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Vegetatively propagated hybrid aspen (Populus tremula × P. tremuloides) is one of the fastest growing tree species in Finland. To study clonal variation in element concentrations in wood, bark and ...

Funding Information: This work was supported by Business Finland corona-co-creation funding for the project Antiviral Fibers?pilot with extracts from Finnish forests (grant: 40699/31/2020). This study was also funded by the Natural Resources Institute Finland?s strategic research funding to the projects ?More, faster, higher quality: potential of short-rotation aspen and willow biomass for novel products in bioeconomy? (AspenWill) and ?Added value potential of new and under-utilized fibre sources in Finnish value networks of green bioeconomy: prefeasibility, prototyping, and market acceptance? (VALUEPOT). In addition, Academy of Finland has supported this study via the project ?Antivirals from Forest Biomasses: Structure, Function, and Applicability? (grant: 342250). Work was also funded by Jane and Aatos Erkko Foundation: Novel probes for discovering antivirals. Publisher Copyright: Copyright © 2021 Tienaho, Reshamwala, Sarjala, Kilpeläinen, Liimatainen, Dou, Viherä-Aarnio, Linnakoski, Marjomäki and Jyske. Earlier studies have shown that the bark of Salix L. species (Salicaceae family) is rich in extractives, such as diverse bioactive phenolic compounds. However, we lack knowledge on the bioactive properties of the bark of willow species and clones adapted to the harsh climate conditions of the cool temperate zone. Therefore, the present study aimed to obtain information on the functional profiles of northern willow clones for the use of value-added bioactive solutions. Of the 16 willow clones studied here, 12 were examples of widely distributed native Finnish willow species, including dark-leaved willow (S. myrsinifolia Salisb.) and tea-leaved willow (S. phylicifolia L.) (3 + 4 clones, respectively) and their natural and artificial hybrids (3 + 2 clones, respectively). The four remaining clones were commercial willow varieties from the Swedish willow breeding program. Hot water extraction of bark under mild conditions was carried out. Bioactivity assays were used to screen antiviral, antibacterial, antifungal,yeasticidal, and antioxidant activities, as well as the total phenolic content of the extracts. Additionally, we introduce a fast and less labor-intensive steam-debarking method for Salix spp. feedstocks. Clonal variation was observed in the antioxidant properties of the bark extracts of the 16 Salix spp. clones. High antiviral activity against a non-enveloped enterovirus, coxsackievirus A9, was found, with no marked differences in efficacy between the native clones. All the clones also showed antibacterial activity against Staphylococcus aureus and Escherichia coli, whereas no antifungal (Aspergillus brasiliensis) or yeasticidal (Candida albicans) efficacy was detected. When grouping the clone extract results into Salix myrsinifolia, Salix phylicifolia, native hybrid, artificial hybrid, and commercial clones, there was a significant difference in the activities between S. phylicifolia clone extracts and commercial clone extracts in the favor of S. phylicifolia in the antibacterial and antioxidant tests. In some antioxidant tests, S. phylicifolia clone extracts were also significantly more active than artificial clone extracts. Additionally, S. myrsinifolia clone extracts showed significantly higher activities in some antioxidant tests than commercial clone extracts and artificial clone extracts. Nevertheless, the bark extracts of native Finnish willow clones showed high bioactivity. The obtained knowledge paves the way towards developing high value-added biochemicals and other functional solutions based on willow biorefinery approaches.

