetylated structures present in white birch (Betula pendula Roth) milled wood lignin

The occurrence and nature of acetate groups in the milled wood lignin (MWL) isolated from birch (Betula pendula Roth) has been addressed by spectroscopic (2D-NMR) and chemical degradative (derivatization followed by reductive cleavage, DFRC) methods. Considerable amounts of acetate groups were present in the MWL preparation. However, 2D-NMR analysis indicated that the lignin polymer is not extensively acetylated and that the major part of the acetate groups is attached to the xylan moieties present in the MWL preparation. Nevertheless, evidence of the presence of minor acetylation of the γ-carbon of the lignin side chain (<3% of both syringyl and guaiacyl lignin units) was provided by DFRC analysis.     

The effects of heavy thinning on stem quality and timber properties of silver birch (Betula pendula Roth)

A study was carried out into the effects of heavy thinning onstem quality and timber properties of silver birch (Betula pendulaRoth). Comparisons were made with lightly thinned and unthinnedbirch stands. Birch stands from the UK were also compared withsimilarly managed stands from Germany and Denmark. In general,trees from lightly thinned and unthinned stands had similarcharacteristics. Heavy thinning enhanced diameter incrementthrough greater ring width in comparison with the other treatments.The timber properties of basic density, grain angle and shrinkagecoefficients in the radial, tangential and longitudinal directions,however, were largely unaffected. More and bigger branches andgreater stem taper were found on trees from heavily thinnedstands, but these stems still met the current Finnish veneerbirch quality and dimension standards. Comparisons between birchstands, managed by heavy and light thinning, from the UK andGermany and Denmark revealed no meaningful differences in timberproperties. The results suggest that veneer dimension and qualitylogs could be produced in the UK in rotations as short as 40years on good sites if heavy thinning is adopted.     

Discoloration and associated fungi in stems of silver birch (Betula pendula Roth.) following logging damage

Study aim was to investigate discoloration and associated fungi in 50‐ to 70‐year‐old stems of Betula pendula damaged by logging. Investigated wounds (on 70 stems) were inflicted 4–18 years previously, and all of them had discoloration with mean radial spread (±SD) 2.9 ± 2.9 cm and longitudinal spread 50.3 ± 24.0 cm. Both wound area and length had a strong impact on a total length of discoloration (r = 0.665 and r = 0.745, respectively; p < 0.001), but had no influence on its spread beyond wound margins (r = ?0.035 and r = ?0.079). There was no correlation between the age of an injury and spread of the discoloration neither in vertical (r = ?0.110) nor in radial (r = ?0.280) direction. Average wound closure rate was 1.2 ± 0.5 cm year ^?1. The difference between discoloration length in stems with open and occluded injuries was insignificant (t‐test, p = 0.769). The principal fungi isolated from the wounds were ascomycetes (Epicoccum, Cadophora, Neonectria, Alternaria spp.) that were able to cause discoloration of wood with little effect on its mechanical properties and capable only for limited spread inside a stem of a living tree. In conclusion, results of the present study demonstrate that the amount of discoloured wood beyond logging wounds in B. pendula is mainly restricted to the damaged portion of a stem and that the associated loss of wood production remains low during about 10 years since the damage.     

Silver birch (Betula pendula Roth.) culture initiation in vitro and genotype determined differences in micropropagation

New Forests - Micropropagation has several advantages over conventional vegetative propagation methods, but it is limited by genotype responsiveness. We assessed the effect of age of the...     

Nitrogen and phosphorus leaching and uptake by container birch seedlings (Betula pendula Roth) grown in three different fertilizations

Leaching of nitrogen (N) and phosphorus (P) through container peatmedium and N and P uptake by seedlings were determined in commercial productionof birch grown with three different types of fertilization. Half of the amountof nutrients was premixed into the peat medium and the other half was appliedasliquid (=fertigated) in the treatment (PF) commonly used in Finland. In twoother treatments all nutrients, either totally (P-VN) or partly (P-N) inslow-release form, were premixed into the peat. Independently of the treatment,the largest proportion (60 to 80%) of total N leached during May and June.During two growing seasons, the amounts of N leached from PF treatments (9 to36kg ha^–1) were as large as the amounts of Nleachedfrom P-VN and P-N treatments (24 to 46 kg ha^–1).Due to fertigations between container trays, however, the total N load perhectare was greatest in the PF treatments. In the soil water 0.5meter beneaththe container area, the N concentration varied from 10 to 60 mgl^–1. The morphological and chemical properties of theseedlings did not differ greatly between treatments.     

