Staff Publications

Staff Publications

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    'Staff publications' is the digital repository of Wageningen University & Research

    'Staff publications' contains references to publications authored by Wageningen University staff from 1976 onward.

    Publications authored by the staff of the Research Institutes are available from 1995 onwards.

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Breeding progress and preparedness for mass-scale deployment of perennial lignocellulosic biomass crops switchgrass, miscanthus, willow and poplar
Clifton-Brown, John ; Harfouche, Antoine ; Casler, Michael D. ; Dylan Jones, Huw ; Macalpine, William J. ; Murphy-Bokern, Donal ; Smart, Lawrence B. ; Adler, Anneli ; Ashman, Chris ; Awty-Carroll, Danny ; Bastien, Catherine ; Bopper, Sebastian ; Botnari, Vasile ; Brancourt-Hulmel, Maryse ; Chen, Zhiyong ; Clark, Lindsay V. ; Cosentino, Salvatore ; Dalton, Sue ; Davey, Chris ; Dolstra, Oene ; Donnison, Iain ; Flavell, Richard ; Greef, Joerg ; Hanley, Steve ; Hastings, Astley ; Hertzberg, Magnus ; Hsu, Tsai Wen ; Huang, Lin S. ; Iurato, Antonella ; Jensen, Elaine ; Jin, Xiaoli ; Jørgensen, Uffe ; Kiesel, Andreas ; Kim, Do Soon ; Liu, Jianxiu ; McCalmont, Jon P. ; McMahon, Bernard G. ; Mos, Michal ; Robson, Paul ; Sacks, Erik J. ; Sandu, Anatolii ; Scalici, Giovanni ; Schwarz, Kai ; Scordia, Danilo ; Shafiei, Reza ; Shield, Ian ; Slavov, Gancho ; Stanton, Brian J. ; Swaminathan, Kankshita ; Trindade, Luisa M. - \ 2019
Global change biology Bioenergy 11 (2019)1. - ISSN 1757-1693 - p. 118 - 151.
bioenergy - feedstocks - lignocellulose - M. sacchariflorus - M. sinensis - Miscanthus - Panicum virgatum - perennial biomass crop - Populus spp. - Salix spp.

Genetic improvement through breeding is one of the key approaches to increasing biomass supply. This paper documents the breeding progress to date for four perennial biomass crops (PBCs) that have high output–input energy ratios: namely Panicum virgatum (switchgrass), species of the genera Miscanthus (miscanthus), Salix (willow) and Populus (poplar). For each crop, we report on the size of germplasm collections, the efforts to date to phenotype and genotype, the diversity available for breeding and on the scale of breeding work as indicated by number of attempted crosses. We also report on the development of faster and more precise breeding using molecular breeding techniques. Poplar is the model tree for genetic studies and is furthest ahead in terms of biological knowledge and genetic resources. Linkage maps, transgenesis and genome editing methods are now being used in commercially focused poplar breeding. These are in development in switchgrass, miscanthus and willow generating large genetic and phenotypic data sets requiring concomitant efforts in informatics to create summaries that can be accessed and used by practical breeders. Cultivars of switchgrass and miscanthus can be seed-based synthetic populations, semihybrids or clones. Willow and poplar cultivars are commercially deployed as clones. At local and regional level, the most advanced cultivars in each crop are at technology readiness levels which could be scaled to planting rates of thousands of hectares per year in about 5 years with existing commercial developers. Investment in further development of better cultivars is subject to current market failure and the long breeding cycles. We conclude that sustained public investment in breeding plays a key role in delivering future mass-scale deployment of PBCs.

Multi-allelic QTL analysis of protein content in a bi-parental population of cultivated tetraploid potato
Klaassen, Michiel T. ; Bourke, Peter M. ; Maliepaard, Chris ; Trindade, Luisa M. - \ 2019
Euphytica 215 (2019)2. - ISSN 0014-2336
Alleles - Haplotypes - Potato - Protein content - QTL analysis - Tetraploid

