Paper birch (Betula papyrifera) shoot selection by moose (Alces alces) following a forest-cleaning experiment

Abstract

In order to maximize food intake per harvesting effort and minimize energy expenditures required to move between feeding patches in nature, herbivores such as moose (Alces alces L.) generally select large plant shoots when browsing in winter. To determine moose preferences for shoots of different morphologies, an experiment was conducted in northern British Columbia in which shoots from birches cut at different times of the growing season were fed in 2 consecutive years to eight human-habituated moose in cafeteria-style feeding trials. The results indicate that moose preferred smaller winter shoots of birches regardless of when the parent plant was cut and also appeared to reject larger shoots containing sylleptic branches. It is argued that the preferences for smaller shoots by moose detected in these trials should be observable under natural conditions, but are generally only supported by literature from some parts of Scandinavia. The findings underscore the importance that factors such as mouth filling per harvesting effort, snow depth and consistency, predators and browse patch distribution must have on foraging decisions made by moose while browsing in the wild. Implications of the findings include the significance of cutting time on the size of shoots produced by birch after cutting, how this affects moose browsing birch and, subsequently, how managers can theoretically use cutting time as a tool in forest cleaning operations to direct the foraging efforts of moose towards or away from forest plantations.     

Effects of root medium pH on water transport in paper birch (Betula papyrifera) seedlings in relation to root temperature and abscisic acid treatments

We investigated the effects of root medium pH on water transport in whole-plant and detached roots of paper birch (Betula papyrifera Marsh.). Exposure of seedling roots to pH 4 and 8 significantly decreased root hydraulic conductivity (Lp) and stomatal conductance (gs), compared with pH 6. When roots of solution-culture-grown (pH 6) seedlings were transferred to pH 4 or 8, their steady-state water flow (Qv) declined within minutes, followed by a decline in gs. The root oxygen uptake rates were not significantly affected by the pH treatments. Treatment of roots with mercuric chloride resulted in a large decrease in Qv at pH 6; the extent of this decrease was similar to that brought about by pH 4 and 8. Lowering root temperature from 21 to 4 degrees C decreased Qv irrespective of medium pH. Low root temperatures did not offset the effects of medium pH 4 on Qv and the roots in this treatment had a high activation energy for water flow. Conversely, roots exposed to pH 8 had a low activation energy, similar to that at pH 6. When 2 micro M abscisic acid, (+/-)-cis-trans-ABA, was added to the root medium, Qv increased in roots that were incubated at pH 6. It also increased slightly in roots incubated at pH 4, but not at pH 8. The increase at pH 4 and 6 was temperature-dependent, occurring at 21 degrees C, but not 4 degrees C. We suggest that the pH treatments are responsible for altering root water flow properties through their effects on the activity of water channels. These results support the concept that ABA effects on water channels are modulated by other, possibly metabolic- and pH-dependent factors.     

Properties of white birch (Betula papyrifera) outer bark particleboards with reinforcement of coarse wood particles in the core layer

? This study proposes substituting traditional raw materials in the surface layers of wood particle-boards with the water resistant white birch (Betula papyrifera) outer bark particles, which can help improve the dimensional stability of manufactured mixed particleboards, thereby alleviating shortages of raw material in a cost-efficient manner.

? Mixed particleboards were fabricated in the laboratory using untreated or alkali treated white birch outer bark particles as substitute material. These particles were resinated successively with three percentages of phenol-formaldehyde resin. Overall, the results of this study clearly demonstrate that the panels could be manufactured using up to 45% of the proposed substitute material and still maintain the required mechanical and physical properties.

? Alkali treatment was used to remove natural wax from bark particles surface which hinders resin adhesion. This treatment negatively affected mechanical and physical properties of finished panels and the variation of phenol-formaldehyde resin percentage in the bark particles significantly affected only their hardness.

? Panel with untreated bark particles in the surface layers, resinated with 5% phenol-formaldehyde resin was selected as the best with the help of a statistical analysis carried out in a factorial complete block design, especially from the dimensional stability criterion.

