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Barley development from seedling to flowering involves both external and internal changes, the latter requiring microscopic observation. Internal changes allow for the classification of preflowering development into three phases: vegetative, early reproductive, and late reproductive. Genetic and environmental factors influence the duration of these phases, impacting grain yield. Photoperiod-sensitivity genes PPD-H1 play amajor role in flowering time, affecting adaptation; however, the effect might also be direct (beyond affecting phenology). In this paper, we aimed to assess how PPD-H1 alleles affect barley development, including the progression of growth phases, leaf emergence, tillering dynamics, and spikelet development. Two experiments (field and controlled conditions) were conducted with a factorial combination of (i) four near-isogenic lines (NILs) for PPD-H1 alleles (ppd-H1 or Ppd-H1) under two contrasting PHYC genetic backgrounds (PhyC-l and PhyC-e) and (ii) two photoperiod conditions (short and long days). As expected, longer photoperiods led to a shorter growth cycle. All subphases of time to flowering, final leaf number, and phyllochron were affected by photoperiod. The effects of PPD-H1 on flowering time depended on the PHYC genetic backgrounds and photoperiod conditions. PPD-H1 effects on flowering time were associated with leaf number and phyllochron; the interplay between leaf number and phyllochron affected mainly the late reproductive phase. We also found that although PPD-H1 did not affect the phyllochron of the first six leaves, the phyllochron of leaves appearing later, when grown under a short photoperiod, was consistently increased in lines carrying the ppd-H1 allele. Tillering dynamics exhibited variability, but PPD-H1 did not affect the final spike number under a 24-h photoperiod. [ABSTRACT FROM AUTHOR]

The prevalence of drought events worldwide emphasizes the importance of screening and cultivating drought-adapted crops. In this study, 206 germplasm resources were used as materials, dry weight as target trait, and two genotyping methods as criteria to evaluate drought adaptability at the seedling establishment stage. The results showed a significant decrease in average dry weight of the tested germplasm resources (from 746.90 mg to 285.40 mg) and rich variation in the responses of dry weight among each genotype to drought (CV=61.14%). In traditional evaluation method, drought resistance coefficient (DC), geometric mean productivity index (GMP), mean productivity index (MP), stress susceptibility index (SSI), stress tolerance index (STI), and tolerance index (TOL) also exhibited diversity in tested genotypes (CV>30%). However, these indices showed varying degrees of explanation for dry weight under stress and nonstress environments and failed to differentiate drought adaptability among genotypes clearly. In new evaluation method, four stress indices were developed to quantify barley seedling production and stability capacities. Compared to traditional stress indices, the stress production index (SI) explained dry weight more comprehensively under stress conditions (R² = 0.98), while the ideal production index (II) explained dry weight better under non-stress conditions (R² = 0.89). Furthermore, the potential index (PI) and elasticity index (EI) eliminated disparities in traditional stress indices and comprehensively clarified the contribution of elasticity and potential to production capacity under drought stress. Ultimately, through grading evaluation and cluster analysis, the tested germplasm resources were effectively categorized, and 11 genotypes were identified as suitable for cultivation in arid areas. Overall, the comprehensive evaluation method based on the newly developed stress indices surpasses the traditional method in screening drought adaptability of crops and serves as a vital tool for identifying high-stability and high-production capacities genotypes in various environments, which is expected to provide practical guidance for barley planting and breeding in arid areas. [ABSTRACT FROM AUTHOR]

Barley stripe or yellowrust (BYR) caused by Puccinia striiformis f. sp. hordei (Psh) is a significant constraint to barley production. The disease is best controlled by genetic resistance, which is considered the most economical and sustainable component of integrated disease management. In this study, we assessed the diversity of resistance to Psh in a panel of international barley genotypes (n = 266) under multiple disease environments (Ecuador, India, and Mexico) using genome-wide association studies (GWASs). Four quantitative trait loci (QTLs) (three on chromosome 1H and one on 7H) associated with resistance to Psh were identified. The QTLs were validated by mapping resistance to Psh in five biparental populations, which detected key genomic regions on chromosomes 1H (populations Pompadour/Zhoungdamei, Pompadour/Zug161, and CI9214/Baudin), 3H (Ricardo/Gus), and 7H (Fumai8/Baronesse). The QTL RpshQ.GWA.1H.1 detected by GWAS and RpshQ.Bau.1H detected using biparental mapping populations co-located were the most consistent and stable across environments and are likely the same resistance region. RpshQ.Bau.1H was saturated using population CI9214/Baudin by enriching the target region, which placed the resistance locus between 7.9 and 8.1 Mbp (flanked by markers sun_B1H_03, 0.7 cM proximal to Rpsh_1H and sun_B1H_KASP_02, 3.2 cM distal on 1HS) in the Morex reference genome v.2. A Kompetitive Allele Specific PCR (KASP) marker sun_B1H_KASP_01 that cosegregated for RpshQ.Bau.1H was developed. The marker was validated on 50 Australian barley cultivars, showingwell-defined allelic discrimination and presence in six genotypes (Baudin, Fathom, Flagship, Grout, Sakurastar, and Shepherd). This marker can be used for reliable marker-assisted selection and pyramiding of resistance to Psh and in diversifying the genetic base of resistance to stripe rust. [ABSTRACT FROM AUTHOR]

In the face of escalating environmental challenges, understanding crop responses to abiotic stress is pivotal for sustainable agriculture. The present studymeticulously investigates the intricate interplay between drought and salinity stress in barley (Hordeum vulgare L.). Employing three distinct barley genotypes--Traveller, Prunella, and Zahna--we scrutinize their physiological, biochemical, and molecular adaptations under stress conditions. Our findings underscore genotype-specific responses, unravelling the multifaceted mechanisms that govern stress tolerance. Chlorophyll content, a vital indicator of photosynthetic efficiency, exhibits significant variations across genotypes. Salinity stress induces a decline in chlorophyll levels, while drought stress triggers a more nuanced response. Stomatal conductance, a key regulator of water loss, also diverges among the genotypes. Traveller displays remarkable stomatal closure under drought, conserving water, whereas Prunella and Zahna exhibit contrasting patterns. Antioxidant enzyme activities, crucial for combating oxidative stress, fluctuate significantly. Activities of superoxide dismutase (SOD) and catalase (CAT) surge under salinity stress, while drought predominantly impacts SOD. Gene expression profiling reveals genotype-specific signatures, with stress-responsive genes modulating adaptive pathways. Correlation analyses revealed the intricate interplay of the physiological and biochemical parameters. Genotype-specific adaptations, coupled with dynamic physiological and molecular responses, underscore the plasticity of barley's stress tolerance mechanisms. Throughout the study, the Zahna genotype demonstrated notable tolerance in terms of performance. These insights hold promise for breeding resilient cultivars, bolstering food security in an increasingly unpredictable climate. By deciphering the barley stress symphony, we contribute to the harmonious orchestration of sustainable agricultural practices. [ABSTRACT FROM AUTHOR]

Three-amino-loop-extension (TALE) family belongs to the homeobox gene superfamily and occurs widely in plants, playing a crucial role in regulating their growth and development. Currently, genome-wide analysis of the TALE family has been completed in many plants. However, the systematic identification and hormone response analysis of the TALE gene family in barley are still lacking. In this study, 21 TALE candidate genes were identified in barley, which can be divided into KNOX and BELL subfamilies. Barley TALE members in the same subfamily of the phylogenetic tree have analogically conserved motifs and gene structures, and segmental duplications are largely responsible for the expansion of the HvTALE family. Analysis of TALE orthologous and homologous gene pairs indicated that the HvTALE family has mainly undergone purifying selective pressure. Through spatial structure simulation, HvKNOX5–HvKNOX6 and HvKNOX5–HvBELL11 complexes are all formed through hydrogen bonding sites on both the KNOX2 and homeodomain (HD) domains of HvKNOX5, which may be essential for protein interactions among the HvTALE family members. Expression pattern analyses reveal the potential involvement of most HvTALE genes in responses to exogenous hormones. These results will lay the foundation for regulation and function analyses of the barley TALE gene family in plant growth and development by hormone regulation. [ABSTRACT FROM AUTHOR]

