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In this paper, we provide direct evidence of the importance of root hairs on pore structure development at the root–soil interface during the early stage of crop establishment. This was achieved by use of high-resolution (c. 5 μm) synchrotron radiation computed tomography (SRCT) to visualise both the structure of root hairs and the soil pore structure in plant–soil microcosms. Two contrasting genotypes of barley (Hordeum vulgare), with and without root hairs, were grown for 8d in microcosms packed with sandy loam soil at 1.2 g cm−3 dry bulk density. Root hairs were visualised within air-filled pore spaces, but not in the fine-textured soil regions. We found that the genotype with root hairs significantly altered the porosity and connectivity of the detectable pore space (> 5 μm) in the rhizosphere, as compared with the no-hair mutants. Both genotypes showed decreasing pore space between 0.8 and 0.1mm from the root surface. Interestingly the root-hair-bearing genotype had a significantly greater soil pore volume-fraction at the root–soil interface. Effects of pore structure on diffusion and permeability were estimated to be functionally insignificant under saturated conditions when simulated using image-based modelling.

Root hairs are a key trait for improving the acquisition of phosphorus (P) by plants. However, it is not known whether root hairs provide significant advantage for plant growth under combined soil stresses, particularly under conditions that are known to restrict root hair initiation or elongation (e.g. compacted or high-strength soils). To investigate this, the root growth and P uptake of root hair genotypes of barley, Hordeum vulgare L. (i.e. genotypes with and without root hairs), were assessed under combinations of P deficiency and high soil strength. Genotypes with root hairs were found to have an advantage for root penetration into high-strength layers relative to root hairless genotypes. In P-deficient soils, despite a 20% reduction in root hair length under high-strength conditions, genotypes with root hairs were also found to have an advantage for P uptake. However, in fertilized soils, root hairs conferred an advantage for P uptake in low-strength soil but not in high-strength soil. Improved root–soil contact, coupled with an increased supply of P to the root, may decrease the value of root hairs for P acquisition in high-strength, high-P soils. Nevertheless, this work demonstrates that root hairs are a valuable trait for plant growth and nutrient acquisition under combined soil stresses. Selecting plants with superior root hair traits is important for improving P uptake efficiency and hence the sustainability of agricultural systems.

There is an urgent need for simple rapid screens of root traits that improve the acquisition of nutrients and water. Temperate cereals produce rhizosheaths of variable weight, a trait first noted on desert species sampled by Tansley over 100 yr ago. This trait is almost certainly important in tolerance to abiotic stress. Here, we screened association genetics populations of barley for rhizosheath weight and derived quantitative trait loci (QTLs) and candidate genes. We assessed whether rhizosheath weight was correlated with plant performance and phosphate uptake under combined drought and phosphorus deficiency. Rhizosheath weight was investigated in relation to root hair length, and under both laboratory and field conditions. Our data demonstrated that rhizosheath weight was correlated with phosphate uptake under dry conditions and that the differences in rhizosheath weight between genotypes were maintained in the field. Rhizosheath weight also varied significantly within barley populations, was correlated with root hair length and was associated with a genetic locus (QTL) on chromosome 2H. Putative candidate genes were identified. Rhizosheath weight is easy and rapid to measure, and is associated with relatively high heritability. The breeding of cereal genotypes for beneficial rhizosheath characteristics is achievable and could contribute to agricultural sustainability in nutrient‐ and water‐stressed environments.

Micronutrient deficiency in cereals is a problem of global significance, severely reducing grain yield and quality in marginal soils. Ancient landraces represent, through hundreds of years of local adaptation to adverse soil conditions, a unique reservoir of genes and unexplored traits for enhancing yield and abiotic stress tolerance. Here we explored and compared the genetic variation in a population of Northern European barley landraces and modern elite varieties, and their tolerance to manganese (Mn) limitation. A total of 135 barley accessions were genotyped and the genetic diversity was explored using Neighbor-Joining clustering. Based on this analysis, a sub-population of genetically diverse landraces and modern elite control lines were evaluated phenotypically for their ability to cope with Mn-deficient conditions, across three different environments increasing in complexity from hydroponics through pot experiments to regional field trials. Genetically a group of Scottish barley landraces (Bere barley) were found to cluster according to their island of origin, and accessions adapted to distinct biogeographical zones with reduced soil fertility had particularly larger Mn, but also zinc (Zn) and copper (Cu) concentrations in the shoot. Strikingly, when grown in an alkaline sandy soil in the field, the locally adapted landraces demonstrated an exceptional ability to acquire and translocate Mn to developing leaves, maintain photosynthesis and generate robust grain yields, whereas modern elite varieties totally failed to complete their life cycle. Our results highlight the importance of gene pools of local adaptation and the value of ancient landrace material to identify and characterize genes that control nutrient use efficiency traits in adverse environments to raise future crop production and improve agricultural sustainability in marginal soils. We propose and discuss a model summarizing the physiological mechanisms involved in the complex trait of tolerance to Mn limitation.

