Silver Birch Genotypes Research Articles

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

Micropropagation has several advantages over conventional vegetative propagation methods, but it is limited by genotype responsiveness. We assessed the effect of age of the mother-tree and the time of explant collection on culture initiation, as well as the multiplication ability and effect of different nutrient media and plant growth regulators on silver birch genotypes. Explants collected from 1‐year‐old trees (66%) and explants collected in spring (64–67%) developed a significantly (both p < 0.001) higher proportion of shoots than those from 15‐year‐old trees (39%) and those collected in mid-summer (31%) and autumn (29%), respectively. In a stabilised culture, the length of the main shoot varied from 1.3 to 7.8 cm between genotypes, and the multiplication rate ranged from 1.0 to 6.8 shoots per explant. Hyperhydrated shoots were present in 17 out of 50 clones, and, among the clones, ranged from 14 to 50%. Cultures on the Murashige and Skoog basal medium had a higher multiplication rate than cultures on a Woody Plant Medium, and the application of zeatin provided better results than 6‐benzylaminopurine. The difference between cytokinin types was 11–29% for the multiplication rate and 21–29% for the length of the main stem. The highest multiplication rate was obtained using a zeatin concentration of 0.5 mg L−1. However, better shoot growth and proliferation had a significant positive relation to shoot hyperhydration (all p < 0.001). Therefore, a medium with an optimal balance between the multiplication rate and the number of hyperhydrated shoots should be carefully selected.

Local adaptation of phenotypic stem traits distinguishes two provenance regions of silver birch in Latvia

Populations of tree species with a wide geographic range, such as silver birch ( Roth), show genetic specialization to native environments, while maintaining high phenotypical plasticity. Accordingly, assessment of local specialization is essential for adaptive management. The aim of the study was to detect geographic patterns of local adaptation of growth and stem quality based on two open-pollinated progeny trials in Latvia testing local material. Two provenance regions differing by continentality were distinguished, which also differed in genetic control of growth traits, likely originating from the post-glacial recolonization of vegetation and subsequent natural adaptation. Heritability of the traits was estimated for each of the distinguished regions, indicating differing patterns of genetic adaptation and potential for future selection. Trees from the more continental inland showed superior growth and possessed higher heritability. The coastal provenance region showed slower growth and intermediate heritability of the respective traits. Moderate to high heritability for stem quality traits was estimated irrespectively of region. Overall, better growth and higher heritability suggests that anthropogenic selection within the best inland provenances may constitute better performing and adaptable breeding population compared to the coastal one. Still, overlapping phenotypical variation and heritability of quality traits implies improved stemwood quality for plywood regardless of the provenance region. High adaptive capacity of silver birch genotypes suggests ability to cope with climatic changes, highlighting its potential for climate-smart forestry.Betula pendula

Spectral Reflectance in Silver Birch Genotypes from Three Provenances in Finland

The goal of this study was to investigate the variation in the leaf spectral reflectance and its association with other leaf traits from 12 genotypes among three provenances of origin (populations) in a common garden for Finnish silver birch trees in 2015 and 2016. The spectral reflectance was measured in the laboratory from the detached leaves in the wavelength range of visible and near-infrared (VNIR, 400–1000 nm) and shortwave infrared (SWIR, 1000–2500 nm). The variation among the provenance was initially visualized with principal component analysis (PCA) and a clear separation among the provenances was detected with the discriminant analysis of principal components (DAPC) and partial least squares discriminant analysis (PLS-DA) depicting a less strong variation among the genotypes within the provenances. Wavelengths contributing to the separation of the genotypes and provenances were identified from the contribution plot of DAPC and the red edge was strongly related to the differences. Chlorophyll content showed clear provenance variation and was associated with the separation among the genotypes and provenances in the DAPC space. The normalized difference vegetation index (NDVI705,750) and chlorophyll reflectance index (CRI) showed clear significance among the provenances, whereas NDVI670,780 showed no variation. The variation in the chlorophyll content and the CRI and red edge-based NDVI indices indicated seasonal variation as the chlorophyll content starts increasing in early June. The correlation of foliar chlorophyll content and the chlorophyll-related spectral indices for the discrimination of provenances and genotypes are reported for the first time in a naturally occurring tree species consecutively for two years.