Plants exhibit an array of drought responses and adaptations, where the trade-off between water loss and CO2 uptake for growth is mediated by regulation of stomatal aperture in response to soil water content (SWC), among other factors. For crop yield stability, the question is how drought timing and response patterns relate to post-drought growth resilience and vigor. We earlier identified, in a few reference varieties of barley that differed by the SWC at which transpiration was curtailed, two divergent water use strategies: water-saving ("isohydric") and water-spending ("anisohydric"). We proposed that an isohydric strategy may reduce risk from spring droughts in climates where the probability of precipitation increases during the growing season, whereas the anisohydric is consistent with environments having terminal droughts, or with those where dry periods are short and not seasonally progressive. Here, we have examined drought response physiology in an 81-line barley (Hordeum vulgare L.) diversity set that spans 20th century European breeding and identified several lines with a third, dynamic strategy. We found a strong positive correlation between vigor and transpiration, the dynamic group being highest for both. However, these lines curtailed daily transpiration at a higher SWC than the isohydric group. While the dynamic lines, particularly cv Hydrogen and Baronesse, were not the most resilient in terms of restoring initial growth rates, their strong initial vigor and high return to initial transpiration rates meant that their growth nevertheless surpassed more resilient lines during recovery from drought. The results will be of use for defining barley physiological ideotypes suited to future climate scenarios. [ABSTRACT FROM AUTHOR]

Many traits play essential roles in determining crop yield. Wide variation for morphological traits exists in Hordeum vulgare L., but the genetic basis of this morphological variation is largely unknown. To understand genetic basis controlling morphological traits affecting yield, a barley doubled haploid population (146 individuals) derived from Clipper X Sahara 3771 was used to map chromosome regions underlying days to awn appearance, plant height, fertile spike number, flag leaf length, spike length, harvest index, seed number per plant, thousands kernel weight, and grain yield. Twentyseven QTLs for nine traits were mapped to the barley genome that described 3-69% of phenotypic variations; and some genomic regions harbor a given QTL for more than one trait. Out of 27 QTLs identified, 19 QTLs were novel. Chromosomal regions on 1H, 2H, 4H, and 6H associated with seed grain yield, and chromosome regions on 2H and 6H had major effects on grain yield (GY). One major QTL for seed number per plant was flanked by marker VRS1-KSUF15 on chromosome 2H. This QTL was also associated with GY. Some loci controlling thousands kernel weight (TKW), fertile spike number (FSN), and GY were the same. The major grain yield QTL detected on linkage PSR167 co-localized with TAM10. Two major QTLs controlling TKW and FSN were also mapped at this locus. Eight QTLs on chromosomes 1H, 2H, 3H, 4H, 5H, 6H, and 7H consistently affected spike characteristics. One major QTL (ANIONT1A-TACMD) on 4H affected both spike length (SL) and spike number explained 9 and 5% of the variation of SL and FSN, respectively. In conclusion, this study could cast some light on the genetic basis of the studied pivotal traits. Moreover, fine mapping of the identified major effect markers may facilitate the application of molecular markers in barley breeding programs. [ABSTRACT FROM AUTHOR]

Salinity has become a major environmental concern of agricultural lands, impairing crop production. The current study aimed to examine the role of zinc oxide nanoparticles (ZnO NPs) in reducing the oxidative stress induced by salinity and the overall improvement in phytochemical properties in barley. A total of nine different barley genotypes were first subjected to salt (NaCl) stress in hydroponic conditions to determine the tolerance among the genotypes. The genotype Annora was found as most sensitive, and the most tolerant genotype was Awaran 02 under salinity stress. In another study, the most sensitive (Annora) and tolerant (Awaran 02) barley genotypes were grown in pots under salinity stress (100 mM). At the same time, half of the pots were provided with the soil application of ZnO NPs (100 mg kg−1 ), and the other half pots were foliar sprayed with ZnO NPs (100 mg L−1 ). Salinity stress reduced barley growth in both genotypes compared to control plants. However, greater reduction in barley growth was found in Annora (sensitive genotype) than in Awaran 02 (tolerant genotype). The exogenous application of ZnO NPs ameliorated salt stress and improved barley biomass, photosynthesis, and antioxidant enzyme activities by reducing oxidative damage caused by salt stress. However, this positive effect by ZnO NPs was observed more in Awaran 02 than in Annora genotype. Furthermore, the foliar application of ZnO NPs was more effective than the soil application of ZnO NPs. Findings of the present study revealed that exogenous application of ZnO NPs could be a promising approach to alleviate salt stress in barley genotypes with different levels of salinity tolerance [ABSTRACT FROM AUTHOR]

Introduction: In the Near East and North Africa (NENA) region, crop production is being affected by various abiotic factors, including freshwater scarcity, climate, and soil salinity. As a result, farmers in this region are in search of salt-tolerant crops that can thrive in these harsh environments, using poor-quality groundwater. The main staple food crop for most of the countries in this region, Tunisia included, is barley. Methods: The present study was designed to investigate the sensitivity and tolerance of six distinct barley genotypes to aridity and salinity stresses in five different natural field environments by measuring their photosynthetic activity. Results and discussion: The results revealed that tolerant genotypes were significantly less affected by these stress factors than sensitive genotypes. The genotypes that were more susceptible to salinity and aridity stress exhibited a significant decline in their photosynthetic activity. Additionally, the fluorescence yields in growth phases J, I, and P declined significantly in the order of humid environment (BEJ), semi-arid site (KAI), and arid environment (MED) and became more significant when salt stress was added through the use of saline water for irrigation. The stress adversely affected the quantum yield of primary photochemistry (jP0), the quantum yield of electron transport (jE0), and the efficiency by trapped excitation (y0) in the vulnerable barley genotypes. Moreover, the performance index (PI) of the photosystem II (PSII) was found to be the most distinguishing parameter among the genotypes tested. The PI of sensitive genotypes was adversely affected by aridity and salinity. The PI of ICARDA20 and Konouz decreased by approximately 18% and 33%, respectively, when irrigated with non-saline water. The reduction was even greater, reaching 39%, for both genotypes when irrigated with saline water. However, tolerant genotypes Souihli and Batini 100/1B were less impacted by these stress factors. The fluorescence study provided insights into the photosynthetic apparatus of barley genotypes under stress. It enabled reliable salinity tolerance screening. Furthermore, the study confirmed that the chlorophyll a fluorescence induction curve had an inflection point (step K) even before the onset of visible signs of stress, indicating physiological disturbances, making chlorophyll fluorescence an effective tool for identifying salinity tolerance in barley. [ABSTRACT FROM AUTHOR]

Miniature inverted-repeat transposable elements (MITEs) constitute a class of class II transposable elements (TEs) that are abundant in plant genomes, playing a crucial role in their evolution and diversity. Barley (Hordeum vulgare), the fourth-most important cereal crop globally, is widely used for brewing, animal feed, and human consumption. However, despite their significance, the mechanisms underlying the insertion or amplification of MITEs and their contributions to barley genome evolution and diversity remain poorly understood. Through our comprehensive analysis, we identified 32,258 full-length MITEs belonging to 2,992 distinct families, accounting for approximately 0.17% of the barley genome. These MITE families can be grouped into four well-known superfamilies (Tc1/Mariner-like, PIF/Harbinger-like, hAT-like, and Mutator-like) and one unidentified superfamily. Notably, we observed two major expansion events in the barley MITE population, occurring approximately 12-13 million years ago (Mya) and 2-3 Mya. Our investigation revealed a strong preference of MITEs for gene-related regions, particularly in promoters, suggesting their potential involvement in regulating host gene expression. Additionally, we discovered that 7.73% miRNAs are derived from MITEs, thereby influencing the origin of certain miRNAs and potentially exerting a significant impact on post-transcriptional gene expression control. Evolutionary analysis demonstrated that MITEs exhibit lower conservation compared to genes, consistent with their dynamic mobility. We also identified a series of MITE insertions or deletions associated with domestication, highlighting these regions as promising targets for crop improvement strategies. These findings significantly advance our understanding of the fundamental characteristics and evolutionary patterns of MITEs in the barley genome. Moreover, they contribute to our knowledge of gene regulatory networks and provide valuable insights for crop improvement endeavors.