Novel bioeconomic approaches call for increasingly faster production of lignocellulosic biomass and its bettertailored use for higher added value. The high-yield capacity and structural properties of willows (Salix spp.) suggest their excellent potential for the production of designed biochar for use in agronomic, electronic and technical applications. All these applications rely on the internal pore structure of biochar. However, we lack an in-depth quantitative understanding of the interlinkages between the feedstock properties and the physical quality of the biochar produced. We studied quantitatively how the clonal and within-plant properties of five different willow clones (hybrids of Salix schwerinii E.L. Wolf) affected the micrometre-scale pore properties of the produced biochars (pyrolyzed at + 462 ?C). The porosity and pore size distribution were analysed before and after slow pyrolysis by X-ray microtomography and image analysis. We also studied the potential of conventional low-cost fibre analysis techniques to be used to predict biochar pore properties directly from fresh feedstock. The total porosity (0.55?0.62) and the pore size distribution of willow wood and derived biochars varied between clones. Approximately two-thirds of the biochar total porosity was associated with pores formed by wood fibres. Pyrolysis levelled off the structural variation detected between and within the clones. Pyrolysis-induced shrinkage reduced the pore sizes and narrowed the pore size distribution. The results suggest that conventional fibre analysis techniques could be utilized to predict biochar homogeneity. Short rotation coppice willows are suitable feedstock to produce homogenous biochar precursor for production of bio-based carbon materials to be used in high value-added technical applications. The structural homogeneity of the feedstock and produced biochar can be enhanced by selecting proper harvesting strategy and clones used in plantations. From the industrial perspective, comprehensive understanding of feedstock properties helps to control quality of the produced biochar.

Even though individuals with a deviant morphology have been elemental in genetics of model species, they have thus far been largely ignored in the studies of forest trees. Here we studied the inheritance of curly-grained and brown-figured wood phenotype in curly birch (Betula pendula var. carelica (Mercklin) Hämet-Ahti). In addition of the figured wood, curly birches display reduced and aberrant growth, indicating that the causative locus (loci) is (are) vital for normal tree development. To explore the genetic basis of this mutation, we studied the inheritance of the curly birch phenotype in a progeny trial (crosses between curly birch parent trees, between curly and normal phenotypes, and from selfings of curly trees). Based on the external morphology, the phenotypes of 11-year-old progeny trees were scored as either curly or wild. Based on the phenotypic segregation ratios, we postulate a simple Mendelian inheritance model for curliness: (i) a one-locus, two-allele model in which the allele coding for curly phenotype is dominant over the allele coding for normal phenotype and (ii) the semi-dominant curly allele is lethal when homozygous. We expect that further studies on the molecular genetic basis of the curly birch phenotype will provide valuable information on the developmental pathways involved in wood formation.

Studies addressing endodormancy release in adult trees are usually carried out using twigs detached from the trees in the experiments. Potential problems caused by cutting the root–shoot connection when detaching the twigs can be avoided by using grafts as the experimental material. We studied the effects of chilling on the endodormancy release in Norway spruce (Picea abies (L.) Karst.) grafts where twigs of 16-, 32- and 80-year-old trees were used as the scions. The grafts were first exposed to chilling in natural conditions and then samples of them were transferred at intervals to a regrowth test in forcing conditions in a greenhouse. The bud burst percentage, BB%, in the forcing conditions generally increased from zero to near 100% with increasing previous chilling accumulation from mid-October until mid-November, indicating that endodormancy was released in almost all of the grafts by mid-November. The days to bud burst, DBB, decreased in the forcing conditions with successively later transfers until the next spring. Neither BB% nor DBB was dependent on the age of the scion. However, in the early phase of ecodormancy release, the microscopic internal development of the buds was more advanced in the grafts representing the 16-year-old than in those representing the 32- or 80-year-old trees. In conclusion, our findings suggest that no major change in the environmental regulation of endodormancy release in Norway spruce takes place when the trees get older. Taken together with earlier findings with Norway spruce seedlings, our results suggest that regardless of the seedling or tree age, the chilling requirement of endodormancy release is met in late autumn. The implications of our findings for Norway spruce phenology under climatic warming and the limitations of our novel method of using grafts as a proxy of trees of different ages are discussed.

This study was conducted to examine the characteristics of dominant height growth and develop site index models for clonal hybrid aspen plantations in southern Finland. Data were obtained from repeatedly measured clonal hybrid aspen trials with varying initial spacing: 2.5 m × 2.5 m (1600 trees ha−1), 3.0 m × 3.0 m (1200 trees ha−1), 3.5 m × 3.5 m (800 trees ha−1), and 5.0 m × 5.0 m (400 trees ha−1). The total number of data points in the analysis was 389 for the age of 3–20. Within the range of observed data, the dominant height grew linearly over age and was significantly different due to the initial planting density; growth was higher when the planting was denser. Using the initial density effect, dominant height growth models were developed based on the Chapman-Richards function through nonlinear mixed-effects modelling. The density variable was found to be statistically the best variable when modifying only the shape parameter of the Chapman-Richards function. All fixed-effects were significant for both models, with and without the density effect. The residual plots of the model did not show any bias over the predicted value, stand age or planting density. The predicted dominant height was higher with increasing initial density. The predicted dominant height increment was faster with higher planting densities until the age of 14 years. The anamorphic site index curves were presented with base age of 20 years including the planting density effect. The overall pattern of site index curves was consistent with those observed in previous studies. The models developed in this study can be used to estimate the dominant height and site index of hybrid aspen plantations in southern Finland.