     

Effects of winter temperatures on two birch (Betula) species

In Massachusetts, low winter temperatures delay the onset of flowering in black birch (Betula lenta L.), but not in gray birch (B. populifolia Marsh.). During the winter of 2006, male inflorescences and twigs of black birch had higher water contents than those of gray birch, and the inflorescences of black birch experienced greater frost kill than those of gray birch. Vessels diameters were greater in black than in gray birch, a difference associated with a higher incidence of winter xylem embolism, as indicated by reduced xylem hydraulic conductance. In both species, recovery of hydraulic conductance in twigs that survived the winter coincided with the development of root pressure. Frost kill to male inflorescences or associated damage to plant tissues may account for the difference between species in the effect of winter temperature on the time of first flowering. In a comparison of 24 birch species, sensitivity of the first flowering date to temperature was also correlated with water content in male inflorescences.     

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.     

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.     

Exploring life growth patterns in birch (Betula pendula)

The conversion of homogeneous plantations of conifers to mixed or broadleaved forests is part of sustainable forest management strategies in Western Europe. For the conversion of pine plantations on sandy soils, birch seems a convenient choice. Yet, thus far, little is known about the growth of mature birches outside Northern Europe. We sampled 32 birch trees in forests throughout the sandy ecoregion in northern Belgium at sites that varied in soil and local stand conditions. Half of the birches showed the characteristic pattern in growth (expressed as basal area increment) with a sharply increasing growth till around the age of 25 years followed by a steep decrease in growth. The growth was strongly related to tree age and crown dimensions, only weakly related to soil conditions, and not related to the local stand conditions.     

Photosynthesis,water and nitrogen use efficiencies of four paper birch (Betula papyrifera) populations grown under different soil moisture and nutrient regimes

A greenhouse experiment was conducted to determine how four populations of paper birch (Betula papyrifera Marsh.), adapted to particular temperature, water and photoperiod regimes responded physiologically to different soil water and nutrient regimes. Seedlings of each of the four populations (Eaglet, LeeCreek, Porcupine and Skeena) from British Columbia were planted in pots and subjected to high and low water and nitrogen regimes for 4 months. Net photosynthesis (A), stomatal conductance (g_s), water use efficiency (WUE) and nitrogen use efficiency (NUE) were measured and compared among treatments. Trade-offs between WUE and NUE were examined. Soil moisture was the major factor affecting A for all the populations. All populations showed relatively high A in the high-water and high-nitrogen (HWHN) treatment, ranging from 8.5 to 9.9 μmol CO₂ m⁻² s⁻¹. Photosynthesis of trees in the low-water and high-nitrogen (LWHN) treatment was about 45% of that in the HWHN treatment. There was a linear increase in A as g_s increased in all populations. A decreased with increasing vapor pressure deficit (VPD). The populations demonstrated significantly different relationships between A and g_s, and VPD, with the LeeCreek population being the most conservative. This may provide a physiological basis for provenance selection and seed transfer. Eaglet and LeeCreek populations had lower WUE at the same level of NUE compared with Porcupine and Skeena populations. Porcupine and Skeena populations may be more suitable for moist sites due to their greater capacity to sequester water quickly, while the LeeCreek and Eaglet populations may be more suitable for drier interior sites where long drought periods can occur.     

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.     

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.     

Influence of light quality on the germination of Betula papyrifera seeds

The influence of light quality on germination of unstratified birch seeds (Betula papyrifera) was investigated in a greenhouse study. Different light regimes were created with five plastic filters of different colour in combination with basic illumination. The filters were adjusted to equal levels of photosynthetically active radiation. The difference in light quality in terms of R : Fr ratio (red : far‐red light ratio) varied from 0.23 to 1.03. Germination was recorded 7, 10, 14, 21, 25, 31 and 36 days after sowing. Response to light quality was significant from 25 days after sowing (p < 0.043), and it was found that differences increased with time. Filters producing light regimes with low R : Fr ratios were found to reduce and delay germination of birch seeds. Germination after 36 days varied from 55% for the control treatment to 11% for the lowest R : Fr ratio. The relationship between seed germination percentage and R : FR ratio was found to be significant (p < 0.017) with an R‐sq value of 88.6%.     

