European soybean to benefit people and the environment.

Europe imports large amounts of soybean that are predominantly used for livestock feed, mainly sourced from Brazil, USA and Argentina. In addition, the demand for GM-free soybean for human consumption is project to increase. Soybean has higher protein quality and digestibility than other legumes, along with high concentrations of isoflavones, phytosterols and minerals that enhance the nutritional value as a human food ingredient. Here, we examine the potential to increase soybean production across Europe for livestock feed and direct human consumption, and review possible effects on the environment and human health. Simulations and field data indicate rainfed soybean yields of 3.1 ± 1.2 t ha-1 from southern UK through to southern Europe (compared to a 3.5 t ha-1 average from North America). Drought-prone southern regions and cooler northern regions require breeding to incorporate stress-tolerance traits. Literature synthesized in this work evidenced soybean properties important to human nutrition, health, and traits related to food processing compared to alternative protein sources. While acknowledging the uncertainties inherent in any modelling exercise, our findings suggest that further integrating soybean into European agriculture could reduce GHG emissions by 37-291 Mt CO2e year-1 and fertiliser N use by 0.6-1.2 Mt year-1, concurrently improving human health and nutrition.

A new Rothamsted long-term field experiment for the twenty-first century: principles and practice.

Agriculture faces potentially competing societal demands to produce food, fiber and fuel while reducing negative environmental impacts and delivering regulating, supporting and cultural ecosystem services. This necessitates a new generation of long-term agricultural field experiments designed to study the behavior of contrasting cropping systems in terms of multiple outcomes. We document the principles and practices of a new long-term experiment of this type at Rothamsted, established at two contrasting sites in 2017 and 2018, and report initial yield data at the crop and system level. The objective of the Large-Scale Rotation Experiment was to establish gradients of system properties and outcomes to improve our fundamental understanding of UK cropping systems. It is composed of four management factors-phased rotations, cultivation (conventional vs reduced tillage), nutrition (additional organic amendment vs standard mineral fertilization) and crop protection (conventional vs smart crop protection). These factors were combined in a balanced design resulting in 24 emergent cropping systems at each site and can be analyzed at the level of the system or component management factors. We observed interactions between management factors and with the environment on crop yields, justifying the systems level, multi-site approach. Reduced tillage resulted in lower wheat yields but the effect varied with rotation, previous-crop and site. Organic amendments significantly increased spring barley yield by 8% on average though the effect again varied with site. The plowed cropping systems tended to produce higher caloric yield overall than systems under reduced tillage. Additional response variables are being monitored to study synergies and trade-offs with outcomes other than yield at the cropping system level. The experiment has been established as a long-term resource for inter-disciplinary research. By documenting the design process, we aim to facilitate the adoption of similar approaches to system-scale agricultural experimentation to inform the transition to more sustainable cropping systems. Supplementary Information: The online version contains supplementary material available at 10.1007/s13593-023-00914-8.

The potential for soybean to diversify the production of plant-based protein in the UK.

Soybean (Glycine max) offers an important source of plant-based protein. Currently much of Europe's soybean is imported, but there are strong economic and agronomic arguments for boosting local production. Soybean is grown in central and eastern Europe but is less favoured in the North due to climate. We conducted field trials across three seasons and two sites in the UK to test the viability of early-maturing soybean varieties and used the data from these trials to calibrate and validate the Rothamsted Landscape Model. Once validated, the model was used to predict the probability soybean would mature and the associated yield for 26 sites across the UK based on weather data under current, near-future (2041-60) and far-future (2081-2100) climate. Two representative concentration pathways, a midrange mitigation scenario (RCP4.5) and a high emission scenario (RCP8.5) were also explored. Our analysis revealed that under current climate early maturing varieties will mature in the south of the UK, but the probability of failure increases with latitude. Of the 26 sites considered, only at one did soybean mature for every realisation. Predicted expected yields ranged between 1.39 t ha-1 and 1.95 t ha-1 across sites. Under climate change these varieties are likely to mature as far north as southern Scotland. With greater levels of CO2, yield is predicted to increase by as much as 0.5 t ha-1 at some sites in the far future, but this is tempered by other effects of climate change meaning that for most sites no meaningful increase in yield is expected. We conclude that soybean is likely to be a viable crop in the UK and for similar climates at similar latitudes in Northern Europe in the future but that for yields to be economically attractive for local markets, varieties must be chosen to align with the growing season.

