The Value of Hybrid Aspen Coppice Investment under Different Discount Rate, Price and Management Scenarios: A Case Study of Estonia

Hybrid aspen is one of the most promising tree species for short-rotation forestry in Northern Europe. After the clearcutting of hybrid aspen plantation, the next generation arises from root and stump sprouts. The economic feasibility of different management strategies of hybrid aspen coppice stands has not yet been comprehensively evaluated in Northern Europe. We compared the land expectation values (LEVs) of hybrid aspen coppice stands managed according to four scenarios: three early thinning methods (corridor, cross-corridor and single-tree) followed by conventional management and intensive bioenergy production (repeated harvests in 5-year rotations) over a 25-year period in hemiboreal Estonia. We considered the historic price volatility of aspen wood assortments under various discount rates (1–20%). We found that the 25-year rotation with different early thinning methods was more profitable than short bioenergy cycles in the case of low discount rates (<5%). The LEV of short coppice cycles for only bioenergy production became more profitable in comparison with those by thinning methods, when higher discount rates (>10%) were applied. Hybrid aspen coppice stands can be managed profitably, but more risks are taken when the management strategy focuses only on bioenergy production.

Saccharification Potential of Transgenic Greenhouse- and Field-Grown Aspen Engineered for Reduced Xylan Acetylation

High acetylation of xylan in hardwoods decreases their value as biorefinery feedstocks. To counter this problem, we have constitutively suppressed RWA genes encoding acetyl-CoA transporters using the 35S promoter, or constitutively and wood-specifically (using the WP promoter) expressed fungal acetyl xylan esterases of families CE1 (AnAXE1) and CE5 (HjAXE), to reduce acetylation in hybrid aspen. All these transformations improved the saccharification of wood from greenhouse-grown trees. Here, we describe the chemical properties and saccharification potential of the resulting lines grown in a five-year field trial, and one type of them (WP:AnAXE1) in greenhouse conditions. Chemically, the lignocellulose of the field- and greenhouse-field-grown plants slightly differed, but the reductions in acetylation and saccharification improvement of engineered trees were largely maintained in the field. The main novel phenotypic observation in the field was higher lignification in lines with the WP promoter than those with the 35S promoter. Following growth in the field, saccharification glucose yields were higher from most transformed lines than from wild-type (WT) plants with no pretreatment, but there was no improvement in saccharification with acid pretreatment. Thus, acid pretreatment removes most recalcitrance caused by acetylation. We found a complex relationship between acetylation and glucose yields in saccharification without pretreatment, suggesting that other variables, for example, the acetylation pattern, affect recalcitrance. Bigger gains in glucose yields were observed in lines with the 35S promoter than in those with the WP promoter, possibly due to their lower lignin content. However, better lignocellulose saccharification of these lines was offset by a growth penalty and their glucose yield per tree was lower. In a comparison of the best lines with each construct, WP:AnAXE1 provided the highest glucose yield per tree from saccharification, with and without pretreatment, WP:HjAXE yields were similar to those of WT plants, and yields of lines with other constructs were lower. These results show that lignocellulose properties of field-grown trees can be improved by reducing cell wall acetylation using various approaches, but some affect productivity in the field. Thus, better understanding of molecular and physiological consequences of deacetylation is needed to obtain quantitatively better results.

Populus PtERF85 Balances Xylem Cell Expansion and Secondary Cell Wall Formation in Hybrid Aspen

Secondary growth relies on precise and specialized transcriptional networks that determine cell division, differentiation, and maturation of xylem cells. We identified a novel role for the ethylene-induced Populus ethylene response factor PtERF85 (Potri.015G023200) in balancing xylem cell expansion and secondary cell wall (SCW) formation in hybrid aspen (Populus tremula x tremuloides). Expression of PtERF85 is high in phloem and cambium cells and during the expansion of xylem cells, while it is low in maturing xylem tissue. Extending PtERF85 expression into SCW forming zones of woody tissues through ectopic expression reduced wood density and SCW thickness of xylem fibers but increased fiber diameter. Xylem transcriptomes from the transgenic trees revealed transcriptional induction of genes involved in cell expansion, translation, and growth. The expression of genes associated with plant vascular development and the biosynthesis of SCW chemical components such as xylan and lignin, was down-regulated in the transgenic trees. Our results suggest that PtERF85 activates genes related to xylem cell expansion, while preventing transcriptional activation of genes related to SCW formation. The importance of precise spatial expression of PtERF85 during wood development together with the observed phenotypes in response to ectopic PtERF85 expression suggests that PtERF85 contributes to the transition of fiber cells from elongation to secondary cell wall deposition.