Protein content is a key quality trait for the potato starch industry. The objective of this study was to identify allele-specific quantitative trait loci (QTLs) for tuber protein content in cultivated potato (Solanum tuberosum L.) at the tetraploid level. We analysed 496 full-sib F1 clones in a 3-year field trial to dissect the complex genetic architecture of soluble tuber protein content. Genotypic data from a 60K single nucleotide polymorphism (SNP) array was used for SNP dosage scoring, constructing homologue specific linkage maps and assembly of a dense integrated chromosomal linkage map. From the integrated map, probabilistic multi-locus identity-by-descent (IBD) haplotypes (alleles) were estimated and used to detect associations between the IBD haplotypes and the phenotypic trait values. Moderate levels of trait heritability were estimated between 40 and 74% that correspond with previous studies. Our contemporary naive analysis identified potential additive QTLs on chromosomes 2, 3, 5 (top arm) and 9 across the years. Moreover, cofactor QTL analysis identified two masked QTLs on chromosomes 1 and 5 (lower arm). The QTLs on chromosomes 2, 5 (lower arm) and 9 are reported here for the first time. The QTLs that we identified on chromosomes 1, 3 and 5 (top arm) show overlap with previous studies for protein content in potato. Collectively the naive QTLs explained 12 to 17% of the phenotypic variance. The underlying alleles of the QTLs provided both positive and negative effects on the phenotype. Our work uncovers the complex genetic architecture of this trait and describes potential breeding strategies for improvement. As protein has emerged as a high-value component from industrial potato starch production, the dissection of the genetic architecture and subsequent improvement of this trait by breeding has great economic and environmental relevance.

Life cycle assessment of ethanol production from miscanthus : A comparison of production pathways at two European sites
Lask, Jan ; Wagner, Moritz ; Trindade, Luisa M. ; Lewandowski, Iris - \ 2019
Global change biology Bioenergy 11 (2019)1. - ISSN 1757-1693 - p. 269 - 288.
biofuel - environmental performance - life cycle assessment - lignocellulosic ethanol - marginal land - miscanthus - perennial crop - pretreatment

Lignocellulosic ethanol represents a renewable alternative to petrol. Miscanthus, a perennial plant that grows on marginal land, is characterized by efficient use of resources and is considered a promising source of lignocellulosic biomass. A life cycle assessment (LCA) was performed to determine the environmental impacts of ethanol production from miscanthus grown on marginal land in Great Britain (Aberystwyth) and an average-yield site in Germany (Stuttgart; functional unit: 1 GJ). As the conversion process has substantial influence on the overall environmental performance, the comparison examined three pretreatment options for miscanthus. Overall, results indicate lower impacts for the production in Stuttgart in comparison with the corresponding pathways in Aberystwyth across the analysed categories. Disparities between the sites were mainly attributed to differences in biomass yield. When comparing the conversion options, liquid hot water treatment resulted in the lowest impacts, followed by dilute sulphuric acid. Dilute sodium hydroxide pretreatment represented the least favourable option. Site-dependent variation in biomass composition and degradability did not have substantial influence on the environmental performance of the analysed pathways. Additionally, implications of replacing petrol with miscanthus ethanol were examined. Ethanol derived from miscanthus resulted in lower impacts with respect to greenhouse gas emissions, fossil resource depletion, natural land transformation and ozone depletion. However, for other categories, including toxicity, eutrophication and agricultural land occupation, net scores were substantially higher than for the fossil reference. Nevertheless, the results indicate that miscanthus ethanol produced via dilute acid and liquid hot water treatment at the site in Stuttgart has the potential to comply with the requirements of the European Renewable energy directive for greenhouse gas emission reduction. For ethanol production at the marginal site, carbon sequestration needs to be considered in order to meet the requirements for greenhouse gas mitigation.

Latitudinal adaptation and genetic insights into the origins of cannabis sativa L.
Zhang, Qingying ; Chen, Xuan ; Guo, Hongyan ; Trindade, Luisa M. ; Salentijn, Elma M.J. ; Guo, Rong ; Guo, Mengbi ; Xu, Yanping ; Yang, Ming - \ 2018
Advances in Intelligent Systems and Computing 871 (2018). - ISSN 2194-5357
Cannabaceae - CpDNA - Genetic diversity - Industrial hemp - Phylogeography

Cannabis is one of the most important industrial crops distributed worldwide. However, the phylogeographic structure and domestication knowledge of this crop remains poorly understood. In this study, sequence variations of five chloroplast DNA (cpDNA) regions were investigated to address these questions. For the 645 individuals from 52 Cannabis accessions sampled (25 wild populations and 27 domesticated populations or cultivars), three haplogroups (Haplogroup H, M, L) were identified and these lineages exhibited distinct high-middle-low latitudinal gradients distribution pattern. This pattern can most likely be explained as a consequence of climatic heterogeneity and geographical isolation. Therefore, we examined the correlations between genetic distances and geographical distances, and tested whether the climatic factors are correlated with the cpDNA haplogroup frequencies of populations. The “isolation-by-distance” models were detected for the phylogeographic structure, and the day-length was found to be the most important factor (among 20 BioClim factors) that influenced the population structures. Considering the distinctive phylogeographic structures and no reproductive isolation among members of these lineages, we recommend that Cannabis be recognized as a monotypic genus typified by Cannabis sativa L., containing three subspecies: subsp. sativa, subsp. Indica, and subsp. ruderalis. Within each haplogroup which possesses a relatively independent distribution region, the wild and domesticated populations shared the most common haplotypes, indicating that there are multiregional origins for the domesticated crop. Contrast to the prevalent Central-Asia-Origin hypothesis of C. saltiva, molecular evidence reveals for the first time that the low latitude haplogroup (Haplogroup L) is the earliest divergent lineage, implying that Cannabis is probably originated in low latitude region.