Breeding progress and preparedness for mass-scale deployment of perennial lignocellulosic biomass crops switchgrass, miscanthus, willow and poplar.

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.

Optimizing the bioenergy water footprint by selecting SRC willow canopy phenotypes: regional scenario simulations.

BACKGROUND AND AIMS: Bioenergy is central for the future energy mix to mitigate climate change impacts; however, its intricate link with the water cycle calls for an evaluation of the carbon-water nexus in biomass production. The great challenge is to optimize trade-offs between carbon harvest and water use by choosing cultivars that combine low water use with high productivity. METHODS: Regional scenarios were simulated over a range of willow genotype × environment interactions for the major UK soil × climate variations with the process-based model LUCASS. Soil available water capacity (SAWC) ranged from 51 to 251 mm and weather represented the north-west (wet, cool), north-east (dry, cool), south-west (wet, warm) and south-east (dry, warm) of the UK. Scenario simulations were evaluated for small/open narrow-leaf (NL) versus large/closed broad-leaf (BL) willow canopy phenotypes using baseline (1965-89) and warmer recent (1990-2014) weather data. KEY RESULTS: The low productivity under baseline climate in the north could be compensated by choosing BL cultivars (e.g. 'Endurance'). Recent warmer climate increased average productivity by 0.5-2.5 t ha-1, especially in the north. The modern NL cultivar 'Resolution' had the smallest and most efficient water use. On marginal soils (SAWC <100 mm), yields remained below an economic threshold of 9 t ha-1 more frequently under baseline than recent climate. In the drought-prone south-east, 'Endurance' yielded less than 'Resolution', which consumed on average 17 mm year-1 less water. Assuming a planting area of 10 000 ha, in droughty years between 1.3 and 4.5 × 106 m3 of water could be saved, with a small yield penalty, for 'Resolution'. CONCLUSIONS: With an increase in air temperature and occasional water scarcities expected with climate change, high-yielding NL cultivars should be the preferred choice for sustainable use of marginal lands and reduced competition with agricultural food crops.

Dry matter losses and quality changes during short rotation coppice willow storage in chip or rod form.

This study compares dry matter losses and quality changes during the storage of SRC willow as chips and as rods. A wood chip stack consisting of approximately 74 tonnes of fresh biomass, or 31 tonnes dry matter (DM) was built after harvesting in the spring. Three weeks later, four smaller stacks of rods with an average weight of 0.8 tonnes, or 0.4 tonnes DM were built. During the course of the experiment temperature recorders placed in the stacks found that the wood chip pile reached 60 °C within 10 days of construction, but the piles of rods remained mostly at ambient temperatures. Dry matter losses were calculated by using pre-weighed independent samples within the stacks and by weighing the whole stack before and after storage. After 6 months the wood chip stack showed a DM loss of between 19.8 and 22.6%, and mean losses of 23.1% were measured from the 17 independent samples. In comparison, the rod stacks showed an average stack DM loss of between 0 and 9%, and between 1.4% and 10.6% loss from the independent samples. Analysis of the stored material suggests that storing willow in small piles of rods produces a higher quality fuel in terms of lower moisture and ash content; however, it has a higher fine content compared to storage in chip form. Therefore, according to the two storage methods tested here, there may be a compromise between maximising the net dry matter yield from SRC willow and the final fine content of the fuel.

Testing the Use of Static Chamber Boxes to Monitor Greenhouse Gas Emissions from Wood Chip Storage Heaps.

This study explores the use of static chamber boxes to detect whether there are fugitive emissions of greenhouse gases (GHGs) from a willow chip storage heap. The results from the boxes were compared with those from 3-m stainless steel probes inserted into the core of the heap horizontally and vertically at intervals. The results from probes showed that there were increases of carbon dioxide (CO2) concentrations in the heap over the first 10 days after heap establishment, which were correlated with a temperature rise to 60 °C. As the CO2 declined, there was a small peak in methane (CH4) concentration in probes orientated vertically in the heap. Static chambers positioned at the apex of the heap detected some CO2 fluxes as seen in the probes; however, the quantities were small and random in nature. A small (maximum 5 ppm) flux in CH4 occurred at the same time as the probe concentrations peaked. Overall, the static chamber method was not effective in monitoring fluxes from the heap as there was evidence that gases could enter and leave around the edges of the chambers during the course of the experiment. In general, the use of standard (25 cm high) static chambers for monitoring fluxes from wood chip heaps is not recommended.