Simultaneous drying and densification of silver birch (Betula pendula L.) veneers: analysis of morphology,thickness swelling,and density profile

In this study, birch (Betula pendula L.) veneers were simultaneously densified and dried using a contact drying method at pressures of 1.5 and 3 MPa at 130 °C and compared with veneer dried in a laboratory-scale convective type dryer. Compression rate, thickness swelling, and the density profiles of the veneers were investigated. Furthermore, the microstructure of densified veneers was studied by scanning electron microscopy (SEM). A maximum veneer compression rate of 9 % was achieved at a pressure of 3 MPa. Under these conditions, the veneers were, on average, densified from 504 to 574 kg m^?3 (approximately 14 %). After water soaking, full set-recovery—recovery to the initial thickness—occurred. However, the swelling rate was lower for the densified veneer. Density profiles measurements showed that densification occurs throughout the veneers. The SEM images showed that the surface of the densified veneers were smoother, whilst no cracks were detected due to densification. Densification seemed to occur in vessels. Typically, rays were bent when there was a vessel nearby.     

Genetic transformation of silver birch (Betula pendula) by particle bombardment

We used in vitro callus and shoot cultures as target material for genetic transformation of silver birch (Betula pendula Roth) by particle bombardment. Cultivation of in vitro shoot cultures before particle bombardment and a long selection period, combined with a high concentration of selective agent after bombardment, led to the production of transformed plantlets that were stable, and no escapes were found among the tree lines produced. Clonal variation in transformation efficiency was found in transient expression of the beta-glucuronidase gene in callus cultures and in plantlets transformed by stable integration of the ribulose-1,5-bisphosphate carboxylase/oxygenase small subunit (RbcS) and neomycin phosphotransferase (npt2) genes.     

Variations in phenology and growth of European white birch (Betula pendula) clones

Phenology can have a profound effect on growth and climatic adaptability of northern tree species. Although the large interannual variations in dates of bud burst and growth termination have been widely discussed, little is known about the genotypic and spatial variations in phenology and how these sources of variation are related to temporal variation. We measured bud burst of eight white birch (Betula pendula Roth) clones in two field experiments daily over 6 years, and determined the termination of growth for the same clones over 2 years. We also measured yearly height growth. We found considerable genetic variation in phenological characteristics among the birch clones. There was large interannual variation in the date of bud burst and especially in the termination of growth, indicating that, in addition to genetic effects, environmental factors have a strong influence on both bud burst and growth termination. Height growth was correlated with timing of growth termination, length of growth period and bud burst, but the relationships were weak and varied among years. We accurately predicted the date of bud burst from the temperature accumulation after January 1, and base temperatures between +2 and -1 degrees C. There was large clonal variation in the duration of bud burst. Interannual variation in bud burst may have important consequences for insect herbivory of birches.     

Twilight far-red treatment advances leaf bud burst of silver birch (Betula pendula)

Bud development of boreal trees in spring, once initiated, is driven by ambient air temperature, but the mechanism triggering bud development remains unclear. We determined if some aspect of the diurnal or seasonal light regime influences initiation of bud burst once the chilling requirement is met. We grew 3-year-old birch plantlets cloned from a mature tree of boreal origin in light conditions realistically simulating the lengthening days of spring at 60 degrees N. To emulate the reduction in red to far-red light (R:FR) ratio between daylight and twilight, one group of plantlets was subjected to reduced R:FR ratio in the morning and evening in addition to progressively lengthening days, whereas the other group was subjected to the same R:FR ratio throughout the day. The reduced R:FR ratio of twilight advanced bud burst by 4 days compared with the reference group (P = 0.04). To assess the interplay between the fulfillment of the chilling requirement and the subsequent response to warming, we fitted a thermal time model to the data with separate parameterizations for the starting dates of heat sum accumulation in each treatment. Least-squares fitting suggested that bud development started in light regimes corresponding to late March, almost two months after the chilling requirement for dormancy release was satisfied. Therefore, shortening night length or increasing day length, or both, appears to be the cue enabling bud development in spring, with twilight quality having an effect on the photoperiodic response. If twilight alone were the cue, the difference in bud burst dates between the experimental groups would have been greater than 4 days. The result gives experimental support for the use of thermal-time models in phenological modeling.     

Seasonal and within-stem variations of neutral lipids in silver birch (Betula pendula) wood

Neutral lipids were analyzed in stem wood of a 7-year-old clone and in five 35-70-year-old mature trees of silver birch (Betula pendula Roth). In young trees and in mature wood of old trees, the free fatty acid fraction comprised less than 5% of the concentration of triacylglycerols (TG). The concentration of free linoleic acid was lowest in March when the young trees were dormant and highest during midsummer and September. In mature trees, the TG concentration increased toward the pith, indicating that living parenchyma cells close to the pith have a large TG storage capacity. The TG concentration (mean 0.51 +/- 0.02% of wood dry mass) remained constant throughout the year in young trees, whereas the concentration of beta-sitosterol, the dominant free sterol (mean 82.5 +/- 0.4% of total free sterols), decreased during spring and early summer when the temperature gradually increased, and increased during autumn when the trees became dormant. In young trees, we detected a seasonal interconversion between the free and esterified forms of beta-sitosterol and campesterol, and within the steryl ester fraction between squalene and betulaprenol-7. The concentration of esterified sterols/isoprenoids was exceptionally high, especially in the inner regions of mature stem wood (mean 0.6 +/- 0.03% of wood dry mass). No heartwood formation was detected.     