Warming and ozone exposure effects on silver birch (Betula pendula Roth) leaf litter quality, microbial growth and decomposition

Climate warming is expected to accelerate decomposition in boreal forests, but the concomitant effects of tropospheric ozone (O3), a phytotoxic greenhouse gas, alone and in combination with warming, are poorly understood. We studied how these two climatic factors affect leaf litter decomposition of two silver birch genotypes. We used field exposure for growing saplings and native and reciprocal transplant experiments for litter incubation to disentangle environmental and litter quality mediated effects of temperature and O3 on litter mass loss. We analysed litter C% and N% and microbial biomass (using qPCR) in fresh litter and after 217 and 257 days of incubation. Warming decreased fresh litter C% and N% and bacterial DNA, whereas elevated O3 increased N% and bacterial and fungal DNA, equally for both genotypes. In contrast, most effects on microbes during litter incubation varied between the two genotypes. Warming effects on microbes were mainly environmental, but despite having effects on litter quality and microbial growth, warming and O3 both had only weak or no effects on litter mass loss. Litter quality and microbial growth in northern birch stands are likely to change due to warming and O3 exposures, but effects on litter decomposition rate may remain weak.

Root morphology, mycorrhizal roots and extramatrical mycelium growth in silver birch (Betula pendula Roth) genotypes exposed to experimental warming and soil moisture manipulations

In the future, boreal forests will be growing in a warmer climate with more fluctuating soil moisture conditions. However, the knowledge about the effects of simultaneous warming and drought on boreal trees, and especially on their belowground compartments, is still scarce. We studied the responses of four silver birch genotypes to experimental warming at normal and low watering levels after two growing seasons. Variables analysed include mycorrhizal short root growth, fungal biomass (ergosterol) in roots and extramatrical mycelium, and root morphology (root tissue density, root length density, specific root length, specific root surface area). In addition, root and shoot mass fractions were determined. Gt26 had more root mass and also more 2–5 mm diameter roots, but less short roots and ergosterol in its roots than the other genotypes. There were no treatment effects on extramatrical mycelium and mycorrhizal infection levels, but root ergosterol concentrations increased in gt12 due to warming. Drought increased root mass at the expense of shoot growth in gt12, but warming did not cause allometric growth changes in any of the genotypes. Our results show some genotype-dependent responses to warming and drought but also significant within population differences in belowground allocation patterns among the silver birch genotypes.

Contrasting responses of silver birch VOC emissions to short- and long-term herbivory

There is a need to incorporate the effects of herbivore damage into future models of plant volatile organic compound (VOC) emissions at leaf or canopy levels. Short-term (a few seconds to 48 h) changes in shoot VOC emissions of silver birch (Betula pendula Roth) in response to feeding by geometrid moths (Erannis defoliaria Hübner) were monitored online by proton transfer reaction time-of-flight mass spectrometry (PTR-TOF-MS). In addition, two separate field experiments were established to study the effects of long-term foliage herbivory (FH, 30-32 days of feeding by geometrids Agriopis aurantiaria (Clerck) and E. defoliaria in two consecutive years) and bark herbivory (BH, 21 days of feeding by the pine weevil (Hylobius abietis L.) in the first year) on shoot and rhizosphere VOC emissions of three silver birch genotypes (gt14, gt15 and Hausjärvi provenance). Online monitoring of VOCs emitted from foliage damaged by geometrid larvae showed rapid bursts of green leaf volatiles (GLVs) immediately after feeding activity, whereas terpenoid emissions had a tendency to gradually increase during the monitoring period. Long-term FH caused transient increases in total monoterpene (MT) emissions from gt14 and sesquiterpene (SQT) emissions from Hausjärvi provenance, mainly in the last experimental season. In the BH experiment, genotype effects were detected, with gt14 trees having significantly higher total MT emissions compared with other genotypes. Only MTs were detected in the rhizosphere samples of both field experiments, but their emission rates were unaffected by genotype or herbivory. The results suggest that silver birch shows a rapid VOC emission response to short-term foliage herbivory, whereas the response to long-term foliage herbivory and bark herbivory is less pronounced and variable at different time points.