Elevated atmospheric CO(2) concentration changes ectomycorrhizal morphotype assemblages in Betula papyrifera

Ectomycorrhizae are extremely diverse, with different species of fungi having very different physiologies and morphologies that, in turn, confer a range of different benefits to the host plant. To test the hypothesis that elevated CO(2) leads to changes in the assemblage of ectomycorrhizae associated with trees, we examined the number and frequency of ectomycorrhizal morphotypes colonizing roots of Betula papyrifera Marsh. saplings grown at an ambient or elevated (700 ppm) atmospheric CO(2) concentration for 24 weeks. Elevated CO(2) resulted in significant changes in the composition of the ectomycorrhizal assemblage toward morphotypes with a higher incidence of emanating hyphae and rhizomorphs. We conclude that B. papyrifera saplings will be able to support a more costly mycorrhization in future elevated-CO(2) atmospheres.     

Micro-variations in yellow birch (Betula alleghaniensis) growth conditions after patch scarification

The abundance of poor quality yellow birch stands resulting from diameter limit cuts poses important regeneration challenges since seed production is uncertain, adequate seedbeds are rare and competition very severe. The present study aims at evaluating variations in seedling growth, temperature, soil humidity and nutrient availability for the whole range of seedbeds created by a patch scarification treatment. In three poor quality stands, soil temperature and humidity, nutrient availability and seedling morphology were monitored on undisturbed forest floor, at different places in 2 m wide scarified patches, and on mounds formed by the displaced material. Even though the study associated some variations in temperature and humidity to different microsite positions, the effect on nutrient availability was more pronounced. Hence, the study found better nutrient availability on mounds in comparison with patches as well as at the border of patches. Soil nutrient availability as measured with in situ ion exchange resin declined very rapidly in the interior of scarified patch (within 5 cm for the border) suggesting that small scarified patches combined with control of vegetation competition should provide a better environment for seedling growth. Growth parameters of yellow birch seedlings were not correlated to measured soil parameters suggesting that there were important effects of uncontrolled factors.     

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.     

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.     

Discolouration of birch wood: analysis of extractives from discoloured surface of vacuum-dried European white birch (Betula pubescens) board

A discolouration that appeared on the surfaces of a European white birch (Betula pubescens) board during vacuum drying was studied by means of colour measurements (CIEL*a*b*), elemental analysis and the analysis of extractives. The discoloured surface layer of the dried board contained substantially more methanol-soluble extractives than did the light-coloured part (50.7 mg g⁻¹ vs. 26.7 mg g⁻¹ dry wood), and the colour difference (ΔE* _ab 20.0) between the two extracts was notable. Characterization by means of GC and ¹³C NMR spectroscopy showed that the extracts contain sugars (mainly glucose and fructose), low-molecular-weight phenolic compounds, proanthocyanidins, Brauns’ lignin and fatty acid esters. Concentrations of sugars, low-molecular-weight phenols, proanthocyanidins and Brauns’ lignin were higher in the discoloured surface layer than in the light-coloured part. The yellowness of the surface layer was associated with the accumulation of low-molecular-weight phenolic extractives, and the redness with Brauns’ lignin and possibly proanthocyanidins.     

Effect of ABA on freezing resistance of Betula papyrifera and Alnus incana woody plant cell suspensions

Treatment of birch (Betula papyrifera Marsh) and alder (Alnus incana (L.) Moench) cell suspension cultures with ABA increased the freezing resistance of the cells. After 7 days of treatment with 10(-5) M ABA, birch cells grown at 23 and 4 degrees C attained an LT(50) of -16.9 and -14.1 degrees C, respectively, whereas control cells had an LT(50) of -9.1 degrees C. In alder cell suspensions, treatment with 10(-5) M ABA at 23 degrees C induced a small increase in freezing resistance from -7.3 to -10.8 degrees C. Exposure to 4 degrees C alone did not induce a significant increase in hardiness in birch cell suspensions. Addition of 10(-5) M ABA to the medium inhibited fresh weight increase over 10 days of 3-g inocula of birch and alder by 70 and 52%, respectively. With the same concentration of ABA in the medium we found different intracellular ABA concentrations in 3- and 6-g inocula. We conclude that the concentration of ABA in the medium does not reflect the intracellular concentration of tissue cultures, and that cultural conditions may influence ABA accumulation by cell cultures.     

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.