Photosynthesis and canopy characteristics in genetically defined families of silver birch (Betula pendula)

Net photosynthetic rates (A) of leaves in upper and lower crown layers (A(upper) and A(lower)), leaf area index (LAI), mean tilt angle (MTA), several leaf characteristics, and volume growth were observed in fast- and slow-growing families of a 14-year-old full-sib and half-sib family progeny test of Betula pendula Roth. Each measure of net photosynthetic rate was calculated after correcting measured net photosynthesis for the effects of environmental variables. The differences in A(upper) and LAI among families were significant. The proportions of the total variance assigned to family for A(upper), A(lower) and LAI were 33.64, 28.93 and 54.99%, respectively. The mean A(upper) and LAI of the fast-growing families were significantly higher than those of the slow-growing families, whereas the mean A(lower) of the fast-growing families was significantly lower than that of the slow-growing families. There were also significant differences among families in leaf size, leaf shape, and the ratios leaf fresh weight/area and leaf dry weight/area. Between 27.55 and 54.55% of the total variance in these characteristics could be assigned to the family effect. Volume growth was positively correlated with A(upper) and LAI, but it was most strongly correlated with A(upper) x LAI.     

Isolation of genes up-regulated by copper in a copper-tolerant birch (Betula pendula) clone

Suppression subtractive hybridization (SSH) was used to isolate genes differentially expressed following exposure to copper (Cu) in a naturally selected Cu-tolerant birch (Betula pendula Roth.) clone originating from a disused lead/zinc smelter. Of the 352 cDNA fragments initially isolated, 108 were up-regulated by Cu, of which 55 showed over twofold induction by macroarray analysis. Searches against protein databases (Blastx) and sequence analysis provided the tentative identity of 21 genes. Three fragments lacked homology to any sequences in the databases. Most of the identified genes are involved in cellular transport, regulation or cell rescue and defense. Several genes have not previously been reported to be up-regulated by Cu, e.g., plasma intrinsic protein 2, glutamine synthetase and multi-drug resistance-associated protein (MRP4). The expression of MRP4, a vacuolar sorting receptor-like protein and an unidentified gene was studied in more detail by quantitative real-time PCR. These genes showed stronger up-regulation by Cu in the roots and shoots of the Cu-tolerant birch clone compared with a less tolerant clone. Clear clonal differences in gene expression were observed, e.g., for the regulator of chromosome condensation family protein, DnaJ protein homolog, vacuolar sorting receptor-like protein and MRP4. These findings contribute to our understanding of Cu tolerance in birch, a pioneer plant in metal-contaminated soils.     

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.     

Long-term dynamics of leaf and root decomposition and nitrogen release in a grey alder (Alnus incana (L.) Moench) and silver birch (Betula pendula Roth.) stands

The decomposition of the leaf litter, fine roots (d?<?2?mm) and coarser roots (2?≤?d?<?5?mm) of grey alder and silver birch, as well as of α-cellulose sheets using the litterbag method was studied in two experimental stands on Podzoluvisol soils in Southern Estonia. For both tree species, the coarser roots decomposed faster than the fine roots, (p?<?.05), tree species did not affect the decomposition rate of the roots (p?>?.5). The nitrogen (N) input to soil from aboveground litter was multiple times higher than the N flux from roots. The remaining relative ash-free mass of the leaves of grey alder and silver birch after three and a half years was similar. After 11 years the remaining relative ash-free mass of the fine and coarser roots of grey alder still accounted for around 10% of the initial value. For silver birch the remaining value was around 20% after 9 years. The litterbag method to underestimates in fertile soils the decomposition of organic matter and thus did not reflect the actual dynamics of decomposition.     

Effects of acid rain treatments on the birch rust,Melampsoridium betulinum and birch,Betula pendula

Effects of acidic rain on birch rust and on the growth of Betula pendula seedlings were studied by irrigating with acidic water at pH 3, 4 and 5.6. The seedlings were inoculated with uredospores of Melampsoridium betulinum and the development of rust and the severity of rust infection was examined. The increasing acidity of the irrigation water did not affect on the development of the rust, but the nitrogen in the acidified water resulted in an extension of seedling growth, and hence the leaves became senescent at a later date in autumn and the rust grew for a longer time.