Exploring natural genetic variation in tomato sucrose synthases on the basis of increased kinetic properties
Dinh, Quy Dung ; Finkers, Richard ; Westphal, Adrie H. ; Dongen, Walter M.A.M. van; Visser, Richard G.F. ; Trindade, Luisa M. - \ 2018
PLoS ONE 13 (2018)10. - ISSN 1932-6203

Sucrose synthase (SuSy) is one key enzyme directly hydrolyzing sucrose to supply substrates for plant metabolism, and is considered to be a biomarker for plant sink strength. Improvement in plant sink strength could lead to enhanced plant growth and yield. Cultivated tomatoes are known to have a narrow genetic diversity, which hampers further breeding for novel and improved traits in new cultivars. In this study, we observed limited genetic variation in SuSy1, SuSy3 and SuSy4 in 53 accessions of cultivated tomato and landraces, but identified a wealth of genetic diversity in 32 accessions of related wild species. The variation in the deduced amino acid sequences was grouped into 23, 22, and 17 distinct haplotypes for SuSy1/3/4, respectively. Strikingly, all known substrate binding sites were highly conserved, as well as most of the phosphorylation sites except in SuSy1. Two SuSy1 and three SuSy3 protein variants were heterologously expressed to study the effect of the amino acid changes on enzyme kinetic properties, i.e. maximal sucrose hydrolyzing capacity (Vmax), affinity for sucrose (Km), and catalytic efficiency (Vmax/Km) at 25°C and 16°C. SuSy1-haplotype#3 containing phosphorylation site Ser-16 did not have an improvement in the kinetic properties compared to the reference SuSy1-haplotype#1 containing Arg-16. Meanwhile SuSy3-haplotype#9 from a wild accession, containing four amino acid changes S53A, S106I, E727D and K741E, showed an increase in Vmax/Km at 16°C compared to the reference SuSy3-haplotype#1. This study demonstrates that SuSy kinetic properties can be enhanced by exploiting natural variation, and the potential of this enzyme to improve sucrose metabolism and eventually sink strength in planta.

Detailed biochemical and morphologic characteristics of the green microalga Neochloris oleoabundans cell wall
Rashidi, Behzad ; Trindade, Luisa M. - \ 2018
Algal Research 35 (2018). - ISSN 2211-9264 - p. 152 - 159.
Biochemical composition - Cell wall - Chlorophyta - Electron microscopy - Green microalgae - Neochloris oleoabundans

Chlorophyta, the group of green algae of which there are >6000 species, manifests a great diversity of intercellular and extracellular components. Building blocks in the cell walls of Chlorophyta are very distinct and they may contain various components. Here, we characterize the cell walls of Neochloris oleoabundans, a Chlorophyte microalga, both in terms of biochemical composition and morphology. N. oleoabundans cell walls are composed of about 24.3% carbohydrates, 31.5% proteins, 22.2% lipids and 7.8% inorganic material, which contrasts to the cell walls of (higher) terrestrial plants in which carbohydrates are by far the main component. We also observed that cell wall carbohydrates are mainly non-cellulosic polysaccharides, essentially composed of rhamnose, galactose, glucuronic acid and glucosamine, of which glucose is only a minor component. The lipids comprising the N. oleoabundans cell walls are generally wax/cutin-like. Electron microscopic studies revealed that N. oleabundans cell walls are approximately 200 nm thick and consist of two main layers: a thinner inner layer and a more electron-dense outer layer. On the outer layer are hair-like structures that are possibly rich in carbohydrates. These findings are an important contribution that enable us to understand the complexity of cell walls in green microalgae.