Introduction: Ramularia leaf spot (RLS) disease is a growing threat to barley cultivation, but with no substantial resistance identified to date. Similarly, the understanding of the lifestyle of Ramularia collo-cygni (Rcc) and the prediction of RLS outbreak severity remain challenging, with Rcc displaying a rather untypical long endophytic phase and a sudden change to a necrotrophic lifestyle. The aim of this study was to provide further insights into the defense dynamics during the different stages of colonization and infection in barley in order to identify potential targets for resistance breeding. Methods: Utilizing the strength of proteomics in understanding plant-pathogen interactions, we performed an integrative analysis of a published transcriptome dataset with a parallel generated proteome dataset. Therefore, we included two spring barley cultivars with contrasting susceptibilities to Rcc and two fungal isolates causing different levels of RLS symptoms. Results: Interestingly, early responses in the pathogen recognition phase of the host were driven by strong responses differing between isolates. An important enzyme in this process is a xylanase inhibitor, which protected the plant from cell wall degradation by the fungal xylanase. At later time points, the differences were driven by cultivar-specific responses, affecting mostly features contributing to the pathogenesis- and senescence-related pathways or photosynthesis. Discussion: This supports the hypothesis of a hemibiotrophic lifestyle of Rcc, with slight differences in trophism of the two analyzed isolates. The integration of these data modalities highlights a strength of protein-level analysis in understanding plant-pathogen interactions and reveals new features involved in fungal recognition and susceptibility in barley cultivars. [ABSTRACT FROM AUTHOR]

In the last century, breeding programs have traditionally favoured yield-related traits, grown under high-input conditions, resulting in a loss of genetic diversity and an increased susceptibility to stresses in crops. Thus, exploiting understudied genetic resources, that potentially harbour tolerance genes, is vital for sustainable agriculture. Northern European barley germplasm has been relatively understudied despite its key role within the malting industry. The European Heritage Barley collection (ExHIBiT) was assembled to explore the genetic diversity in European barley focusing on Northern European accessions and further address environmental pressures. ExHIBiT consists of 363 springbarley accessions, focusing on two-row type. The collection consists of landraces (~14%), old cultivars (~18%), elite cultivars (~67%) and accessions with unknown breeding history (~1%), with 70% of the collection from Northern Europe. The population structure of the ExHIBiT collection was subdivided into three main clusters primarily based on the accession's year of release using 26,585 informative SNPs based on 50k iSelect single nucleotide polymorphism (SNP) array data. Power analysis established a representative core collection of 230 genotypically and phenotypically diverse accessions. The effectiveness of this core collection for conducting statistical and association analysis was explored by undertaking genome-wide association studies (GWAS) using 24,876 SNPs for nine phenotypic traits, four of which were associated with SNPs. Genomic regions overlapping with previously characterised flowering genes (HvZTLb) were identified, demonstrating the utility of the ExHIBiT core collection for locating genetic regions that determine important traits. Overall, the ExHIBiT core collection represents the high level of untapped diversity within Northern European barley, providing a powerful resource for researchers and breeders to address future climate scenarios. [ABSTRACT FROM AUTHOR]

In a pot experiment, we investigated synergistic interaction of N and P fertilisation on barley biomass (Hordeum vulgare) on both shoot and root level with the aim to determine whether N-P interaction would be the same for all levels of N and P fertilisation. We further aimed to determine whether there was a critical level of N and/or P fertilisation rate, above which, a decrease in resource allocation to roots (as nutrient availability increased) could be demonstrated. Barley plants were grown from seed on a nutrient poor substrate and subjected to a two-way NxP fertilisation gradient using a modified Hoagland fertilisation solution. We observed N-P interactions in shoot and root biomass, and N and P useefficiencies. A synergistic response in biomass was observed only above a critical level of P fertilisation when P was not limiting growth. Furthermore, we found that the same incremental increase in N:P ratio of applied fertiliser elicited different responses in shoot and root biomass depending on P treatment and concluded that barley plants were less able to cope with increasing stoichiometric imbalance when P was deficient. We provide, for the first time, stoichiometric evidence that critical levels for synergistic interactions between N-P may exist in crop plants. [ABSTRACT FROM AUTHOR]

Introduction: Induced modification of plant gene expression is of both fundamental and applied importance. Cis-acting regulatory elements (CREs) are major determinants of the spatiotemporal strength of gene expression. Yet, there are few examples where induced genetic variation in predetermined CREs has been exploited to improve or investigate crop plants. Methods: The digital PCR based FIND-IT technology was applied to discover barley mutants with CRE variants in the promoter of the nutritional important barley grain phytase (PAPhy_a) gene. Results and discussion: Mutants with higher or lower gene expression and ultimately higher or lower mature grain phytase activity (MGPA), respectively, were discovered. Field trials and inositol phosphate profiling during germination showed that PAPhy_a does not influence agronomic performance under the trial conditions but it does shorten the lag time of phosphate mobilization during germination. Higher endogenous MGPA is an improvement of grain quality for feed use as it improves the phosphate bioavailability for monogastric animals. Moreover, as the targeted CRE motifs of the PAPhy_a promoter are shared with a range of seed expressed genes like key cereal and legume storage genes, the current results demonstrates a concept for modulating individual gene expression levels of a range of seed genes. [ABSTRACT FROM AUTHOR]

Barley (Hordeum vulgare L.) is used in malt production for brewing applications. Barley malting involves a process of controlled germination that modifies the grain by activating enzymes to solubilize starch and proteins for brewing. Initially, the grain is submerged in water to raise grain moisture, requiring large volumes of water. Achieving grain modification at reduced moisture levels can contribute to the sustainability of malting practices. This study combined proteomics, bioinformatics, and biochemical phenotypic analysis of two malting barley genotypes with observed differences in water uptake and modification efficiency. We sought to reveal the molecular mechanisms at play during controlled germination and explore the roles of protein groups at 24 h intervals across the first 72 h. Overall, 3,485 protein groups were identified with 793 significant differentially abundant (DAP) within and between genotypes, involved in various biological processes, including protein synthesis, carbohydrate metabolism, and hydrolysis. Functional integration into metabolic pathways, such as glycolysis, pyruvate, starch and sucrose metabolism, revealed a metabolic rerouting due to low oxygen enforced by submergence during controlled germination. This SWATH-MS study provides a comprehensive proteome reference, delivering new insights into the molecular mechanisms underlying the impacts of low oxygen during controlled germination. It is concluded that continued efficient modification of malting barley subjected to submergence is largely due to the capacity to reroute energy to maintain vital processes, particularly protein synthesis. [ABSTRACT FROM AUTHOR]

Barley stripe or yellow rust (BYR) caused by Puccinia striiformis f. sp. hordei (Psh) is a significant constraint to barley production. The disease is best controlled by genetic resistance, which is considered the most economical and sustainable component of integrated disease management. In this study, we assessed the diversity of resistance to Psh in a panel of international barley genotypes (n = 266) under multiple disease environments (Ecuador, India, and Mexico) using genome-wide association studies (GWASs). Four quantitative trait loci (QTLs) (three on chromosome 1H and one on 7H) associated with resistance to Psh were identified. The QTLs were validated by mapping resistance to Psh in five biparental populations, which detected key genomic regions on chromosomes 1H (populations Pompadour/Zhoungdamei, Pompadour/Zug161, and CI9214/Baudin), 3H (Ricardo/Gus), and 7H (Fumai8/Baronesse). The QTL RpshQ.GWA.1H.1 detected by GWAS and RpshQ.Bau.1H detected using biparental mapping populations co-located were the most consistent and stable across environments and are likely the same resistance region. RpshQ.Bau.1H was saturated using population CI9214/Baudin by enriching the target region, which placed the resistance locus between 7.9 and 8.1 Mbp (flanked by markers sun_B1H_03, 0.7 cM proximal to Rpsh_1H and sun_B1H_KASP_02, 3.2 cM distal on 1HS) in the Morex reference genome v.2. A Kompetitive Allele Specific PCR (KASP) marker sun_B1H_KASP_01 that co-segregated for RpshQ.Bau.1H was developed. The marker was validated on 50 Australian barley cultivars, showing well-defined allelic discrimination and presence in six genotypes (Baudin, Fathom, Flagship, Grout, Sakurastar, and Shepherd). This marker can be used for reliable marker-assisted selection and pyramiding of resistance to Psh and in diversifying the genetic base of resistance to stripe rust.

In the face of escalating environmental challenges, understanding crop responses to abiotic stress is pivotal for sustainable agriculture. The present study meticulously investigates the intricate interplay between drought and salinity stress in barley (Hordeum vulgare L.). Employing three distinct barley genotypes—Traveller, Prunella, and Zahna—we scrutinize their physiological, biochemical, and molecular adaptations under stress conditions. Our findings underscore genotype-specific responses, unravelling the multifaceted mechanisms that govern stress tolerance. Chlorophyll content, a vital indicator of photosynthetic efficiency, exhibits significant variations across genotypes. Salinity stress induces a decline in chlorophyll levels, while drought stress triggers a more nuanced response. Stomatal conductance, a key regulator of water loss, also diverges among the genotypes. Traveller displays remarkable stomatal closure under drought, conserving water, whereas Prunella and Zahna exhibit contrasting patterns. Antioxidant enzyme activities, crucial for combating oxidative stress, fluctuate significantly. Activities of superoxide dismutase (SOD) and catalase (CAT) surge under salinity stress, while drought predominantly impacts SOD. Gene expression profiling reveals genotype-specific signatures, with stress-responsive genes modulating adaptive pathways. Correlation analyses revealed the intricate interplay of the physiological and biochemical parameters. Genotype-specific adaptations, coupled with dynamic physiological and molecular responses, underscore the plasticity of barley’s stress tolerance mechanisms. Throughout the study, the Zahna genotype demonstrated notable tolerance in terms of performance. These insights hold promise for breeding resilient cultivars, bolstering food security in an increasingly unpredictable climate. By deciphering the barley stress symphony, we contribute to the harmonious orchestration of sustainable agricultural practices.