Hybrid aspen ( Ã ) is one of the fastest growing tree species in Finland. During the mid-1990s, a breeding programme was started with the aim of selecting clones that were superior in producing pulpwood. Hybrid aspen can also be grown as a short-rotation crop for bioenergy. To study clonal variation in wood and bark properties, seven clones were selected from a 12-year-old field trial located in southern Finland. From each clone, five trees were harvested and samples were taken from stem wood, stem bark and branches to determine basic density, effective heating value, moisture and ash content. Vertical within-tree variation in moisture content and basic density was also studied. The differences between clones were significant for almost all studied properties. For all studied properties there was a significant difference between wood and bark. Wood had lower ash content (0.5% vs. 3.9%), basic density (378 kg m vs. 450 kg m) and effective heating value (18.26 MJ kg vs. 19.24 MJ kg), but higher moisture content (55% vs. 49%) than bark. The values for branches were intermediate. These results suggest that the properties of hybrid aspen important for energy use could be improved by clonal selection. However, selecting clones based on fast growth only may be challenging since it may lead to a decrease in hybrid aspen wood density.Populus tremulaP. tremuloidesâ3â3â1â1

We studied the effect of seed transfers on survival, yield and stem defects of silver birch (Betula pendula Roth) of North European origin in five parallel provenance trials in Finland. The trials were located at Loppi (60°39′N) in southern Finland, at Kannonkoski (62°58′N), Ilomantsi (62°58′N) and Toholampi (63°47′N) in central Finland and at Rovaniemi (66°21′N) in northern Finland. Four trials were growing on moist upland forest sites and one on agricultural land. The material consisted of altogether 38 stand seed origins from Finland, Sweden, Estonia, Great Britain and Russia ranging between latitudes 53° and 67°N. Survival, height, diameter at breast height, stem volume/ha and number of stem defects per tree were assessed when the trees were 19 years old. Seed transfer distance was calculated either as latitudinal or temperature sum difference between the seed origin and the trial location. According to coefficients of determination (R2), the latitude-based transfer models gave a slightly better fit with the data than the models based on temperature sum differences. Significant differences were found among the origins regarding all studied traits in all trials. In all traits, the pattern of variation in relation to the transfer distance was curvilinear, and the effect of transfer distance was significant. A long transfer from both directions led to a reduction in all measured traits. The optimal transfer distance and direction varied depending on the trait and covariate used in the models. According to the latitudinal model, local or slightly more northern origins had highest survival and lowest number of stem defects. Height growth and yield was at highest with the local or slightly more southern origins. A northward transfer of ca. 2° of latitude increased yield, while a southward transfer or a longer northward transfer, decreased the yield. According to the temperature sum model, origins within the limits of approximately ±200 d.d. compared to the trial location produced the highest yield. Origins with similar or slightly lower temperature sums compared to the trial locations had highest survival and best stem quality. The current seed transfer guidelines for silver birch in Finland, which recommend transfer distances of 150 km or 150 d.d. at maximum in southward or northward direction, seem still valid. Because of the low gain in expected yield by even moderate northward transfers and the uncertainties in the success of more southern origins in varying future climate, we recommend no changes in seed transfer guidelines.