Editorial: Optimizing miscanthus for the sustainable bioeconomy: From genes to products
Farrar, Kerrie ; Heaton, Emily A. ; Trindade, Luisa M. - \ 2018
Frontiers in Plant Science 9 (2018). - ISSN 1664-462X
Bioeconomy - Biomass quality - Crop yield and phenology - Environmental impact - Establishment and crop management - Miscanthus
Heterologous expression of two Arabidopsis starch dikinases in potato
Xu, Xuan ; Dees, Dianka ; Huang, Xing Feng ; Visser, Richard G.F. ; Trindade, Luisa M. - \ 2018
Starch-Stärke 70 (2018)1-2. - ISSN 0038-9056
Glucan - Granule morphology - Starch metabolism - Storage starch - Transitory starch - Water dikinase
Starch phosphate esters influence physiochemical properties of starch granules that are essential both for starch metabolism and industrial use of starches. To modify properties of potato starch and understand the effect of starch phosphorylation on starch metabolism in storage starch, the starch dikinases from Arabidopsis thaliana, glucan water dikinase 2 and 3 (AtGWD2 and AtGWD3), were heterologously expressed in potato tubers (Solanum tuberosum) from two genetic backgrounds: the amylose-containing clone Kardal and the amylose-free mutant amf. Modified starches showed altered granule morphology, but no significant changes in starch phosphate content were observed. Genes involved in starch metabolism did not show altered expression in the transgenic lines relatively to the control; however, sucrose synthase 4 (SUSY4) was upregulated in transgenic tubers with AtGWD2 expression. Taken together, these results demonstrated that expression of AtGWD2 and AtGWD3 are not suitable for engineering starch with a high phosphate content in potato tubers but can provide new insights in the process of starch phosphorylation in the amyloplast.
Improving sustainability of maize to ethanol processing by plant breeding and process optimization
Slegers, P.M. ; Torres Salvador, A.F. ; Boxtel, A.J.B. van; Trindade, L.M. - \ 2017
In: Proceedings of the 25th European Biomass Conference. - Stockholm : EUBCE (European Biomass Conference and Exhibition Proceedings ) - ISBN 9788889407172 - p. 1031 - 1033.
Efficient management of plant resources is essential for a sustainable biobased economy. The biomass conversion efficiency and sustainability performance depend greatly on the choice of feedstock and the applied processing technology. The aim of this research was to enhance the biomass use of maize stover for bioethanol production, by combining plant breeding of the maize feedstock with various pretreatment severities and applying an exploratory assessment of the environmental and economic impacts. We found that systematic genetic gains of cell wall digestibility can lead to significant advances in the total glucose productivity and also in the sustainability performance. The best maize characteristics tested led to a total glucose productivity of 3.7 ton per hectare using mild processing conditions. This matches the highest realizable yields under severe processing conditions. In the best scenarios the environmental and economic impacts of operating conditions were reduced by 15% compared to the benchmark.
A tandem CBM25 domain of α-amylase from Microbacterium aurum as potential tool for targeting proteins to starch granules during starch biosynthesis
Huang, Xing Feng ; Nazarian, Farhad ; Vincken, Jean Paul ; Visser, Richard G.F. ; Trindade, Luisa M. - \ 2017
BMC Biotechnology 17 (2017)1. - ISSN 1472-6750
CBM20 - CBM25 - Starch - Starch binding domain - Transgenic potato
Background: Starch-binding domains from carbohydrate binding module family 20 have been used as a tool for starch engineering. Previous studies showed that expression of starch binding domain fusion proteins in planta resulted in modified starch granule structures and physicochemical properties. However, although 13 carbohydrate binding module families have been reported to contain starch-binding domains, only starch-binding domains from carbohydrate binding module family 20 have been well studied and introduced into plants successfully. In this study, two fragments, the tandem CBM25 domain and the tandem CBM25 with multiple fibronectin type III (FN3) domains of the α-amylase enzyme from Microbacterium aurum, were expressed in the tubers of a wild type potato cultivar (cv. Kardal) and an amylose-free (amf) potato mutant. Results: The (CBM25)2 and FN3 protein were successfully accumulated in the starch granules of both Kardal and amf transformants. The accumulation of (CBM25)2 protein did not result in starch morphological alterations in Kardal but gave rise to rough starch granules in amf, while the FN3 resulted in morphological changes of starch granules (helical starch granules in Kardal and rough surface granules in amf) but only at a very low frequency. The starches of the different transformants did not show significant differences in starch size distribution, apparent amylose content, and physico-chemical properties in comparison to that of untransformed controls. Conclusion: These results suggest that the starch-binding domains from carbohydrate binding module family 25 can be used as a novel tool for targeting proteins to starch granules during starch biosynthesis without side-effects on starch morphology, composition and properties.
Effect of the middle lamella biochemical composition on the non-linear behaviour of technical fibres of hemp under tensile loading using strain mapping
Fuentes, C.A. ; Willekens, P. ; Petit, J. ; Thouminot, C. ; Müssig, J. ; Trindade, L.M. ; Vuure, A.W. Van - \ 2017
Composites Part A: Applied Science and Manufacturing 101 (2017). - ISSN 1359-835X - p. 529 - 542.
A. Biocomposite - A. Natural fibres - B. Fibre deformation - Strain mapping
This manuscript describes the effects of alterations in biochemical composition on structural morphology and the mechanical behaviour of technical fibres of hemp used for composite applications. First, the strength and apparent Young's modulus distribution of technical fibres of hemp of 96 hemp samples, corresponding to 32 different hemp accessions cultivated in 3 locations, were analysed using Weibull distribution. From these, 2 samples (one with high and one with low fibre strength) were selected for further analysis. Next, full-field strain measurement at the micro-scale during tensile loading via digital image correlation analysis was used for evaluating both, the stress-strain behaviour at a global scale and the local mechanical behaviour heterogeneity at a micro-scale, along a technical fibre of hemp. The analysis reveals 2 typical types of tensile stress-strain curves, and a complex and very irregular pattern of strain concentrations, which are associated to the technical fibre strength. The non-linear behaviour of the stress-strain curve is explained by the development of shear strain at the elementary fibre (botanically defined as the individual cell) interphases. Micro tomography and biochemical analysis of the technical fibre microstructure showed that alterations in cell wall composition, in particular substitution of pectin, leads to changes in the non-linear behaviour of technical fibres of hemp under tensile loading.
Environmental influences on the growing season duration and ripening of diverse Miscanthus germplasm grown in six countries
Nunn, Christopher ; Hastings, Astley Francis St John ; Kalinina, Olena ; Özgüven, Mensure ; Schüle, Heinrich ; Tarakanov, Ivan G. ; Weijde, Tim van der; Anisimov, Aleksander A. ; Iqbal, Yasir ; Kiesel, Andreas ; Khokhlov, Nikolay F. ; McCalmont, Jon P. ; Meyer, Heike ; Mos, Michal ; Schwarz, Kai Uwe ; Trindade, Luisa M. ; Lewandowski, Iris ; Clifton-Brown, John - \ 2017
Frontiers in Plant Science 8 (2017). - ISSN 1664-462X - 14 p.
Miscanthus - Modeling - Multi-location - Ripening - Senescence