Plants exhibit an array of drought responses and adaptations, where the trade-off between water loss and CO2 uptake for growth is mediated by regulation of stomatal aperture in response to soil water content (SWC), among other factors. For crop yield stability, the question is how drought timing and response patterns relate to post-drought growth resilience and vigor. We earlier identified, in a few reference varieties of barley that differed by the SWC at which transpiration was curtailed, two divergent water use strategies: water-saving (“isohydric”) and water-spending (“anisohydric”). We proposed that an isohydric strategy may reduce risk from spring droughts in climates where the probability of precipitation increases during the growing season, whereas the anisohydric is consistent with environments having terminal droughts, or with those where dry periods are short and not seasonally progressive. Here, we have examined drought response physiology in an 81-line barley (Hordeum vulgare L.) diversity set that spans 20th century European breeding and identified several lines with a third, dynamic strategy. We found a strong positive correlation between vigor and transpiration, the dynamic group being highest for both. However, these lines curtailed daily transpiration at a higher SWC than the isohydric group. While the dynamic lines, particularly cv Hydrogen and Baronesse, were not the most resilient in terms of restoring initial growth rates, their strong initial vigor and high return to initial transpiration rates meant that their growth nevertheless surpassed more resilient lines during recovery from drought. The results will be of use for defining barley physiological ideotypes suited to future climate scenarios.

The comeback of wheat stem rust in Europe, caused by Puccinia graminis f. sp. tritici, and the prevalence of the alternate (sexual) host in local areas have recently regained attention as a potential threat to European wheat production. The aim of this study was to investigate a potential epidemiological link between the aecia found on an indigenous barberry species and stem rust infections on nearby cereals and grasses. Aecial infections collected from Berberis vulgaris subsp. seroi were inoculated on a panel of susceptible genotypes of major cereal crop species. In total, 67 stem rust progeny isolates were recovered from wheat (51), barley (7), and rye (9), but none from oat, indicating the potential of barberry derived isolates to infect multiple cereals. Molecular genotyping of the progeny isolates and 20 cereal and grass stem rust samples collected at the same locations and year, revealed a clear genetic relatedness between the progeny isolated from barberry and the stem rust infections found on nearby cereal and grass hosts. Analysis of Molecular Variance indicated that variation between the stem rust populations accounted for only 1%. A Principal Components Analysis using the 62 detected multilocus genotypes also demonstrated a low degree of genetic variation among isolates belonging to the two stem rust populations. Lastly, pairwise comparisons based on fixation index (Fst), Nei's genetic distances and number of effective migrants (Nm) revealed low genetic differentiation and high genetic exchange between the two populations. Our results demonstrated a direct epidemiological link and functionality of an indigenous barberry species as the sexual host of P. graminis in Spain, a factor that should be considered when designing future strategies to prevent stem rust in Europe and beyond. [ABSTRACT FROM AUTHOR]

Improving barley grain quality is a major goal in barley breeding. In this study, a total of 35 papers focusing on quantitative trait loci (QTLs) mapping for barley quality traits published since 2000 were collected. Among the 454 QTLs identified in these studies, 349 of them were mapped onto high-density consensus maps, which were used for QTL meta-analysis. Through QTL metaanalysis, the initial QTLs were integrated into 41 meta-QTLs (MQTLs) with an average confidence interval (CI) of 1. 66 cM, which is 88.9% narrower than that of the initial QTLs. Among the 41 identified MQTLs, 25 were subsequently validated in publications using genome-wide association study (GWAS). From these 25 validated MQTLs, ten breeder’s MQTLs were selected. Synteny analysis comparing barley and wheat MQTLs revealed orthologous relationships between eight breeder’s MQTLs and 45 wheat MQTLs. Additionally, 17 barley homologs associated with rice quality traits were identified within the regions of the breeder’s MQTLs through comparative analysis. The findings of this study provide valuable insights for molecular marker-assisted breeding and the identification of candidate genes related to quality traits in barley. [ABSTRACT FROM AUTHOR]

Polyphenol oxidase (PPO) is an oxidoreductase. In damaged plant tissues, it catalyzes enzymatic browning by oxidizing o-diphenols to highly reactive oquinones, which polymerize producing heterogeneous dark polymer melanin. In intact tissues, functions of PPO are not well understood. The aim of the study was to investigate the barley PPO gene family and to reveal the possible involvement of Ppo genes in melanization of barley grain, which is controlled by the Blp1 gene. Based on known barley Ppo genes on chromosome 2H (Ppo1 and Ppo2), two additional genes--Ppo3 and Ppo4--were found on chromosomes 3H and 4H, respectively. These genes have one and two exons, respectively, contain a conserved tyrosinase domain and are thought to be functional. Comparative transcriptional analyzes of the genes in samples of developing grains (combined hulls and pericarp tissues) were conducted in two barley lines differing by melanin pigmentation. The genes were found to be transcribed with increasing intensity (while grains mature) independently from the grain color, except for Ppo2, which is transcribed only in black-grained line i:BwBlp1 accumulating melanin in grains. Analysis of this gene's expression in detached hulls and pericarps showed its elevated transcription in both tissues in comparison with yellow ones, while it was significantly higher in hulls than in pericarp. Segregation analysis in two F2 populations obtained based on barley genotypes carrying dominant Blp1 and recessive ppo1 (I) and dominant Blp1 and recessive ppo1 and ppo2 (II) was carried out. In population I, only two phenotypic classes corresponding to parental black and white ones were observed; the segregation ratio was 3 black to 1 white, corresponding to monogenic. In population II, aside from descendants with black and white grains, hybrids with a gray phenotype -- light hulls and dark pericarp -- were observed; the segregation ratio was 9 black to 3 gray to 4 white, corresponding to the epistatic interaction of two genes. Most hybrids with the gray phenotype carry dominant Blp1 and a homozygous recessive allele of Ppo2. Based on transcription and segregation assays one may conclude involvement of Ppo2 but not Ppo1 in melanin formation in barley hulls. [ABSTRACT FROM AUTHOR]

Barley is an important crop worldwide known for its adaptation to harsh environments and used in multiple forms as feed, food and beverages. Its productivity is affected by major abiotic and biotic stresses. Scald caused by hemibiotrophic fungus Rhynchosporium commune is a major foliar disease in many parts of the world. Host plant resistance is targeted by breeders to efficiently control this disease. An association mapping panel of 316 spring barley genotypes (AM2017) was screened for seedling resistance in greenhouse against three R. commune isolates and for adult plant resistance in three field locations in Morocco. The phenotyping results showed different numbers of entries with resistant and moderately resistant reactions at both seedling and adult plant stages. The reactions differed between the isolates with the highest percentage of resistant genotypes observed for isolate SC-S611 (49.4%) and highest percentage of susceptible genotypes (73.8%) for isolate SC1122. At adult plant stage, the highest percentage of scald resistant genotypes (64.5%) was observed at Rommani site compared to 56% at Guich site and only 28.8% at Marchouch site. Seven genotypes were resistant at the seedling and adult plant stages. Genome wide association study (GWAS) revealed 102 MTA (15 QTL) at the seedling stage, and 25 MTA (12 QTL) associated with scald resistance at the adult plant stage. In addition, the sequences of 92 out of 102 at SRT, and 24 out of 25 significant SNP markers at APR were located in genomic regions enriched with functional proteins involved in diverse cellular processes including disease resistance. These markers span over all chromosomes with the majority of SNPs located on 3H and 7H. This study has verified 18 QTL reported in previous studies. In addition, it was successful in identifying new sources of resistance and novel genomic regions which could help in enhancing scald resistance in barley breeding programs. [ABSTRACT FROM AUTHOR]

IntroductionIn the Near East and North Africa (NENA) region, crop production is being affected by various abiotic factors, including freshwater scarcity, climate, and soil salinity. As a result, farmers in this region are in search of salt-tolerant crops that can thrive in these harsh environments, using poor-quality groundwater. The main staple food crop for most of the countries in this region, Tunisia included, is barley.MethodsThe present study was designed to investigate the sensitivity and tolerance of six distinct barley genotypes to aridity and salinity stresses in five different natural field environments by measuring their photosynthetic activity.Results and discussionThe results revealed that tolerant genotypes were significantly less affected by these stress factors than sensitive genotypes. The genotypes that were more susceptible to salinity and aridity stress exhibited a significant decline in their photosynthetic activity. Additionally, the fluorescence yields in growth phases J, I, and P declined significantly in the order of humid environment (BEJ), semi-arid site (KAI), and arid environment (MED) and became more significant when salt stress was added through the use of saline water for irrigation. The stress adversely affected the quantum yield of primary photochemistry (φP0), the quantum yield of electron transport (φE0), and the efficiency by trapped excitation (ψ0) in the vulnerable barley genotypes. Moreover, the performance index (PI) of the photosystem II (PSII) was found to be the most distinguishing parameter among the genotypes tested. The PI of sensitive genotypes was adversely affected by aridity and salinity. The PI of ICARDA20 and Konouz decreased by approximately 18% and 33%, respectively, when irrigated with non-saline water. The reduction was even greater, reaching 39%, for both genotypes when irrigated with saline water. However, tolerant genotypes Souihli and Batini 100/1B were less impacted by these stress factors.The fluorescence study provided insights into the photosynthetic apparatus of barley genotypes under stress. It enabled reliable salinity tolerance screening. Furthermore, the study confirmed that the chlorophyll a fluorescence induction curve had an inflection point (step K) even before the onset of visible signs of stress, indicating physiological disturbances, making chlorophyll fluorescence an effective tool for identifying salinity tolerance in barley.