Silver birch ( Roth) seed origins from the Baltic countries and from Finland were compared in terms of growth, wood density, bark thickness and the incidence of darkened core wood, frost cracks and decay, and the effect of seed origin latitude was examined in two Finnish provenance trials. The material consisted of 21 stand and single tree origins ranging from latitudes 54° to 63°N from the Baltic countries and Finland. The trials, measured at the age of 22 years, were located at Tuusula (60°21´N), southern Finland and at Viitasaari (63°11´N), central Finland. The Baltic origins were superior to the Finnish ones in the southern trial regarding height, whereas in central Finland the Finnish origins grew better. There was no consistent difference between the Baltic and the Finnish group of origins in wood density. Bark thickness decreased with increasing seed origin latitude. The Baltic origins had significantly thicker bark than the Finnish origins. A moderate positive correlation was detected between the seed origin latitude and the incidence of darkened core wood in the southern trial, where the darkened core wood was more common in the Finnish origins than in the Baltic ones. The highest proportion of trees with frost cracks was detected in the south-western Latvian origins growing in central Finland. Seed transfers from the Baltic would have an increasing effect on the bark thickness of birch logs, but no or only minor effects on wood density. Based on our results, there is no reason to recommend the use of non-native Baltic seed origins in Finland instead of the native locally adapted seed sources.Betula pendula

Summary In Europe, two commercially important treelike birch species occur naturally: silver birch ( Betula pendula Roth) and downy birch ( Betula pubescens Ehrh.). Both species have a wide natural distribution area on the Eurasian continent, ranging from the Atlantic to eastern Siberia. Although birches occur throughout almost the whole of Europe, the most abundant birch resources are in the temperate and boreal forests of Northern Europe. In the Baltic and Nordic countries, the proportion of birch out of the total volume of the growing stock varies between 11 and 28 per cent. In Northern Europe, birch is commercially the most important broadleaved tree species. Birches are light-demanding early successional pioneer species, which grow both in mixed stands and in pure stands. This article provides an overview of the most important ecological characteristics and typical growth and yield patterns of birch, based on European scientifi c literature. Growth and yield research on birch has been relatively active in Northern Europe, where numerous growth and yield models have been developed during the last decades. In this paper, a list of published scientifi c articles on growth modelling is provided and is grouped according to the different types of model. When growing in forest stands, birches have a relatively straight slender stem form. The current practices and silvicultural recommendations, based on research directed at high-quality timber production in silver birch stands, are reviewed. Although the emphasis is on even-aged pure silver birch stands, the management of mixed stands as well as the silviculture of downy birch and curly birch are also briefl y discussed.

We studied the light and stereomicroscopic structure of developing vegetative buds from a 16-year-old Norway spruce [Picea abies (L.) Karst.] of southern Finnish origin in relation to temperature sum and to externally visible changes in the buds before and during bud burst in forcing conditions. Branches were collected on 17 January and transferred to the greenhouse where they were first subjected to preforcing conditions (darkness, +4 degrees C) for 7 days and then to the forcing conditions (day length 12 h, +20 degrees C). Buds were sampled 20 times between 17 January and 13 February. Air temperature was recorded hourly throughout the study period. The first microscopic change was a temporary increase in the size and number of lipid droplets before the onset of temperature sum (Tor = +5 degrees C) accumulation. From the 4th to the 9th day under the forcing conditions, tracheids started to develop from the base up to the top of the bud. This was closely synchronized with an observed morphological change in the shape of needle tip from rounded to pointed ones. Development from the first visible change in the bud scales on the 12th forcing day to bud burst took 9 days when the temperature sum was 313 d.d. The temperature sums in our experiment overestimated the requirements of temperature sum for bud development phases measured in the field. Bud development could be divided into four structural phases. The first two phases, i.e., morphological changes in the primary needles, occurred without any externally visible changes in the buds. Thus, these phases have a potential for testing and improving the phenological models, which, up to now, have mainly been based on the bud burst observation by the naked eye.