Societal Impact Statement Bere is an ancient barley (Hordeum vulgare L.) that was once widely grown in northern Britain, where its ability to grow on poor soils and under challenging climatic conditions made it a valuable staple. By the end of the 20th century, Bere had largely been replaced by higher‐yielding modern varieties and only survived in cultivation on a few Scottish islands. This article reviews the recent revival of Bere, driven by its use in high‐value food and drink products and multidisciplinary research into its genetics, valuable sustainability traits and potential for developing resilient barley varieties. Summary In Britain, modern cereal varieties have mostly replaced landraces. A remarkable exception occurs on several Scottish islands where Bere, an ancient 6‐row barley (Hordeum vulgare L.), is grown as a monocrop or in mixtures. In the Outer Hebrides, the mixture is grown for animal feed, and cultivating it with traditional practices is integral to the conservation of Machair, an important coastal dune ecosystem. In Orkney, Bere is grown as a monocrop, and in situ conservation has recently been strengthened by improved agronomy and new markets for grain to produce unique foods and beverages from beremeal (flour) and malt. In parallel, a recently assembled collection of British and North European barley landraces has allowed the genotypic and phenotypic characterisation of Bere and several associated multidisciplinary studies. Genotyping demonstrated Bere's unique identity compared with most other barleys in the collection, indicating an earlier introduction to Scotland than the Norse settlement (c. 9th century AD) suggested previously. Valuable traits found in some Bere accessions include disease resistance, an early heading date (reflecting a short period from sowing to harvest), the ability to grow on marginal, high pH soils deficient in manganese and tolerance to salinity stress. These traits would have been important in the past for grain production under the region's challenging soil and Atlantic‐maritime climatic conditions. We discuss these results within the context of Bere as a genetic, heritage and commercial resource and as a future source of sustainability traits for barley improvement. Bere is an ancient barley (Hordeum vulgare L.) that was once widely grown in northern Britain, where its ability to grow on poor soils and under challenging climatic conditions made it a valuable staple. By the end of the 20th century, Bere had largely been replaced by higher‐yielding modern varieties and only survived in cultivation on a few Scottish islands. This article reviews the recent revival of Bere, driven by its use in high‐value food and drink products and multidisciplinary research into its genetics, valuable sustainability traits and potential for developing resilient barley varieties.

Background and Aims: Plant acquisition of endogenous forms of soil phosphorus (P) could reduce external P requirements in agricultural systems. This study investigated the interaction of citrate and phytase exudation in controlling the accumulation of P and depletion of soil organic P by transgenic Nicotiana tabacum plants. Methods: N. tabacum plant lines including wild-type, vector controls, transgenic plants with single-trait expression of a citrate transporter (A. thaliana frd3) or fungal phytases (phyA: A. niger, P. lycii) and crossed plant lines expressing both traits, were characterized for citrate efflux and phytase exudation. Monocultures and intercropped combinations of single-trait plants were grown in a low available P soil (12 weeks). Plant biomass, shoot P accumulation, rhizosphere soil pH and citrate-extractable-P fractions were determined. Land Equivalent Ratio and complementarity effect was determined in intercropped treatments and multiple-linearregression was used to predict shoot P accumulation based on plant exudation and soil P depletion. Results: Crossed plant lines with co-expression of citrate and phytase accumulated more shoot P than single-trait and intercropped plant treatments. Shoot P accumulation was predicted based on phytase-labile soil P, citrate efflux, and phytase activity (Rsq=0.58, P < .0001). Positive complementarity occurred between intercropped citrate-and phytase-exuding plants, with the greatest gains in shoot P occurring in plant treatments with A. niger phyA expression. Conclusions: We show for the first time that trait synergism associated with the exudation of citrate and phytase by tobacco can be linked to the improved acquisition of P and the depletion of soil organic P.