The development of models to predict yield potential and quality of a Miscanthus crop must consider climatic limitations andthe duration of growing season. As a biomass crop, yield and quality are impacted by the timing of plant developmental transitions such as flowering andsenescence. Growth models are available for the commercially grown clone Miscanthus x giganteus (Mxg), but breeding programs have been working to expand the germplasmavailable, including development of interspecies hybrids. The aimof this study was to assess the performance of diverse germplasmbeyond the range of environments considered suitable for a Miscanthus crop to be grown. To achieve this, six field sites were planted as part of the EU OPTIMISC project in 2012 in a longitudinal gradient from West to East: Wales—Aberystwyth, Netherlands—Wageningen, Stuttgart—Germany, Ukraine—Potash, Turkey—Adana, and Russia—Moscow. Each field trial contained three replicated plots of the same 15 Miscanthus germplasmtypes. Through the 2014 growing season, phenotypic traits were measured to determine the timing of developmental stages key to ripening; the tradeoff between growth (yield) and quality (biomass ash and moisture content). The hottest site (Adana) showed an accelerated growing season, with emergence, flowering and senescence occurring before the other sites. However, the highest yields were produced at Potash, where emergence was delayed by frost and the growing season was shortest. Flowering triggers varied with species and only in Mxg was strongly linked to accumulated thermal time. Our results show that a prolonged growing season is not essential to achieve high yields if climatic conditions are favorable and in regions where the growing season is bordered by frost, delaying harvest can improve quality of the harvested biomass.

Genetic complexity of miscanthus cell wall composition and biomass quality for biofuels
Weijde, Tim van der; Lessa Alvim Kamei, Claire ; Severing, Eduard ; Torres Salvador, Andres Francisco ; Gomez, Leonardo D. ; Dolstra, Oene ; Maliepaard, Chris A. ; McQueen-Mason, Simon J. ; Visser, Richard G.F. ; Trindade, Luisa M. - \ 2017
BMC Genomics 18 (2017). - ISSN 1471-2164 - 15 p.
Biofuel - Biomass quality - Cell wall composition - Conversion efficiency - Genetic map - Miscanthus - Quantitative trait loci (QTL) - Saccharification efficiency - Yield