In the last century, breeding programs have traditionally favoured yield-related traits, grown under high-input conditions, resulting in a loss of genetic diversity and an increased susceptibility to stresses in crops. Thus, exploiting understudied genetic resources, that potentially harbour tolerance genes, is vital for sustainable agriculture. Northern European barley germplasm has been relatively understudied despite its key role within the malting industry. The European Heritage Barley collection (ExHIBiT) was assembled to explore the genetic diversity in European barley focusing on Northern European accessions and further address environmental pressures. ExHIBiT consists of 363 spring-barley accessions, focusing on two-row type. The collection consists of landraces (~14%), old cultivars (~18%), elite cultivars (~67%) and accessions with unknown breeding history (~1%), with 70% of the collection from Northern Europe. The population structure of the ExHIBiT collection was subdivided into three main clusters primarily based on the accession’s year of release using 26,585 informative SNPs based on 50k iSelect single nucleotide polymorphism (SNP) array data. Power analysis established a representative core collection of 230 genotypically and phenotypically diverse accessions. The effectiveness of this core collection for conducting statistical and association analysis was explored by undertaking genome-wide association studies (GWAS) using 24,876 SNPs for nine phenotypic traits, four of which were associated with SNPs. Genomic regions overlapping with previously characterised flowering genes (HvZTLb) were identified, demonstrating the utility of the ExHIBiT core collection for locating genetic regions that determine important traits. Overall, the ExHIBiT core collection represents the high level of untapped diversity within Northern European barley, providing a powerful resource for researchers and breeders to address future climate scenarios.

In a pot experiment, we investigated synergistic interaction of N and P fertilisation on barley biomass (Hordeum vulgare) on both shoot and root level with the aim to determine whether N-P interaction would be the same for all levels of N and P fertilisation. We further aimed to determine whether there was a critical level of N and/or P fertilisation rate, above which, a decrease in resource allocation to roots (as nutrient availability increased) could be demonstrated. Barley plants were grown from seed on a nutrient poor substrate and subjected to a two-way NxP fertilisation gradient using a modified Hoagland fertilisation solution. We observed N-P interactions in shoot and root biomass, and N and P use-efficiencies. A synergistic response in biomass was observed only above a critical level of P fertilisation when P was not limiting growth. Furthermore, we found that the same incremental increase in N:P ratio of applied fertiliser elicited different responses in shoot and root biomass depending on P treatment and concluded that barley plants were less able to cope with increasing stoichiometric imbalance when P was deficient. We provide, for the first time, stoichiometric evidence that critical levels for synergistic interactions between N-P may exist in crop plants.

IntroductionInduced modification of plant gene expression is of both fundamental and applied importance. Cis-acting regulatory elements (CREs) are major determinants of the spatiotemporal strength of gene expression. Yet, there are few examples where induced genetic variation in predetermined CREs has been exploited to improve or investigate crop plants. MethodsThe digital PCR based FIND-IT technology was applied to discover barley mutants with CRE variants in the promoter of the nutritional important barley grain phytase (PAPhy_a) gene. Results and discussionMutants with higher or lower gene expression and ultimately higher or lower mature grain phytase activity (MGPA), respectively, were discovered. Field trials and inositol phosphate profiling during germination showed that PAPhy_a does not influence agronomic performance under the trial conditions but it does shorten the lag time of phosphate mobilization during germination. Higher endogenous MGPA is an improvement of grain quality for feed use as it improves the phosphate bioavailability for monogastric animals. Moreover, as the targeted CRE motifs of the PAPhy_a promoter are shared with a range of seed expressed genes like key cereal and legume storage genes, the current results demonstrates a concept for modulating individual gene expression levels of a range of seed genes.

Barley (Hordeum vulgare L.) is used in malt production for brewing applications. Barley malting involves a process of controlled germination that modifies the grain by activating enzymes to solubilize starch and proteins for brewing. Initially, the grain is submerged in water to raise grain moisture, requiring large volumes of water. Achieving grain modification at reduced moisture levels can contribute to the sustainability of malting practices. This study combined proteomics, bioinformatics, and biochemical phenotypic analysis of two malting barley genotypes with observed differences in water uptake and modification efficiency. We sought to reveal the molecular mechanisms at play during controlled germination and explore the roles of protein groups at 24 h intervals across the first 72 h. Overall, 3,485 protein groups were identified with 793 significant differentially abundant (DAP) within and between genotypes, involved in various biological processes, including protein synthesis, carbohydrate metabolism, and hydrolysis. Functional integration into metabolic pathways, such as glycolysis, pyruvate, starch and sucrose metabolism, revealed a metabolic rerouting due to low oxygen enforced by submergence during controlled germination. This SWATH-MS study provides a comprehensive proteome reference, delivering new insights into the molecular mechanisms underlying the impacts of low oxygen during controlled germination. It is concluded that continued efficient modification of malting barley subjected to submergence is largely due to the capacity to reroute energy to maintain vital processes, particularly protein synthesis.

Introduction: Breeding barley cultivars adapted to drought requires in-depth knowledge on physiological drought responses. Methods: We used a high-throughput functional phenotyping platform to examine the response of four high-yielding European spring barley cultivars to a standardized drought treatment imposed around flowering. Results: Cv. Chanell showed a non-conserving water-use behavior with high transpiration and maximum productivity under well-watered conditions but rapid transpiration decrease under drought. The poor recovery upon reirrigation translated to large yield losses. Cv. Baronesse showed the most water-conserving behavior, with the lowest pre-drought transpiration and the most gradual transpiration reduction under drought. Its good recovery (resilience) prevented large yield losses. Cv. Formula was less conserving than cv. Baronesse and produced low yet stable yields. Cv. RGT's dynamic water use with high transpiration under ample water supply and moderate transpiration decrease under drought combined with high resilience secured the highest and most stable yields. Discussion: Such a dynamic water-use behavior combined with higher drought resilience and favorable root traits could potentially create an ideotype for intermediate drought. Prospective studies will examine these results in field experiments and will use the newly gained understanding on water use in barley to improve process descriptions in crop simulation models to support crop model-aided ideotype design. [ABSTRACT FROM AUTHOR]

Introduction: Breeding barley cultivars adapted to drought requires in-depth knowledge on physiological drought responses. Methods: We used a high-throughput functional phenotyping platform to examine the response of four high-yielding European spring barley cultivars to a standardized drought treatment imposed around flowering. Results: Cv. Chanell showed a non-conserving water-use behavior with high transpiration and maximum productivity under well-watered conditions but rapid transpiration decrease under drought. The poor recovery upon reirrigation translated to large yield losses. Cv. Baronesse showed the most water-conserving behavior, with the lowest pre-drought transpiration and the most gradual transpiration reduction under drought. Its good recovery (resilience) prevented large yield losses. Cv. Formula was less conserving than cv. Baronesse and produced low yet stable yields. Cv. RGT's dynamic water use with high transpiration under ample water supply and moderate transpiration decrease under drought combined with high resilience secured the highest and most stable yields. Discussion: Such a dynamic water-use behavior combined with higher drought resilience and favorable root traits could potentially create an ideotype for intermediate drought. Prospective studies will examine these results in field experiments and will use the newly gained understanding on water use in barley to improve process descriptions in crop simulation models to support crop model--aided ideotype design. [ABSTRACT FROM AUTHOR]

Hulless barley (Hordeum vulgare L. var. nudum Hook. f.), belonging to the genus Gramineae, has high and steady output and thus considered as a principal food crop by Tibetan people. Hulless barley grain can be used for food, brewing, and functional health product development, while its straw serves as an essential supplementary forage and is a crucial cereal crop. Lodging can reduce the yield and quality of barley grain and straw, and it hinders mechanical harvesting. It is a significant factor affecting high and stable yields of barley. Unlike other Poaceae plants (such as rice, wheat), hulless barley is mainly grown in high-altitude regions, where it is susceptible to low temperatures, strong winds, and heavy rainfall. As a result, its stem lodging resistance is relatively weak, making it prone to lodging during the growth period. In this study, we observed that the lignin concentration and the contents of lignin monomers (H, S, and G), and neutral detergent fibre of the lodging-resistant variety Kunlun14 were substantially greater than those of the lodging-sensitive variety Menyuanlianglan. We performed the weighted gene co-expression network analysis (WGCNA) and Short Time-series Expression Miner (STEM) analysis of both the lodging-resistant and lodging-sensitive varieties. Through transcriptome sequencing analysis at different developmental stages, combined with the previously annotated genes related to lodging resistance, a total of 72 DEGs were identified. Among these DEGs, 17 genes were related to lignin, cellulose, and hemicellulose synthesis or regulation, including five transcription factors about NAC, MYB and WRKY. Our results provide a basis for further exploring the molecular mechanism of stem lodging resistance in hulless barley and provide valuable gene resources for stem lodging resistance molecular breeding. [ABSTRACT FROM AUTHOR]