Aims The rhizosheath is defined as the weight of soil adhering strongly to roots on excavation, and current interest in this trait as a potential tolerance mechanism to abiotic stress has prompted us to explore the extent of its occurrence throughout the angiosperm phylogeny. Methods Here we describe a robust, novel method which was used to screen species for the presence/absence and strength of a rhizosheath. We correlate the latter with root hair length to provide insight into some of the factors affecting its formation. We go on to compare experimental data with previous observations in the literature. Results Results of a glasshouse screen demonstrate that rhizosheaths exist in species from many angiosperm orders, and the frequency of their occurrence and their strength and size are related. No correlation between root hair length and rhizosheath size was found, except when root hairs were extremely short, but the presence of root hairs was required for rhizosheath formation. Conclusions The rhizosheath is present in species from many angiosperm orders. Potential to enhance the trait is likely to exist in a range of crop species and could help contribute to future agricultural sustainability.

To enhance the sustainability of agriculture it is imperative that the use of P-fertilisers by temperate cereal crops be improved. This can be achieved both by agronomic and genetic approaches. While many studies have demonstrated genotypic variation in P-use efficiency in a number of cereal species the robustness of this genetic variation in contrasting environments is rarely considered. In this paper we describe an experiment in which we compare the P-nutrition of winter and spring barley genotypes from an association genetic-mapping population grown in a field trial with different cultivation treatments (conventional plough vs. minimum tillage) which had been established over a number of years. We demonstrate that, while there is significant variation between genotypes in their P nutrition, this variation is not comparable between cultivation treatments and only one winter barley genotype (cv. Gleam) has beneficial P-use efficiency traits in both cultivation systems. Analysis of the association genetic-mapping population demonstrated that there was a strong environmental component in the genotypic variation, with more significant associations of shoot P concentration with known SNP (Single Nucleotide Polymorphism) markers when the population was grown in minimum tillage treatments. These data suggest that it may be possible to identify genetic components to variation in P nutrition in barley, but that a large interaction with environmental variables may limit the usefulness of any genes or markers discovered for improving P-use efficiency to the conditions under which the screening was performed.

• Soil adjacent to roots has distinct structural and physical properties from bulk soil, affecting water and solute acquisition by plants. Detailed knowledge on how root activity and traits such as root hairs affect the three-dimensional pore structure at a fine scale is scarce and often contradictory. • Roots of hairless barley (Hordeum vulgare L. cv Optic) mutant (NRH) and its wildtype (WT) parent were grown in tubes of sieved (<250 μm) sandy loam soil under two different water regimes. The tubes were scanned by synchrotron-based X-ray computed tomography to visualise pore structure at the soil–root interface. Pore volume fraction and pore size distribution were analysed vs distance within 1 mm of the root surface. • Less dense packing of particles at the root surface was hypothesised to cause the observed increased pore volume fraction immediately next to the epidermis. The pore size distribution was narrower due to a decreased fraction of larger pores. There were no statistically significant differences in pore structure between genotypes or moisture conditions. • A model is proposed that describes the variation in porosity near roots taking into account soil compaction and the surface effect at the root surface.

Key message Characterisation and genetic mapping of a key gene defining root morphology in bread wheat. Root morphology is central to plants for the efficient uptake up of soil water and mineral nutrients. Here we describe a conditional mutant of hexaploid wheat ( Triticum aestivum L.) that when grown in soil with high Ca 2+ develops a larger rhizosheath accompanied with shorter roots than the wild type. In wheat, rhizosheath size is a reliable surrogate for root hair length and this was verified in the mutant which possessed longer root hairs than the wild type when grown in high Ca 2+ soil. We named the mutant Stumpy and showed it to be due to a single semi-dominant mutation. The short root phenotype at high Ca 2+ was due to reduced cellular elongation which might also explain the long root hair phenotype. Analysis of root cell walls showed that the polysaccharide composition of Stumpy roots is remodelled when grown at non-permissive (high) Ca 2+ concentrations. The mutation mapped to chromosome 7B and sequencing of the 7B chromosomes in both wild type and Stumpy identified a candidate gene underlying the Stumpy mutation. As part of the process to determine whether the candidate gene was causative, we identified wheat lines in a Cadenza TILLING population with large rhizosheaths but accompanied with normal root length. This finding illustrates the potential of manipulating the gene to disconnect root length from root hair length as a means of developing wheat lines with improved efficiency of nutrient and water uptake. The Stumpy mutant will be valuable for understanding the mechanisms that regulate root morphology in wheat.