Populus ERF85 balances xylem cell expansion and secondary cell wall formation in hybrid aspen

Secondary growth relies on precise and specialized transcriptional networks that determine cell division, differentiation, and maturation of xylem cells. We identify a novel role for the ethylene induced Populus ETHYLENE RESPONSE FACTOR ERF85 (Potri.015G023200) in balancing xylem cell expansion and secondary cell wall (SCW) formation in hybrid aspen (Populus tremula x tremuloides). Expression of ERF85 is high in phloem and cambium cells and during expansion of xylem cells, while it is low in maturing xylem tissue. Extending ERF85 expression into SCW forming zones of woody tissues through ectopic expression reduced wood density and SCW thickness of xylem fibers but increased fiber diameter. Xylem transcriptomes from the transgenic trees revealed transcriptional induction of genes involved in cell expansion, translation and growth. Expression of genes associated with plant vascular development and biosynthesis of SCW chemical components such as xylan and lignin, was downregulated in the transgenic trees. Our results suggest that ERF85 activates genes related with xylem cell expansion, while preventing transcriptional activation of genes related to SCW formation. The importance of precise spatial expression of ERF85 during wood development together with the observed phenotypes in response to ectopic ERF85 expression suggests that ERF85 functions as a switch between different phases of xylem differentiation during wood formation.

Plasticity and stability of hybrid aspen clones in 14 field trials over Sweden, Finland and north-west Russia

The study aimed at estimating the genotype × environment (G × E) interaction, stability and plasticity parameters for height growth and survival of hybrid aspen (Populus tremula L. × Populus tremuloides Michx.) clones. A set of 17 hybrid aspen clones were studied at age four in 11 field trials (series 1) and a set of 12 clones were studied at age 12 in three field trials (series 2) in Sweden, Finland and north-west Russia. There was an intermediate and significant correlation between height ranks at four and 12 years, showing modest G × E interaction for tree height. The corresponding correlation for survival was weak and insignificant, indicating a stronger G × E interaction than for height. The stability parameter (S) for height showed large differences among clones. The plasticity index (PI) values for height ranged from 0.6 to 0.8 among clones, demonstrating high levels of phenotypic plasticity.

Isoprene and monoterpene emissions from alder, aspen and spruce short-rotation forest plantations in the United Kingdom

An expansion of bioenergy has been proposed to help reduce fossil-fuel greenhouse gas emissions, and short-rotation forestry (SRF) can contribute to this expansion. However, SRF plantations could also be sources of biogenic volatile organic compound (BVOC) emissions, which can impact atmospheric air quality. In this study, emissions of isoprene and 11 monoterpenes from the branches and forest floor of hybrid aspen, Italian alder and Sitka spruce stands in an SRF field trial in central Scotland were measured during two years (2018–2019) and used to derive emission potentials for different seasons. Sitka spruce was included as a comparison as it is the most extensive plantation species in the UK. Winter and spring emissions of isoprene and monoterpenes were small compared to those in summer. Sitka spruce had a standardised mean emission rate of 15 µgCg-1h-1 for isoprene in the dry and warm summer of 2018 – more than double the emissions in 2019. However, standardised mean isoprene emissions from hybrid aspen were similar across both years, approximately 23 µgCg-1h-1, and standardised mean isoprene emissions from Italian alder were very low. Mean standardised total monoterpene emissions for these species followed a similar pattern of higher standardised emissions in the warmer year: Sitka spruce emitting 4.5 and 2.3 µgCg-1h-1 for 2018 and 2019, aspen emitting 0.3 and 0.09 µgCg-1h-1, and Italian alder emitting 1.5 and 0.2 µgCg-1h-1, respectively. In contrast to these foliage emissions, the forest floor was only a small source of monoterpenes, typically 1 or 2 orders of magnitude lower than foliage emissions on a unit of ground area basis. Estimates of total annual emissions from each plantation type per hectare were derived using the MEGAN 2.1 model. The modelled total BVOC (isoprene and monoterpenes) emissions of SRF hybrid aspen plantations were approximately half those of Sitka spruce for plantations of the same age. Italian alder SRF emissions were 20 times smaller than from Sitka spruce. The expansion of bioenergy plantations to 0.7 Mha has been suggested for the UK to help achieve net-zero greenhouse gas emissions by 2050. The model estimates show that, with such an expansion, total UK BVOC emissions would increase between <1 % and 35 %, depending on the tree species planted. Whereas increases might be small on a national scale, regional increases might have a larger impact on local air quality.

How clonal differences and within-tree heterogeneity affect pore properties of hybrid aspen wood and biochar?