Dry Matter Losses and Greenhouse Gas Emissions From Outside Storage of Short Rotation Coppice Willow Chip.

This study examined the dry matter losses and the greenhouse gas (GHG) concentrations within two short rotation coppice (SRC) willow wood chip storage heaps. One heap was built on a grassland area (East Midlands) and the other (Rothamsted) on a concrete hard standing. A series of 1- and 3-m probes were embedded in the heaps in order to retrieve gas samples for analysis, and pre-weighed net bags were positioned in the core of the heap to detect dry matter losses. The bagged samples showed dry matter losses of 18 and 19 % in the East Midlands and Rothamsted heaps after 210 and 97 days storage, respectively. The Rothamsted heap showed a whole-heap dry matter loss of 21 %. During this time, the wood chips dried from 54 to 39 % moisture content in the East Midlands heap and 50 to 43 % at Rothamsted. The results from analysing the whole Rothamsted heap indicated an overall loss of 1.5 GJ per tonne stored, although measurements from bagged samples in the core suggested that the chips dried sufficiently to have a minimal energy loss from storage. The process of mixing the heap, however, led to incorporation of wet outer layers and hence the average moisture content was higher in an average sample of chip. After establishment of the heaps, the temperature rose rapidly and this correlated with a peak in carbon dioxide (CO2) concentration within the heap. A peak in methane (CH4) concentration was also detected in both heaps, though more noticeably in the East Midlands heap after around 55 days. In both instances, the peak CH4 concentration occurred as CO2 concentrations dropped, suggesting that after an active period of aerobic decomposition in the first 2 months of storage, the conditions in the heap became anaerobic. The results from this study suggest that outside wood chip storage is not an efficient method of storing biomass, though this may be location-specific as there are some studies showing lower dry matter losses. It is necessary to explore other methods of harvesting SRC to minimise losses and optimise land use efficiency. Further research is required to detect whether there are fugitive emissions of CH4 from wood chip heaps, as this will compromise the net GHG savings from utilising the biomass stored in this way.

Dry Matter Losses and Methane Emissions During Wood Chip Storage: the Impact on Full Life Cycle Greenhouse Gas Savings of Short Rotation Coppice Willow for Heat.

A life cycle assessment (LCA) approach was used to examine the greenhouse gas (GHG) emissions and energy balance of short rotation coppice (SRC) willow for heat production. The modelled supply chain includes cutting multiplication, site establishment, maintenance, harvesting, storage, transport and combustion. The relative impacts of dry matter losses and methane emissions from chip storage were examined from a LCA perspective, comparing the GHG emissions from the SRC supply chain with those of natural gas for heat generation. The results show that SRC generally provides very high GHG emission savings of over 90 %. The LCA model estimates that a 1, 10 and 20 % loss of dry matter during storage causes a 1, 6 and 11 % increase in GHG emissions per MWh. The GHG emission results are extremely sensitive to emissions of methane from the wood chip stack: If 1 % of the carbon within the stack undergoes anaerobic decomposition to methane, then the GHG emissions per MWh are tripled. There are some uncertainties in the LCA results, regarding the true formation of methane in wood chip stacks, non-CO2 emissions from combustion, N2O emissions from leaf fall and the extent of carbon sequestered under the crop, and these all contribute a large proportion of the life cycle GHG emissions from cultivation of the crop.

High yielding biomass genotypes of willow (Salix spp.) show differences in below ground biomass allocation.

Willows (Salix spp.) grown as short rotation coppice (SRC) are viewed as a sustainable source of biomass with a positive greenhouse gas (GHG) balance due to their potential to fix and accumulate carbon (C) below ground. However, exploiting this potential has been limited by the paucity of data available on below ground biomass allocation and the extent to which it varies between genotypes. Furthermore, it is likely that allocation can be altered considerably by environment. To investigate the role of genotype and environment on allocation, four willow genotypes were grown at two replicated field sites in southeast England and west Wales, UK. Above and below ground biomass was intensively measured over two two-year rotations. Significant genotypic differences in biomass allocation were identified, with below ground allocation differing by up to 10% between genotypes. Importantly, the genotype with the highest below ground biomass also had the highest above ground yield. Furthermore, leaf area was found to be a good predictor of below ground biomass. Growth environment significantly impacted allocation; the willow genotypes grown in west Wales had up to 94% more biomass below ground by the end of the second rotation. A single investigation into fine roots showed the same pattern with double the volume of fine roots present. This greater below ground allocation may be attributed primarily to higher wind speeds, plus differences in humidity and soil characteristics. These results demonstrate that the capacity exists to breed plants with both high yields and high potential for C accumulation.