We studied the variation in critical night length for bud set in two photoperiodic ecotypes (two latitudinally distant stands) of silver birch (Betula pendula Roth) in three phytotron experiments. Seeds from 21 open-pollinated mother trees in a southern (Tuusula, 60 degrees N) and a northern (Kittilä, 67 degrees N) Finnish stand were germinated and grown for 4 weeks in a 24-h photoperiod in a greenhouse and then moved to different night length treatments at 18 degrees C for 4 to 6 weeks. Night lengths from 5 to 8.5 h were used for southern origin seedlings and from 1 to 4.5 h for northern origin seedlings. At the end of the treatments, apical bud set was observed and the percentage of seedlings with bud set calculated for each treatment and tree progeny. The critical night lengths (CNL) for 50% bud set were determined separately for seedlings from each mother tree by regression analysis. In both ecotypes, the mean percentage of seedlings with bud set was lowest for the shortest night lengths and increased rapidly as night lengths increased. Mean CNL with its 95% confidence interval for the southern and northern ecotypes was 6.3 +/- 0.2 and 3.1 +/- 0.3 h, respectively. The CNL of the two ecotypes differed significantly in three experiments. Within-ecotype variance of the CNL was significantly higher in the northern ecotype (0.484) than in the southern ecotype (0.150). Significant differences in CNL were detected between individual mother trees of the southern ecotype, but not between mother trees of the northern ecotype. The ranking of individual mother trees, based on CNL, differed in the three experiments.

The first genetic linkage map for silver birch, Betula pendula Roth, was constructed by using a pseudo-testcross mapping strategy and integration of linkage data from 3 unrelated 2-generation pedigrees. The map is based on the genetic inheritance and segregation of 82 amplified fragment length polyhmorphisms and 19 microsatellite markers, and was constructed by simultaneously comparing the performance of CRI-MAP and OUTMAP packages. The analysis revealed 16 linkage groups, and the total map coverage is 1561 cM (Kosambi units). Average map distance between adjacent markers is 15.5 cM. Linkage groups range between 6 and 18 loci and from 81.2 to 326.5 cM; the remaining 9 linkage groups consist of 2 or 3 loci ranging from 6.3 to 42.4 cM. The uncertainty of the map is illustrated with sensitivity analysis. This initial map can serve as a basis for developing a more detailed genetic map.Key words: Betula pendula, linkage map, microsatellite, AFLP, CRI-MAP, OUTMAP.

The overwintering of trees in northern areas depends on processes regulated by photoperiod and temperature. To identify the physiological and genetic factors involved in this environmental control, three latitudinal ecotypes of pubescent birch (Betula pubescens Ehrh.) growing in a common garden experiment were used. Each ecotype responded to the shortening of the photoperiod according to its specific critical daylength, resulting in the induction of freezing tolerance and dehydration of buds first in the northern ecotype, followed by the central and southern ecotypes, respectively. By contrast, there was no clear difference in the timing of dormancy release, bud rehydration, and deacclimation in the spring, suggesting that these traits were controlled mainly by temperature. To elucidate the role of dehydrins (DHN) in the overwintering process, two DHN genomic clones were isolated from pubescent birch and expression of the corresponding genes, both in field and under controlled conditions, was characterized. BpuDhn1 was found to encode an Y(n)K(n)-type of basic DHN, while BpuDhn2 encoded an acidic, SK(n)-type of DHN. In field-grown trees the level of BpuDhn1 increased in buds during the autumn, while the level of BpuDhn2 was highest during the coldest winter months. Under controlled conditions BpuDhn1 increased in response to the combined effect of short daylength and low, non-freezing temperatures whereas the expression of BpuDhn2 was mainly controlled by low temperature while photoperiod had less effect on its expression. These results suggest that DHNs participate in the sensitive environmental regulation of the overwintering process in birch.

Woody plants in the temperate and boreal zone undergo annual cycle of growth and dormancy under seasonal changes. Growth cessation and dormancy induction in autumn are prerequisites for the development of substantial cold hardiness in winter. During evolution, woody plants have developed different ecotypes that are closely adapted to the local climatic conditions. In this study, we employed distinct photoperiodic ecotypes of silver birch (Betula pendula Roth) to elucidate differences in these adaptive responses under seasonal changes. In all ecotypes, short day photoperiod (SD) initiated growth cessation and dormancy development, and induced cold acclimation. Subsequent low temperature (LT) exposure significantly enhanced freezing tolerance but removed bud dormancy. Our results suggested that dormancy and freezing tolerance might partially overlap under SD, but these two processes were regulated by different mechanisms and pathways under LT. Endogenous abscisic acid (ABA) levels were also altered under seasonal changes; the ABA level was low during the growing season, then increased in autumn, and decreased in winter. Compared with the southern ecotype, the northern ecotype was more responsive to seasonal changes, resulting in earlier growth cessation, cold acclimation and dormancy development in autumn, higher freezing tolerance and faster dormancy release in winter, and earlier bud flush and growth initiation in spring. In addition, although there was no significant ecotypic difference in ABA level during growing season, the rates and degrees of ABA alterations were different between the ecotypes in autumn and winter, and could be related to ecotypic differences in dormancy and freezing tolerance.