Triticeae, the wheatgrass tribe, includes several major cereal crops and their wild relatives. Major cropswithin the Triticeae are wheat, barley and rye which are important for human consumption, animal feed, and rangeland protection. Species within this tribe are known for their large genomes and complex genetic histories. Powered by recent advances in sequencing technology, researchers worldwide have made progress in elucidating the genomes of Triticeae crops. In addition to assemblies of high-quality reference genomes, pan-genome studies have just started to capture the genomic diversities of these species, shedding light on our understanding of the genetic basis of domestication and environmental adaptation of Triticeae crops. In this review, we focus on recent signs of progress in genome sequencing, pan-genome analyses, and resequencing analysis of Triticeae crops. We also propose future research avenues in Triticeae crop genomes, including identifying genome structure variations, the association of genomic regions with desired traits, mining functions of the non-coding area, introgression of high-quality genes from wild Triticeae resources, genome editing, and integration of genomic resources. [ABSTRACT FROM AUTHOR]

Increasing attention is paid to providing new tools to breeders for targeted breeding for specific root traits that are beneficial in low-fertility, drying soils; however, such information is not available for barley (Hordeum vulgare L.). A panel of 191 barley accessions (originating from Australia, Europe, and Africa) was phenotyped for 26 root and shoot traits using the semi-hydroponic system and genotyped using 21 062 high-quality single nucleotide polymorphism (SNP) markers generated by genotyping-by-sequencing (GBS). The population structure analysis of the barley panel identified six distinct groups. We detected 1199 significant (P<0.001) marker-trait associations (MTAs) with r² values up to 0.41. The strongest MTAs were found for root diameter in the top 20 cm and the longest root length. Based on the physical locations of these MTAs in the barley reference genome, we identified 37 putative QTLs for the root traits, and three QTLs for shoot traits, with nine QTLs located in the same physical regions. The genomic region 640-653 Mb on chromosome 7H was significant for five root length-related traits, where 440 annotated genes were located. The putative QTLs for various root traits identified in this study may be useful for genetic improvement regarding the adaptation of new barley cultivars to suboptimal environments and abiotic stresses. [ABSTRACT FROM AUTHOR]

Plant hormones play important roles in seed development; however, transcriptional regulation of their metabolism and levels of the respective bioactive forms during barley seed development is poorly understood. To this end, this study performed a comprehensive analysis of changes in the expression patterns phytohormone metabolism genes and levels of the respective bioactive forms in the embryo and endosperm tissues. Our study showed the presence of elevated levels of abscisic acid (ABA), bioactive forms of gibberellins (GAs), jasmonate (JA) and cytokinins (CKs), auxin and salicylic acid (SA) in the endosperm and embryo tissues at early stage of seed filling (SF). The levels of all hormones in both tissues, except that of ABA, decreased to low levels during SF. In contrast, embryonic ABA level increased during SF and peaked at physiological maturity (PM) while the endospermic ABA was maintained at a similar level observed during SF. Although its level decreased high amount of ABA was still present in the embryo during post-PM. We detected low levels of ABA in the endosperm and all the other hormones in both tissues during post-PM phase except the relatively higher levels of jasmonoyl-isoleucine and SA detected at late stage of post-PM. Our data also showed that spatiotemporal changes in the levels of plant hormones during barley seed development are mediated by the expression of specific genes involved in their respective metabolic pathways. These results indicate that seed development in barley is mediated by spatiotemporal modulation in the metabolism and levels of plant hormones. [ABSTRACT FROM AUTHOR]

In cereal crops, such as barley (Hordeum vulgare L.), the ability to appropriately respond to environmental cues is an important factor for yield stability and thus for agricultural production. Reactive oxygen species (ROS), such as hydrogen peroxide (H2O2), are key components of signal transduction cascades involved in plant adaptation to changing environmental conditions. H2O2-mediated stress responses include the modulation of expression of stress-responsive genes required to cope with different abiotic and biotic stresses. Despite its importance, knowledge of the effects of H2O2 on the barley transcriptome is still scarce. In this study, we identified global transcriptomic changes induced after application of 10 mM H2O2 to five-day-old barley plants. In total, 1883 and 1001 differentially expressed genes (DEGs) were identified in roots and leaves, respectively. Most of these DEGs were organ-specific, with only 209 DEGs commonly regulated and 37 counter-regulated between both plant parts. A GO term analysis further confirmed that different processes were affected in roots and leaves. It revealed that DEGs in leaves mostly comprised genes associated with hormone signaling, response to H2O2 and abiotic stresses. This includes many transcriptions factors and small heat shock proteins. DEGs in roots mostly comprised genes linked to crucial aspects of H2O2 catabolism and oxidant detoxification, glutathione metabolism, as well as cell wall modulation. These categories include many peroxidases and glutathione transferases. As with leaves, the H2O2 response category in roots contains small heat shock proteins, however, mostly different members of this family were affected and they were all regulated in the opposite direction in the two plant parts. Validation of the expression of the selected commonly regulated DEGs by qRT-PCR was consistent with the RNA-seq data. The data obtained in this study provide an insight into the molecular mechanisms of oxidative stress responses in barley, which might also play a role upon other stresses that induce oxidative bursts. [ABSTRACT FROM AUTHOR]

Corn-leaf aphid (CLA), Rhopalosiphum maidis (Fitch) (Hemiptera: Aphididae) is a serious economic pest of barley worldwide. Breeding for aphid resistance in plants is considered a cost-effective and environmentally safe approach for aphid control, compared to the use of chemical pesticides. One of the challenges in breeding for aphid resistance is the identification of resistant plant genotypes, which can be achieved through the use of molecular markers. In the present study, a set of aphid specific 10 simple-sequence repeats (SSR) markers were used to investigate genetic diversity and population structure analyses in 109 barley genotypes against R. maidis. Three statistical methods viz., multivariate hierarchical clustering based on Jaccard's similarity coefficient, principal coordinate analysis (PCoA) and the Bayesian approach were utilized to classify the 109 barley genotypes. The analyses revealed four subpopulations i.e., SubPop1, SubPop2, SubPop3 and SubPop4 with 19, 46, 20 and 24 genotypes including admixtures, respectively and represented 17.43%, 42.2%, 18.34% and 22.01% genotypes of the total population size, respectively. The studied SSR markers produced 67 polymorphic bands, with an average of 6.7 and ranging from 3 to 12 bands. Heterozygosity (H) was found to be highest in SSR28 (0.64) and lowest in SSR27 (0.89). The observed genetic diversity index varied from 0.10 to 0.34 (with an average of 0.19). Major allele frequency varied from 74.08% to 94.80%. On an average, 87.52% of the 109 barley genotypes shared a common major allele at any locus. Based on the Aphid Infestation Index (AII), only 2 genotypes were found to be resistant against CLA. SubPop2 also had lowest mean aphid population (28.83), widest genetic similarity index (0.60-1.00) and highest genetic similarity coefficient (0.82), which highlighted its potential for inclusion in future CLA resistance breeding programs. [ABSTRACT FROM AUTHOR]

Genome-wide prediction is a powerful tool in breeding. Initial results suggest that genome-wide approaches are also promising for enhancing the use of the genebank material: predicting the performance of plant genetic resources can unlock their hidden potential and fill the information gap in genebanks across the world and, hence, underpin prebreeding programs. As a proof of concept, we evaluated the power of across-genebank prediction for extensive germplasm collections relying on historical data on flowering/heading date, plant height, and thousand kernel weight of 9,344 barley (Hordeum vulgare L.) plant genetic resources from the German Federal Ex situ Genebank for Agricultural and Horticultural Crops (IPK) and of 1,089 accessions from the International Center for Agriculture Research in the Dry Areas (ICARDA) genebank. Based on prediction abilities for each trait, three scenarios for predictive characterization were compared: 1) a benchmark scenario, where test and training sets only contain ICARDA accessions, 2) across-genebank predictions using IPK as training and ICARDA as test set, and 3) integrated genebank predictions that include IPK with 30% of ICARDA accessions as a training set to predict the rest of ICARDA accessions. Within the population of ICARDA accessions, prediction abilities were low to moderate, which was presumably caused by a limited number of accessions used to train the model. Interestingly, ICARDA prediction abilities were boosted up to ninefold by using training sets composed of IPK plus 30% of ICARDA accessions. Pervasive genotype × environment interactions (GEIs) can become a potential obstacle to train robust genome-wide prediction models across genebanks. This suggests that the potential adverse effect of GEI on prediction ability was counterbalanced by the augmented training set with certain connectivity to the test set. Therefore, across-genebank predictions hold the promise to improve the curation of the world’s genebank collections and contribute significantly to the long-term development of traditional genebanks toward biodigital resource centers. [ABSTRACT FROM AUTHOR]