X-ray micro-computed tomography in willow reveals tissue patterning of reaction wood and delay in programmed cell death.

BACKGROUND: Variation in the reaction wood (RW) response has been shown to be a principle component driving differences in lignocellulosic sugar yield from the bioenergy crop willow. The phenotypic cause(s) behind these differences in sugar yield, beyond their common elicitor, however, remain unclear. Here we use X-ray micro-computed tomography (µCT) to investigate RW-associated alterations in secondary xylem tissue patterning in three dimensions (3D). RESULTS: Major architectural alterations were successfully quantified in 3D and attributed to RW induction. Whilst the frequency of vessels was reduced in tension wood tissue (TW), the total vessel volume was significantly increased. Interestingly, a delay in programmed-cell-death (PCD) associated with TW was also clearly observed and readily quantified by µCT. CONCLUSIONS: The surprising degree to which the volume of vessels was increased illustrates the substantial xylem tissue remodelling involved in reaction wood formation. The remodelling suggests an important physiological compromise between structural and hydraulic architecture necessary for extensive alteration of biomass and helps to demonstrate the power of improving our perspective of cell and tissue architecture. The precise observation of xylem tissue development and quantification of the extent of delay in PCD provides a valuable and exciting insight into this bioenergy crop trait.

Insights into nitrogen allocation and recycling from nitrogen elemental analysis and 15N isotope labelling in 14 genotypes of willow.

Minimizing nitrogen (N) fertilization inputs during cultivation is essential for sustainable production of bioenergy and biofuels. The biomass crop willow (Salix spp.) is considered to have low N fertilizer requirements due to efficient recycling of nutrients during the perennial cycle. To investigate how successfully different willow genotypes assimilate and allocate N during growth, and remobilize and consequently recycle N before the onset of winter dormancy, N allocation and N remobilization (to and between different organs) were examined in 14 genotypes of a genetic family using elemental analysis and (15)N as a label. Cuttings were established in pots in April and sampled in June, August and at onset of senescence in October. Biomass yield of the trees correlated well with yields recorded in the field. Genotype-specific variation was observed for all traits measured and general trends spanning these sampling points were identified when trees were grouped by biomass yield. Nitrogen reserves in the cutting fuelled the entirety of the canopy establishment, yet earlier cessation of this dependency was linked to higher biomass yields. The stem was found to be the major N reserve by autumn, which constitutes a major source of N loss at harvest, typically every 2-3 years. These data contribute to understanding N remobilization in short rotation coppice willow and to the identification of traits that could potentially be selected for in breeding programmes to further improve the sustainability of biomass production.

Reaction wood - a key cause of variation in cell wall recalcitrance in willow.

BACKGROUND: The recalcitrance of lignocellulosic cell wall biomass to deconstruction varies greatly in angiosperms, yet the source of this variation remains unclear. Here, in eight genotypes of short rotation coppice willow (Salix sp.) variability of the reaction wood (RW) response and the impact of this variation on cell wall recalcitrance to enzymatic saccharification was considered. RESULTS: A pot trial was designed to test if the 'RW response' varies between willow genotypes and contributes to the differences observed in cell wall recalcitrance to enzymatic saccharification in field-grown trees. Biomass composition was measured via wet chemistry and used with glucose release yields from enzymatic saccharification to determine cell wall recalcitrance. The levels of glucose release found for pot-grown control trees showed no significant correlation with glucose release from mature field-grown trees. However, when a RW phenotype was induced in pot-grown trees, glucose release was strongly correlated with that for mature field-grown trees. Field studies revealed a 5-fold increase in glucose release from a genotype grown at a site exposed to high wind speeds (a potentially high RW inducing environment) when compared with the same genotype grown at a more sheltered site. CONCLUSIONS: Our findings provide evidence for a new concept concerning variation in the recalcitrance to enzymatic hydrolysis of the stem biomass of different, field-grown willow genotypes (and potentially other angiosperms). Specifically, that genotypic differences in the ability to produce a response to RW inducing conditions (a 'RW response') indicate that this RW response is a primary determinant of the variation observed in cell wall glucan accessibility. The identification of the importance of this RW response trait in willows, is likely to be valuable in selective breeding strategies in willow (and other angiosperm) biofuel crops and, with further work to dissect the nature of RW variation, could provide novel targets for genetic modification for improved biofuel feedstocks.