Background: Miscanthus sinensis is a high yielding perennial grass species with great potential as a bioenergy feedstock. One of the challenges that currently impedes commercial cellulosic biofuel production is the technical difficulty to efficiently convert lignocellulosic biomass into biofuel. The development of feedstocks with better biomass quality will improve conversion efficiency and the sustainability of the value-chain. Progress in the genetic improvement of biomass quality may be substantially expedited by the development of genetic markers associated to quality traits, which can be used in a marker-assisted selection program. Results: To this end, a mapping population was developed by crossing two parents of contrasting cell wall composition. The performance of 182 F1 offspring individuals along with the parents was evaluated in a field trial with a randomized block design with three replicates. Plants were phenotyped for cell wall composition and conversion efficiency characters in the second and third growth season after establishment. A new SNP-based genetic map for M. sinensis was built using a genotyping-by-sequencing (GBS) approach, which resulted in 464 short-sequence uniparental markers that formed 16 linkage groups in the male map and 17 linkage groups in the female map. A total of 86 QTLs for a variety of biomass quality characteristics were identified, 20 of which were detected in both growth seasons. Twenty QTLs were directly associated to different conversion efficiency characters. Marker sequences were aligned to the sorghum reference genome to facilitate cross-species comparisons. Analyses revealed that for some traits previously identified QTLs in sorghum occurred in homologous regions on the same chromosome. Conclusion: In this work we report for the first time the genetic mapping of cell wall composition and bioconversion traits in the bioenergy crop miscanthus. These results are a first step towards the development of marker-assisted selection programs in miscanthus to improve biomass quality and facilitate its use as feedstock for biofuel production.

Extending miscanthus cultivation with novel germplasm at six contrasting sites
Kalinina, Olena ; Nunn, Christopher ; Sanderson, Ruth ; Hastings, Astley F.S. ; Weijde, Tim van der; Özgüven, Mensure ; Tarakanov, Ivan ; Schüle, Heinrich ; Trindade, Luisa M. ; Dolstra, Oene ; Schwarz, Kai Uwe ; Iqbal, Yasir ; Kiesel, Andreas ; Mos, Michal ; Lewandowski, Iris ; Clifton-Brown, John C. - \ 2017
Frontiers in Plant Science 8 (2017). - ISSN 1664-462X
Establishment - Marginal land - Miscanthus - Multi-location field trials - Novel hybrids - Productivity

Miscanthus is a genus of perennial rhizomatous grasses with C4 photosynthesis which is indigenous in a wide geographic range of Asian climates. The sterile clone, Miscanthus × giganteus (M. × giganteus), is a naturally occurring interspecific hybrid that has been used commercially in Europe for biomass production for over a decade. Although, M. × giganteus has many outstanding performance characteristics including high yields and low nutrient offtakes, commercial expansion is limited by cloning rates, slow establishment to a mature yield, frost, and drought resistance. In this paper, we evaluate the performance of 13 novel germplasm types alongside M. × giganteus and horticultural “Goliath” in trials in six sites (in Germany, Russia, The Netherlands, Turkey, UK, and Ukraine). Mean annual yields across all the sites and genotypes increased from 2.3 ± 0.2 t dry matter ha−1 following the first year of growth, to 7.3 ± 0.3, 9.5 ± 0.3, and 10.5 ± 0.2 t dry matter ha−1 following the second, third, and fourth years, respectively. The highest average annual yields across locations and four growth seasons were observed for M. × giganteus (9.9 ± 0.7 t dry matter ha−1) and interspecies hybrid OPM-6 (9.4 ± 0.6 t dry matter ha−1). The best of the new hybrid genotypes yielded similarly to M. × giganteus at most of the locations. Significant effects of the year of growth, location, species, genotype, and interplay between these factors have been observed demonstrating strong genotype × environment interactions. The highest yields were recorded in Ukraine. Time needed for the crop establishment varied depending on climate: in colder climates such as Russia the crop has not achieved its peak yield by the fourth year, whereas in the hot climate of Turkey and under irrigation the yields were already high in the first growing season. We have identified several alternatives to M. × giganteus which have provided stable yields across wide climatic ranges, mostly interspecies hybrids, and also Miscanthus genotypes providing high biomass yields at specific geographic locations. Seed-propagated interspecific and intraspecific hybrids, with high stable yields and cheaper reliable scalable establishment remain a key strategic objective for breeders.