Barley is an important crop worldwide known for its adaptation to harsh environments and used in multiple forms as feed, food and beverages. Its productivity is affected by major abiotic and biotic stresses. Scald caused by hemibiotrophic fungus Rhynchosporium commune is a major foliar disease in many parts of the world. Host plant resistance is targeted by breeders to efficiently control this disease. An association mapping panel of 316 spring barley genotypes (AM2017) was screened for seedling resistance in greenhouse against three R. commune isolates and for adult plant resistance in three field locations in Morocco. The phenotyping results showed different numbers of entries with resistant and moderately resistant reactions at both seedling and adult plant stages. The reactions differed between the isolates with the highest percentage of resistant genotypes observed for isolate SC-S611 (49.4%) and highest percentage of susceptible genotypes (73.8%) for isolate SC-1122. At adult plant stage, the highest percentage of scald resistant genotypes (64.5%) was observed at Rommani site compared to 56% at Guich site and only 28.8% at Marchouch site. Seven genotypes were resistant at the seedling and adult plant stages. Genome wide association study (GWAS) revealed 102 MTA (15 QTL) at the seedling stage, and 25 MTA (12 QTL) associated with scald resistance at the adult plant stage. In addition, the sequences of 92 out of 102 at SRT, and 24 out of 25 significant SNP markers at APR were located in genomic regions enriched with functional proteins involved in diverse cellular processes including disease resistance. These markers span over all chromosomes with the majority of SNPs located on 3H and 7H. This study has verified 18 QTL reported in previous studies. In addition, it was successful in identifying new sources of resistance and novel genomic regions which could help in enhancing scald resistance in barley breeding programs.

Increasing attention is paid to providing new tools to breeders for targeted breeding for specific root traits that are beneficial in low-fertility, drying soils; however, such information is not available for barley (Hordeum vulgare L.). A panel of 191 barley accessions (originating from Australia, Europe, and Africa) was phenotyped for 26 root and shoot traits using the semi-hydroponic system and genotyped using 21 062 high-quality single nucleotide polymorphism (SNP) markers generated by genotyping-by-sequencing (GBS). The population structure analysis of the barley panel identified six distinct groups. We detected 1199 significant (P

In cereal crops, such as barley (Hordeum vulgare L.), the ability to appropriately respond to environmental cues is an important factor for yield stability and thus for agricultural production. Reactive oxygen species (ROS), such as hydrogen peroxide (H2O2), are key components of signal transduction cascades involved in plant adaptation to changing environmental conditions. H2O2-mediated stress responses include the modulation of expression of stress-responsive genes required to cope with different abiotic and biotic stresses. Despite its importance, knowledge of the effects of H2O2 on the barley transcriptome is still scarce. In this study, we identified global transcriptomic changes induced after application of 10 mM H2O2 to five-day-old barley plants. In total, 1883 and 1001 differentially expressed genes (DEGs) were identified in roots and leaves, respectively. Most of these DEGs were organ-specific, with only 209 DEGs commonly regulated and 37 counter-regulated between both plant parts. A GO term analysis further confirmed that different processes were affected in roots and leaves. It revealed that DEGs in leaves mostly comprised genes associated with hormone signaling, response to H2O2 and abiotic stresses. This includes many transcriptions factors and small heat shock proteins. DEGs in roots mostly comprised genes linked to crucial aspects of H2O2 catabolism and oxidant detoxification, glutathione metabolism, as well as cell wall modulation. These categories include many peroxidases and glutathione transferases. As with leaves, the H2O2 response category in roots contains small heat shock proteins, however, mostly different members of this family were affected and they were all regulated in the opposite direction in the two plant parts. Validation of the expression of the selected commonly regulated DEGs by qRT-PCR was consistent with the RNA-seq data. The data obtained in this study provide an insight into the molecular mechanisms of oxidative stress responses in barley, which might also play a role upon other stresses that induce oxidative bursts.

Fusarium crown rot (FCR) causes significant grain yield loss in winter cereals around the world. Breeding for resistance and/or tolerance to FCR has been slow with relatively limited success. In this study, multi-species experiments were used to demonstrate an improved method to quantify FCR infection levels at plant maturity using quantitative PCR (qPCR), as well as the genotype yield retention using residual regression deviation. Using qPCR to measure FCR infection allowed a higher degree of resolution between genotypes than traditional visual stem basal browning assessments. The results were consistent across three environments with different levels of disease expression. The improved measure of FCR infection along with genotype yield retention allows for partitioning of both tolerance and partial resistance. Together these methods offer new insights into FCR partial resistance and its relative importance to tolerance in bread wheat and barley. This new approach offers a more robust, unbiased way to select for both FCR traits within breeding programs. Key message: Genetic gain for tolerance and partial resistance against Fusarium crown rot (FCR) in winter cereals has been impeded by laborious and variable visual measures of infection severity. This paper presents results of an improved method to quantify FCR infection that are strongly correlated to yield loss and reveal previously unrecognised partial resistance in barley and wheat varieties. [ABSTRACT FROM AUTHOR]

Introduction: Multiple nodes and dwarfmutants in barley are a valuable resource for identifying genes that control shoot branching, vegetative growth and development. Methods: In this study, physiological, microscopic and genetic analysis were conducted to characterize and fine-map the underling gene of a barley mutant with Multiple Stem Nodes and Spikes and Dwarf (msnsd), which was selected from EMS- and 60Co-treated barley cv. Edamai 934. Results and discussion: The msnsd mutant had more stem nodes, lower plant height and a shorter plastochron than Edamai 934. Moreover, the mutant had two or more spikes on each tiller. Microscopic analysis showed that the dwarf phenotype of msnsd resulted from reduced cell lengths and cell numbers in the stem. Further physiological analysis showed that msnsd was GA³-deficient, with its plant height increasing after external GA³ application. Genetic analysis revealed that a single recessive nuclear gene, namely, HvMSNSD, controlled the msnsd phenotype. Using a segregating population derived from Harrington and the msnsd mutant, HvMSNSD was fine-mapped on chromosome 5H in a 200 kb interval using bulked segregant analysis (BSA) coupled with RNA-sequencing (BSRseq), with a C-T substitution in the exon of HvTCP25 co-segregating with the msnsd phenotype. RNA-seq analysis showed that a gene encoding gibberellin 2-oxidase 8, a negative regulator of GA biosynthesis, was upregulated in the msnsd mutant. Several known genes related to inflorescence development that were also upregulated and enriched in the msnsd mutant. Collectively, we propose that HvMSNSD regulates the plastochron and morphology of reproductive organs, likely by coordinating GA homeostasis and changed expression of floral development related genes in barley. This study offers valuable insights into the molecular regulation of barley plant architecture and inflorescence development. [ABSTRACT FROM AUTHOR]

Genome editing strategies in barley (Hordeum vulgare L.) typically rely on Agrobacterium-mediated genetic transformation for the delivery of required genetic reagents involving tissue culture techniques. These approaches are genotype-dependent, time-consuming, and labor-intensive, which hampers rapid genome editing in barley. More recently, plant RNA viruses have been engineered to transiently express short guide RNAs facilitating CRISPR/Cas9-based targeted genome editing in plants that constitutively express Cas9. Here, we explored virus-induced genome editing (VIGE) based on barley stripe mosaic virus (BSMV) in Cas9-transgenic barley. Somatic and heritable editing in the ALBOSTRIANS gene (CMF7) resulting in albino/variegated chloroplast-defective barley mutants is shown. In addition, somatic editing in meiosis-related candidate genes in barley encoding ASY1 (an axis-localized HORMA domain protein), MUS81 (a DNA structure-selective endonuclease), and ZYP1 (a transverse filament protein of the synaptonemal complex) was achieved. Hence, the presented VIGE approach using BSMV enables rapid somatic and also heritable targeted gene editing in barley. [ABSTRACT FROM AUTHOR]