Genetic improvement of willow for bioenergy and biofuels.

Willows (Salix spp.) are a very diverse group of catkin-bearing trees and shrubs that are widely distributed across temperate regions of the globe. Some species respond well to being grown in short rotation coppice (SRC) cycles, which are much shorter than conventional forestry. Coppicing reinvigorates growth and the biomass rapidly accumulated can be used as a source of renewable carbon for bioenergy and biofuels. As SRC willows re-distribute nutrients during the perennial cycle they require only minimal nitrogen fertilizer for growth. This results in fuel chains with potentially high greenhouse gas reductions. To exploit their potential for renewable energy, willows need to be kept free of pests and diseases and yields need to be improved without significantly increasing the requirements for fertilizers and water. The biomass composition needs to be optimized for different end-uses. Yields also need to be sustainable on land less productive for food crops to reduce conflicts over land use. Advances in understanding the physiology and growth of willow, and in the identification of genes underlying key traits, are now at the stage where they can start to be used in breeding programs to help achieve these goals.

Measurement of key compositional parameters in two species of energy grass by Fourier transform infrared spectroscopy.

Two energy grass species, switch grass, a North American tuft grass, and reed canary grass, a European native, are likely to be important sources of biomass in Western Europe for the production of biorenewable energy. Matching chemical composition to conversion efficiency is a primary goal for improvement programmes and for determining the quality of biomass feed-stocks prior to use and there is a need for methods which allow cost effective characterisation of chemical composition at high rates of sample through-put. In this paper we demonstrate that nitrogen content and alkali index, parameters greatly influencing thermal conversion efficiency, can be accurately predicted in dried samples of these species grown under a range of agronomic conditions by partial least square regression of Fourier transform infrared spectra (R(2) values for plots of predicted vs. measured values of 0.938 and 0.937, respectively). We also discuss the prediction of carbon and ash content in these samples and the application of infrared based predictive methods for the breeding improvement of energy grasses.

Quantification of hydroxycinnamic acids and lignin in perennial forage and energy grasses by Fourier-transform infrared spectroscopy and partial least squares regression.

Levels of lignin and hydroxycinnamic acid wall components in three genera of forage grasses (Lolium,Festuca and Dactylis) have been accurately predicted by Fourier-transform infrared spectroscopy using partial least squares models correlated to analytical measurements. Different models were derived that predicted the concentrations of acid detergent lignin, total hydroxycinnamic acids, total ferulate monomers plus dimers, p-coumarate and ferulate dimers in independent spectral test data from methanol extracted samples of perennial forage grass with accuracies of 92.8%, 86.5%, 86.1%, 59.7% and 84.7% respectively, and analysis of model projection scores showed that the models relied generally on spectral features that are known absorptions of these compounds. Acid detergent lignin was predicted in samples of two species of energy grass, (Phalaris arundinacea and Pancium virgatum) with an accuracy of 84.5%.

Bioenergy from plants and the sustainable yield challenge.

Bioenergy from plants, particularly from perennial grasses and trees, could make a substantial contribution to alleviation of global problems in climate change and energy security if high yields can be sustained. Here, yield traits in a range of key bioenergy crops are reviewed, from which several targets for future improvement can be identified. Some are already the focus of genetically modified (GM) and non-GM approaches. However, the efficient growth strategies of perennial bioenergy crops rely on newly assimilated and recycled carbon and remobilized nitrogen in a continually shifting balance between sources and sinks. This balance is affected by biotic (e.g. pest, disease) and abiotic (e.g. drought) stresses. Future research should focus on three main challenges: changing (photo)thermal time sensitivity to lengthen the growing season without risking frost damage or limiting remobilization of nutritional elements following senescence; increasing aboveground biomass without depleting belowground reserves required for next year's growth and thus without increasing the requirement for nutrient applications; and increasing aboveground biomass without increasing water use.