Silver birch telomeres shorten in tissue culture

Shortening of telomeres has been connected with ageing and loss of cell replication or regeneration capacity. The aim of the present study was to examine potential variation in the length of telomeric repeats in silver birch (Betula pendula Roth) using clonal materials consisting of different-aged outdoor trees and tissue cultures of seven genotypes. The overall average length of telomeres was 13.6 kb (±0.3), the minimum length of repeats in the different genotypes varied from 5.9 kb (±0.5) to 9.6 kb (±0.6), and the maximum length varied from 15.3 kb (±1.1) to 22.8 kb (±0.4). When germinated seeds and leaf and cambium samples from 15- and 80-year-old trees were compared, no correlation of ageing and the length of telomeric repeats was found. Positional variation in the telomere length was, however, observed in the cambium of mature trees, the stem base having longer repeats than the upper parts of the tree. Tissue cultures were found to have shorter telomeres than outdoor trees: prolonged culture, callus culture and stressful conditions were all observed to shorten telomeric repeats and should thus be avoided in birch micropropagation. There were significant differences among the studied silver birch genotypes in their telomere length, and these differences were consistent over the sample types. This is the first report on variation of telomeric repeats in a long-living organism studied with clonal materials.

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.

Differential gene expression in senescing leaves of two silver birch genotypes in response to elevated CO2 and tropospheric ozone

Long-term effects of elevated CO(2) and O(3) concentrations on gene expression in silver birch (Betula pendula Roth) leaves were studied during the end of the growing season. Two birch genotypes, clones 4 and 80, with different ozone growth responses, were exposed to 2x ambient CO(2) and/or O(3) in open-top chambers (OTCs). Microarray analyses were performed after 2 years of exposure, and the transcriptional profiles were compared to key physiological characteristics during leaf senescence. There were genotypic differences in the responses to CO(2) and O(3). Clone 80 exhibited greater transcriptional response and capacity to alter metabolism, resulting in better stress tolerance. The gene expression patterns of birch leaves indicated contrasting responses of senescence-related genes to elevated CO(2) and O(3). Elevated CO(2) delayed leaf senescence and reduced associated transcriptional changes, whereas elevated O(3) advanced leaf senescence because of increased oxidative stress. The combined treatment demonstrated that elevated CO(2) only temporarily alleviated the negative effects of O(3). Gene expression data alone were insufficient to explain the O(3) response in birch, and additional physiological and biochemical data were required to understand the true O(3) sensitivity of these clones.

Genetic and environmental determinants of silver birch growth and herbivore resistance

Plant growth and herbivore resistance is affected by both environmental and genetic factors. The aim of this study was to examine (1) whether genotype explains variation in herbivore resistance, plant growth and tolerance of feeding within natural silver birch populations, (2) whether there is significant interaction between birch genotype and environment or the study year, and (3) whether there are defence costs in field grown juvenile silver birch saplings. The 22 genotypes studied were assigned insect exposure and insect removal treatments in two different environments. Resistance to insect herbivory was measured as the amount of undamaged leaves of the saplings and tolerance of feeding as a difference in relative height increment of the saplings between insect exposure and removal treatments in relation to the amount of damage (i.e. opposite number of resistance). Our study shows that a local silver birch population has substantial genotypic variation in resistance to insect herbivores and this variation is not affected by the environment or the study year. Genotypic variation was not observed for tolerance of feeding. We also showed that, in contrast to genotypic variation in resistance, the genotypic variation in growth among silver birch genotypes strongly depended on the environment, i.e. silver birch genotypes have a high degree of phenotypic plasticity in their growth. Therefore, defence costs in terms of lower growth rate appear to be highly dependent on the environment. To conclude, our results indicate that silver birch populations have a good potential to adapt to possibly increasing herbivory in boreal forests in warmer future climates.

Mortality in seedling populations of Silver Birch: genotypic variation and herbivore effects

Summary Variation among mature plants reflects mortality that has occurred during the seedling phase. Our earlier studies have shown variation in resistance to insect and vole herbivory among Silver Birch genotypes that represent variation in a naturally regenerated birch stand. The aim of this study was to examine whether seedling mortality varied among these genotypes, and whether insect and field vole herbivory affected mortality. The 20 genotypes studied competed with each other in dense stands, which were randomly assigned to insect and vole exposure and fertilization treatments. Insect herbivory and fertilization increased mortality significantly (by 98 and 68%, respectively). The fertilization effect was probably due to accelerated self‐thinning among fast‐growing seedlings. Although vole browsing removed considerable biomass, it did not affect seedling mortality. Mortality was context‐dependent. The genotypes differed both in their overall mortality and in their response to insect herbivory. However, the effect of insects on a genotype depended more on its success in competition than on its resistance: even small amounts of feeding were detrimental to short, shaded seedlings, while taller seedlings in the canopy were affected less, although they were eaten more. It is concluded that moderate levels of insect herbivory can increase seedling mortality considerably. Furthermore, herbivory can change the genetic structure of birch populations through selective mortality, which in dense stands is dependent on competitive traits.