Site-specific management of miscanthus genotypes for combustion and anaerobic digestion : A comparison of energy yields
Kiesel, Andreas ; Nunn, Christopher ; Iqbal, Yasir ; Weijde, Tim Van der; Wagner, Moritz ; Özgüven, Mensure ; Tarakanov, Ivan ; Kalinina, Olena ; Trindade, Luisa M. ; Clifton-Brown, John ; Lewandowski, Iris - \ 2017
Frontiers in Plant Science 8 (2017). - ISSN 1664-462X
Biogas - Biomass - Energy yield - Harvest time - Moisture content - Substrate-specific methane yield - Yield
In Europe, the perennial C4 grass miscanthus is currently mainly cultivated for energy generation via combustion. In recent years, anaerobic digestion has been identified as a promising alternative utilization pathway. Anaerobic digestion produces a higher-value intermediate (biogas), which can be upgraded to biomethane, stored in the existing natural gas infrastructure and further utilized as a transport fuel or in combined heat and power plants. However, the upgrading of the solid biomass into gaseous fuel leads to conversion-related energy losses, the level of which depends on the cultivation parameters genotype, location, and harvest date. Thus, site-specific crop management needs to be adapted to the intended utilization pathway. The objectives of this paper are to quantify (i) the impact of genotype, location and harvest date on energy yields of anaerobic digestion and combustion and (ii) the conversion losses of upgrading solid biomass into biogas. For this purpose, five miscanthus genotypes (OPM 3, 6, 9, 11, 14), three cultivation locations (Adana, Moscow, Stuttgart), and up to six harvest dates (August-March) were assessed. Anaerobic digestion yielded, on average, 35% less energy than combustion. Genotype, location, and harvest date all had significant impacts on the energy yield. For both, this is determined by dry matter yield and ash content and additionally by substrate-specific methane yield for anaerobic digestion and moisture content for combustion. Averaged over all locations and genotypes, an early harvest in August led to 25%and a late harvest to 45%conversion losses. However, each utilization option has its own optimal harvest date, determined by biomass yield, biomass quality, and cutting tolerance. By applying an autumn green harvest for anaerobic digestion and a delayed harvest for combustion, the conversion-related energy loss was reduced to an average of 18%. This clearly shows that the delayed harvest required tomaintain biomass quality for combustion is accompanied by high energy losses through yield reduction over winter. The pre-winter harvest applied in the biogas utilization pathway avoids these yield losses and largely compensates for the conversion-related energy losses of anaerobic digestion.
Transgenic modification of potato pectic polysaccharides also affects type and level of cell wall xyloglucan
Huang, Jie Hong ; Jiang, Rui ; Kortstee, Anne ; Dees, Dianka C.T. ; Trindade, Luisa M. ; Gruppen, Harry ; Schols, Henk A. - \ 2017
Journal of the Science of Food and Agriculture 97 (2017)10. - ISSN 0022-5142 - p. 3240 - 3248.
Cell wall polysaccharides - Non-targeted modification - Oligomers - Solanum tuberosum
BACKGROUND: Genes encoding pectic enzymes were introduced into wild-type potato Karnico. Cell wall materials were extracted from Karnico and transgenic lines expressing β-galactosidase (β-Gal-14) or rhamnogalacturonan lyase (RGL-18). Pectic polysaccharides from the β-Gal-14 transgenic line exhibited rhamnogalacturonan-I structural elements with shorter galactan side chains, whereas the RGL-18 transgenic line had less rhamnogalacturonan-I structures than Karnico. Xyloglucan in primary cell walls interacts with pectin and other cell wall polysaccharides and controls cell growth. RESULTS: Xyloglucan extracts from transgenic lines had different levels of monosaccharides compared to wild-type. Most XXGG-type xyloglucans from Karnico and RGL-18 alkali-extractable extracts predominantly consisted of XXGG and XSGG building blocks. Karnico and RGL-18 4molL-1 extracts had small proportions of the XXXG-type xyloglucan, whereas β-Gal-14 extracts also contained the XXXG-type xyloglucan. The peak ratios of XSGG/XXGG were 1.9, 2.4 and 1.1 for 4molL-1 extracts of Karnico, RGL-18 and β-Gal-14 lines, respectively. CONCLUSION: After transgenic modification on pectin, the xyloglucan building blocks may have been changed. The β-Gal-14 lines mostly present XXXG-type repeating units instead of the XXGG-type in 4molL-1 extracts. The ratio of XSGG/XXGG repeating units also changed, indicating that the transgenic modification of pectin altered xyloglucan structure during plant development.
Stability of cell wall composition and saccharification efficiency in Miscanthus across diverse environments
Weijde, Tim van der; Dolstra, Oene ; Visser, Richard G.F. ; Trindade, Luisa M. - \ 2017
Frontiers in Plant Science 7 (2017). - ISSN 1664-462X
Biomass quality - Ethanol - Genotype-by-environment interaction - GGE biplot - Miscanthus - Multi-location trial - Near-infrared spectroscopy (NIRS) - Stability

To investigate the potential effects of differences between growth locations on the cell wall composition and saccharification efficiency of the bioenergy crop miscanthus, a diverse set of 15 accessions were evaluated in six locations across Europe for the first 3 years following establishment. High-throughput quantification of cellulose, hemicellulose and lignin contents, as well as cellulose and hemicellulose conversion rates was achieved by combining near-infrared reflectance spectroscopy (NIRS) and biochemical analysis. Prediction models were developed and found to predict biomass quality characteristics with high accuracy. Location significantly affected biomass quality characteristics in all three cultivation years, but location-based differences decreased toward the third year as the plants reached maturity and the effect of location-dependent differences in the rate of establishment reduced. In all locations extensive variation in accession performance was observed for quality traits. The performance of the different accessions in the second and third cultivation year was strongly correlated, while accession performance in the first cultivation year did not correlate well with performance in later years. Significant genotype-by-environment (G × E) interactions were observed for most traits, revealing differences between accessions in environmental sensitivity. Stability analysis of accession performance for calculated ethanol yields suggested that selection for good and stable performance is a viable approach. Environmental influence on biomass quality is substantial and should be taken into account in order to match genotype, location and end-use of miscanthus as a lignocellulose feedstock.