Production of applicable and homogeneous biochar for soil amendment purposes would benefit from knowledge on how feedstock heterogeneity impacts key biochar pore properties and how the properties are transformed due to pyrolysis. This study aimed (1) to quantify how clonal differences and within-tree heterogeneity of a hybrid aspen feedstock (wood) impact biochar pore properties and (2) to estimate how pore properties of non-pyrolysed wood materials are transformed when pyrolysed into biochar. The study was conducted by collecting samples from a hybrid aspen (Populus tremula L. × Populus tremuloides Michx.) clonal field trial. Key pore properties of non-pyrolysed and pyrolysed wood samples were quantified with 3D X-ray imaging and quantitative image analyses. The results demonstrated how pyrolysis shifted distinctively bi-modal pore size distributions of the wood samples towards smaller pore size regions. The bi-modal wood tissue structure controlled the pore structure also in the biochars. Due to decreasing cell wall thicknesses, the pyrolysis increased the porosity of the materials. While the thermal process homogenized differences in the wall thicknesses, the thicknesses of the feedstock were also shown to control the resulting thicknesses in the biochars. Mechanisms of biochar pore property formation can be considered important when designing applicable biochars for a chosen purpose. Clonal differences and within-tree heterogeneity had a direct impact only on the wall thicknesses and the pore diameters of vessels. These impacts can be of interest when planning feedstock utilization in biochar production. However, the results suggest that relatively homogeneous biochar can be produced from hybrid aspen feedstocks.

Organic nitrogen nutrition: LHT1.2 protein from hybrid aspen (Populus tremula L. x tremuloides Michx) is a functional amino acid transporter and a homolog of Arabidopsis LHT1

The contribution of amino acids (AAs) to soil nitrogen (N) fluxes is higher than previously thought. The fact that AA uptake is pivotal for N nutrition in boreal ecosystems highlights plant AA transporters as key components of the N cycle. At the same time, very little is known about AA transport and respective transporters in trees. Tree genomes may contain thirteen or more genes encoding the LYSINE HISTIDINE TRANSPORTER (LHT) family proteins, and this complicates the study of their significance for tree N use efficiency. With the strategy of obtaining a tool to study N use efficiency, our aim was to identify and characterize a relevant AA transporter in hybrid aspen (Populus tremula L. x tremuloides Michx.). We identified PtrLHT1.2, the closest homolog of Arabidopsis thaliana AtLHT1, which is expressed in leaves, stems and roots. Complementation of a yeast AA uptake mutant verified the function of PtrLHT1.2 as an AA transporter. Furthermore, PtrLHT1.2 was able to fully complement the phenotypes of the Arabidopsis AA uptake mutant lht1 aap5, including early leaf senescence-like phenotype, reduced growth, decreased plant N levels and reduced root AA uptake. AA uptake studies finally showed that PtrLHT1.2 is a high affinity transporter for neutral and acidic AAs. Thus, we identified a functional AtLHT1 homolog in hybrid aspen, which harbors the potential to enhance overall plant N levels and hence increase biomass production. This finding provides a valuable tool for N nutrition studies in trees and opens new avenues to optimizing tree N use efficiency.

Stem Volume and Quality of Hybrid and Common Aspen in the Clonal Archive

Hybrid aspen as a fast growing tree species with short rotation is of interest for the creation of plantation forest crops in intensive forest management. Hybrid aspen wood is used to produce paper, lumber, and plywood, as well as biofuel. The research purpose is to compare the volume and quality of the stem of hybrid and common aspen, and to assess interclonal variability by these features in cold climate areas. The research results of 42 hybrid and 10 common aspen clones in the clonal archive of the Institute of Biology of Komi Scientific Centre of the Ural Branch of the Russian Academy of Sciences are presented. The archive was laid out in 2009 by 1-year-old rooted seedlings with a closed root system with the placement of plants 3×3 m. Donor plants were obtained at the Haapastensyrjä Tree Breeding Station of the Natural Resources Institute Finland. Scales have been developed for assessing plant health and stem straightness. Herewith, the height of trees, the presence of sores, scars and frost clefts on their stems were recorded. The share of tree-like and shrub-like plants was 70 and 5 %, respectively, with an average survival of 75 % at the 10-year biological age. Hybrid aspen was statistically significant in terms of superiority to standard aspen in diameter by 65 %, height by 49 %, and stem volume by more than 3 times. The share of slightly crooked stems was about 7 %, and the share of stems with bark cracks was about 3 %. Good survival, stem quality, and rapid growth of hybrid aspen made it possible to conclude that it is a promising species for plantation reforestation in the taiga zone.