Introduction: During drought, plants close their stomata at a critical soil water content (SWC), together with making diverse physiological, developmental, and biochemical responses. Methods: Using precision-phenotyping lysimeters, we imposed pre-flowering drought on four barley varieties (Arvo, Golden Promise, Hankkija 673, and Morex) and followed their physiological responses. For Golden Promise, we carried out RNA-seq on leaf transcripts before and during drought and during recovery, also examining retrotransposon BARE1expression. Transcriptional data were subjected to network analysis. Results: The varieties differed by their critical SWC (ϴcrit), Hankkija 673 responding at the highest and Golden Promise at the lowest. Pathways connected to drought and salinity response were strongly upregulated during drought; pathways connected to growth and development were strongly downregulated. During recovery, growth and development pathways were upregulated; altogether, 117 networked genes involved in ubiquitin-mediated autophagy were downregulated. Discussion: The differential response to SWC suggests adaptation to distinct rainfall patterns. We identified several strongly differentially expressed genes not earlier associated with drought response in barley. BARE1 transcription is strongly transcriptionally upregulated by drought and downregulated during recovery unequally between the investigated cultivars. The downregulation of networked autophagy genes suggests a role for autophagy in drought response; its importance to resilience should be further investigated. [ABSTRACT FROM AUTHOR]

Cereal grains have been domesticated largely from food grains to feed and malting grains. Barley (Hordeum vulgare L.) remains unparalleled in its success as a primary brewing grain. However, there is renewed interest in "alternative" grains for brewing (and distilling) due to attention being placed on flavor, quality, and health (i.e., gluten issues) aspects that they may offer. This review covers basic and general information on "alternative grains" for malting and brewing, as well as an in-depth look at several major biochemical aspects of these grains including starch, protein, polyphenols, and lipids. These traits are described in terms of their effects on processing and flavor, as well as the prospects for improvement through breeding. These aspects have been studied extensively in barley, but little is known about the functional properties in other crops for malting and brewing. In addition, the complex nature of malting and brewing produces a large number of brewing targets but requires extensive processing, laboratory analysis, and accompanying sensory analysis. However, if a better understanding of the potential of alternative crops that can be used in malting and brewing is needed, then significantly more research is required. [ABSTRACT FROM AUTHOR]

Drought stress can significantly affect plant growth and development. Biochar (BC) and plant growth-promoting rhizobacteria (PGPR) have been found to increase plant fertility and development under drought conditions. The single effects of BC and PGPR in different plant species have been widely reported under abiotic stress. However, there have been relatively few studies on the positive role of PGPR, BC, and their combination in barley (Hordeum vulgare L.). Therefore, the current study investigated the effects of BC from Parthenium hysterophorus, drought tolerant PGPR (Serratia odorifera), and the combination of BC + PGPR on the growth, physiology, and biochemical traits of barley plants under drought stress for two weeks. A total of 15 pots were used under five treatments. Each pot of 4 kg soil comprised the control (T0, 90% water), drought stress alone (T1, 30% water), 35 mL PGPR/kg soil (T2, 30% water), 2.5%/kg soil BC (T3, 30% water), and a combination of BC and PGPR (T4, 30% water). Combined PGPR and BC strongly mitigated the negative effects of drought by improving the shoot length (37.03%), fresh biomass (52%), dry biomass (62.5%), and seed germination (40%) compared to the control. The PGPR + BC amendment treatment enhanced physiological traits, such as chlorophyll a (27.9%), chlorophyll b (35.3%), and total chlorophyll (31.1%), compared to the control. Similarly, the synergistic role of PGPR and BC significantly (p< 0.05) enhanced the antioxidant enzyme activity including peroxidase (POD), catalase (CAT), and superoxide dismutase (SOD) to alleviate the toxicity of ROS. The physicochemical properties (N, K, P, and EL) of the soils were also enhanced by (85%, 33%, 52%, and 58%) respectively, under the BC + PGPR treatment compared to the control and drought stress alone. The findings of this study have suggested that the addition of BC, PGPR, and a combination of both will improve the soil fertility, productivity, and antioxidant defense systems of barley under drought stress. Therefore, BC from the invasive plant P. hysterophorus and PGPR can be applied to water-deficient areas to improve barley crop production. [ABSTRACT FROM AUTHOR]

Low-temperature stress (LTS) is among the major abiotic stresses affecting the geographical distribution and productivity of the most important crops. Understanding the genetic basis of photosynthetic variation under cold stress is necessary for developing more climate-resilient barley cultivars. To that end, we investigated the ability of chlorophyll fluorescence parameters (FVFM, and FVF0) to respond to changes in the maximum quantum yield of Photosystem II photochemistry as an indicator of photosynthetic energy. A panel of 96 barley spring cultivars from different breeding zones of Canada was evaluated for chlorophyll fluorescence-related traits under cold acclimation and freeze shock stresses at different times. Genome-wide association studies (GWAS) were performed using a mixed linear model (MLM). We identified three major and putative genomic regions harboring 52 significant quantitative trait nucleotides (QTNs) on chromosomes 1H, 3H, and 6H for low-temperature tolerance. Functional annotation indicated several QTNs were either within the known or close to genes that play important roles in the photosynthetic metabolites such as abscisic acid (ABA) signaling, hydrolase activity, protein kinase, and transduction of environmental signal transduction at the posttranslational modification levels. These outcomes revealed that barley plants modified their gene expression profile in response to decreasing temperatures resulting in physiological and biochemical modifications. Cold tolerance could influence a long-term adaption of barley in many parts of the world. Since the degree and frequency of LTS vary considerably among production sites. Hence, these results could shed light on potential approaches for improving barley productivity under low-temperature stress. [ABSTRACT FROM AUTHOR]

Nitrogen (N) is one of the most expensive nutrients to supply, therefore, improving the efficiency of N use is essential to reduce the cost of commercial fertilization in plant production. Since cells cannot store reduced N as NH3 or NH4+, polyamines (PAs), the low molecular weight aliphatic nitrogenous bases, are important N storage compounds in plants. Manipulating polyamines may provide a method to increase nitrogen remobilization efficiency. Homeostasis of PAs is maintained by intricate multiple feedback mechanisms at the level of biosynthesis, catabolism, efflux, and uptake. The molecular characterization of the PA uptake transporter (PUT) in most crop plants remains largely unknown, and knowledge of polyamine exporters in plants is lacking. Bi-directional amino acid transporters (BATs) have been recently suggested as possible PAs exporters for Arabidopsis and rice, however, detailed characterization of these genes in crops is missing. This report describes the first systematic study to comprehensively analyze PA transporters in barley (Hordeum vulgare, Hv), specifically the PUT and BAT gene families. Here, seven PUTs (HvPUT1-7) and six BATs (HvBAT1-6) genes were identified as PA transporters in the barley genome and the detailed characterization of these HvPUT and HvBAT genes and proteins is provided. Homology modeling of all studied PA transporters provided 3D structures prediction of the proteins of interest with high accuracy. Moreover, molecular docking studies provided insights into the PA-binding pockets of HvPUTs and HvBATs facilitating improved understanding of the mechanisms and interactions involved in HvPUT/HvBAT-mediated transport of PAs. We also examined the physiochemical characteristics of PA transporters and discuss the function of PA transporters in barley development, and how they help barley respond to stress, with a particular emphasis on leaf senescence. Insights gained here could lead to improved barley production via modulation of polyamine homeostasis. [ABSTRACT FROM AUTHOR]

Roots are the hidden parts of plants, anchoring their above-ground counterparts in the soil. They are responsible for water and nutrient uptake and for interacting with biotic and abiotic factors in the soil. The root system architecture (RSA) and its plasticity are crucial for resource acquisition and consequently correlate with plant performance while being highly dependent on the surrounding environment, such as soil properties and therefore environmental conditions. Thus, especially for crop plants and regarding agricultural challenges, it is essential to perform molecular and phenotypic analyses of the root system under conditions as near as possible to nature (#asnearaspossibletonature). To prevent root illumination during experimental procedures, which would heavily affect root development, Dark-Root (D-Root) devices (DRDs) have been developed. In this article, we describe the construction and different applications of a sustainable, affordable, flexible, and easy to assemble open-hardware bench-top LEGO® DRD, the DRD-BIBLOX (Brick Black Box). The DRD-BIBLOX consists of one or more 3D-printed rhizoboxes, which can be filled with soil while still providing root visibility. The rhizoboxes sit in a scaffold of secondhand LEGO® bricks, which allows root development in the dark and non-invasive root tracking with an infrared (IR) camera and an IR light-emitting diode (LED) cluster. Proteomic analyses confirmed significant effects of root illumination on barley root and shoot proteomes. Additionally, we confirmed the significant effect of root illumination on barley root and shoot phenotypes. Our data therefore reinforces the importance of the application of field conditions in the lab and the value of our novel device, the DRD-BIBLOX. We further provide a DRD-BIBLOX application spectrum, spanning from investigating a variety of plant species and soil conditions and simulating different environmental conditions and stresses, to proteomic and phenotypic analyses, including early root tracking in the dark. [ABSTRACT FROM AUTHOR]