Engineering potato starch with a higher phosphate content
Xu, Xuan ; Huang, Xing Feng ; Visser, Richard G.F. ; Trindade, Luisa M. - \ 2017
PLoS ONE 12 (2017)1. - ISSN 1932-6203
Phosphate esters are responsible for valuable and unique functionalities of starch for industrial applications. Also in the cell phosphate esters play a role in starch metabolism, which so far has not been well characterized in storage starch. Laforin, a human enzyme composed of a carbohydrate-binding module and a dual-specificity phosphatase domain, is involved in the dephosphorylation of glycogen. To modify phosphate content and better understand starch (de)phosphorylation in storage starch, laforin was engineered and introduced into potato (cultivar Kardal). Interestingly, expression of an (engineered) laforin in potato resulted in significantly higher phosphate content of starch, and this result was confirmed in amylose-free potato genetic background (amf). Modified starches exhibited altered granule morphology and size compared to the control. About 20±30% of the transgenic lines of each series showed red-staining granules upon incubation with iodine, and contained higher phosphate content than the blue-stained starch granules. Moreover, low amylose content and altered gelatinization properties were observed in these red-stained starches. Principle component and correlation analysis disclosed a complex correlation between starch composition and starch physico-chemical properties. Ultimately, the expression level of endogenous genes involved in starch metabolism was analysed, revealing a compensatory response to the decrease of phosphate content in potato starch. This study provides a new perspective for engineering starch phosphate content in planta by making use of the compensatory mechanism in the plant itself.
Starch phosphorylation plays an important role in starch biosynthesis
Xu, Xuan ; Dees, Dianka ; Dechesne, Annemarie ; Huang, Xing Feng ; Visser, Richard G.F. ; Trindade, Luisa M. - \ 2017
Carbohydrate Polymers 157 (2017). - ISSN 0144-8617 - p. 1628 - 1637.
Freeze-thaw stability - Glucan - Starch metabolism - Starch phosphate content - Storage starch - Water dikinase
Starch phosphate esters are crucial in starch metabolism and render valuable functionality to starches for various industrial applications. A potato glucan, water dikinase (GWD1) was introduced in tubers of two different potato genetic backgrounds: an amylose-containing line Kardal and the amylose-free mutant amf. In both backgrounds, this resulted in two contrasting effects, a number of plants showed higher phosphate content compared to the respective control, while others lines exhibited lower phosphate content, thereby generating two series of starches with broad-scale variation in phosphate content. The results of systematic analyses on these two series of starches revealed that starch phosphate content strongly influenced starch granule morphology, amylose content, starch fine structure, gelatinization characteristics and freeze-thaw stability of starch gels. Further analyses on the expression level of genes involved in starch metabolism suggested that starch phosphorylation regulates starch synthesis by controlling the carbon flux into starch while simultaneously modulating starch-synthesizing genes.
Evaluation of both targeted and non-targeted cell wall polysaccharides in transgenic potatoes
Huang, Jie Hong ; Kortstee, Anne ; Dees, Dianka C.T. ; Trindade, Luisa M. ; Visser, Richard G.F. ; Gruppen, Harry ; Schols, Henk A. - \ 2017
Carbohydrate Polymers 156 (2017). - ISSN 0144-8617 - p. 312 - 321.
Galacturonosyltransferase 1 - Pectin biosynthesis - Plant cell wall - Rhamnosyltransferase - Solanum tuberosum

In this study, we analyze 31 transgenic lines and their respective untransformed background lines to determine the transgene effects on targeted structures including the pectin components rhamnogalacturonan I (RG-I) and homogalacturonan (HG), neutral side chains (galactan/arabinangalactan), acetylation of pectin, and cellulose level. Modification arising from the pectin backbone- or pectin side chain transgenic lines either increased or decreased the HG:RG-I ratio, side chain length, and methyl esterification of pectin in the tuber cell wall. The pectin esterification transgenic line exhibited only limited side effects. The cellulose level-targeting transgenic lines yielded an unexpectedly high HG:RG-I ratio and longer pectic side chains. These results clearly demonstrate that in effects of a transgene are not restricted to the direct activity of the targeted enzyme but have consequences for the structure of the cell wall matrix. Analysis of whole cell wall structure is therefore necessary to assess the complete effect, direct and indirect, of a transgene.

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