A number of environmental cues including short day photoperiod (SD) and low temperature (LT) are known to interact in triggering growth cessation, cold acclimation and other adaptive responses in temperate-zone tree species. Proper timing of these responses is particularly important for survival of trees in the boreal and subarctic regions. Therefore, we used a northern tree species, silver birch (Betula pendula Roth) as an experimental model to investigate the effect of SD and LT on development of freezing tolerance and on levels of endogenous abscisic acid (ABA) in short-term experiments under controlled conditions. We characterized differences in SD and LT-induced cold acclimation between three different climatic ecotypes from southern, central and northern habitats. The results demonstrated that cold acclimation was rapidly triggered by exposing the plants to SD or LT, and that a combination of the different treatments had an additive effect on freezing tolerance. Freezing tolerance induction was not uniform in the different tissues, the buds and leaves developed freezing tolerance more rapidly than the stem, and the young leaves had a higher freezing tolerance than the old leaves. The ability of the leaves to respond to SD and LT and similarity of the bud and leaf responses indicate that birch leaves provide a rapid and convenient system for studies on molecular mechanisms of cold acclimation. Development of freezing tolerance was dependent on the climatic ecotype, the northern ecotype was clearly more responsive to both SD and LT than the two more southern ecotypes. Development of freezing tolerance induced by SD and LT was accompanied by transient changes in ABA levels. These alterations in ABA levels were ecotype-dependent, the northern ecotype reacting more strongly to the environmental cues.

The timing of budburst of temperate trees is known to be controlled by complicated interactions of temperature and photoperiod. To improve the phenological models of budburst, better knowledge of the internal bud development preceding budburst in relation to environmental cues is needed. We studied the effect of accumulated chilling and forcing temperatures on the internal development of vegetative buds preceding budburst in Norway spruce [Picea abies (L.) Karst.]. Branches from 17-year-old trees of southern Finnish origin were transferred eight times at 1- to 2-week intervals from October to December 2007 from the field at Punkaharju (61°48 ′N, 29°20′E) to the greenhouse with forcing conditions (day length 12 h, +20 °C). After seven different durations of forcing, the developmental phase and primordial shoot growth of the buds were analysed at the stereomicroscopic level. Air temperature was recorded hourly throughout the study period. The accumulated chilling unit sum had a significant effect on the temperature sum that was required to attain a certain developmental phase; a higher amount of chilling required a lower amount of forcing. The variation in the rate of development of different buds within each sample branch in relation to the chilling unit and forcing temperature sum was low. Regarding primordial shoot growth, there was also an inverse relation between accumulated chilling and forcing, i.e., a higher accumulated chilling unit sum before forcing required a lower temperature sum to initiate primordial shoot growth and resulted in a stronger effect of accumulated forcing. A second-order regression model with an interaction of chilling and forcing explained the variation of primordial shoot growth with high precision (R 2 = 0.88). However, further studies are required to determine the final parameter values to be used in phenological modelling.

The general applicability of RAPDs (random amplified polymorphic DNAs) as genetic molecular markers for Betulapendula Roth was evaluated. On average 2.5 clear, potentially useful marker bands were generated per primer from the DNA of a randomly selected birch tree using a set of 60 commercial 10-mer primers. The resulting fragments were typically between 200 and 700 base pairs in length. A novel pooled-progeny analysis of parentage was applied for a posteriori confirmation of parents of an old breeding experiment. The method that saves labour in several laboratory steps is regarded as useful for screening possible errors in breeding. The present work dealt with a case in which one parent was known and there were two candidates for the other. The correct parent was independently confirmed using nine different RAPD loci. The validity of the pooled-progeny application was confirmed by conventional segregation analysis. In this connection, segregation of RAPD alleles at additional loci in the B. pendula cross was also studied.