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Malt production is one of the important uses of barley, and its quality differs greatly depending on the barley varieties used. In this study, ultraperformance liquid chromatography coupled to quadrupole time-of-flight mass spectrometry technology was used to investigate the temporal changes of metabolites during malting in two barley varieties: Franklin (malt barley) and Yerong (non-malt barley). Also, differences in metabolite profiles were compared in the kilned malt between two other malt barley varieties (Copeland and Planet) and two non-malt varieties (ZD10 and Hua30). Results showed that degradation of trisaccharide and accumulation of UDP-glucose and mannose-1-phosphate are the key metabolic events during steeping, with Franklin showing earlier and greater changes. Earlier increase of sugars and amino acids in Franklin is associated with its faster germination rate. Comparative metabolome analysis of kilned malt from the different barley varieties indicated that malt barley accumulated more sugars, hordatine-glucoside, and oxoproline, and non-malt barley accumulated more polyphenols and monogalactosylmonoacylglycerol. These results improved the understanding of the genotypic difference in the formation of malt quality at the metabolomic level.

Vacuolar H+-pumping pyrophosphatases (VPs) provide a proton gradient for Na+ sequestration in the tonoplast; however, the regulatory mechanisms of VPs in developing salt tolerance have not been fully elucidated. Here, we cloned a barley (Hordeum vulgare) VP gene (HVP10) that was identified previously as the HvNax3 gene. Homology analysis showed VP10 in plants had conserved structure and sequence and likely originated from the ancestors of the Ceramiales order of Rhodophyta (Cyanidioschyzon merolae). HVP10 was mainly expressed in roots and upregulated in response to salt stress. After salt treatment for 3 weeks, HVP10 knockdown (RNA interference) and knockout (CRISPR/Cas9 gene editing) barley plants showed greatly inhibited growth and higher shoot Na+ concentration, Na+ transportation rate and xylem Na+ loading relative to wild-type (WT) plants. Reverse transcription quantitative polymerase chain reaction and microelectronic Ion Flux Estimation results indicated that HVP10 likely modulates Na+ sequestration into the root vacuole by acting synergistically with Na+/H+ antiporters (HvNHX1 and HvNHX4) to enhance H+ efflux and K+ maintenance in roots. Moreover, transgenic rice (Oryza sativa) lines overexpressing HVP10 also showed higher salt tolerance than the WT at both seedling and adult stages with less Na+ translocation to shoots and higher grain yields under salt stress. This study reveals the molecular mechanism of HVP10 underlying salt tolerance and highlights its potential in improving crop salt tolerance.

Frequent climate change-induced drought events are detrimental environmental stresses affecting global crop production and ecosystem health. Several efforts have facilitated crop breeding for resilient varieties to counteract stress. However, progress is hampered due to the complexity of drought tolerance; a greater variety of novel genes are required across varying environments. Tibetan annual wild barley is a unique and precious germplasm that is well adapted to abiotic stress and can provide elite genes for crop improvement in drought tolerance.To identify the genetic basis and unique mechanisms for drought tolerance in Tibetan wild barley.Whole genome resequencing and comparative RNA-seq approaches were performed to identify candidate genes associated with drought tolerance via investigating the genetic diversity and transcriptional variation between cultivated and Tibetan wild barley. Bioinformatics, population genetics, and gene silencing were conducted to obtain insights into ecological adaptation in barley and functions of key genes.Over 20 million genetic variants and a total of 15,361 significantly affected genes were identified in our dataset. Combined genomic, transcriptomic, evolutionary, and experimental analyses revealed 26 water deficit resilience-associated genes in the drought-tolerant wild barley XZ5 with unique genetic variants and expression patterns. Functional prediction revealed Tibetan wild barley employs effective regulators to activate various responsive pathways with novel genes, such as Zinc-Induced Facilitator-Like 2 (HvZIFL2) and Peroxidase 11 (HvPOD11), to adapt to water deficit conditions. Gene silencing and drought tolerance evaluation in a natural barley population demonstrated that HvZIFL2 and HvPOD11 positively regulate drought tolerance in barley.Our findings reveal functional genes that have been selected across barley's complex history of domestication to thrive in water deficit environments. This will be useful for molecular breeding and provide new insights into drought-tolerance mechanisms in wild relatives of major cereal crops.

This study displayed the transcriptional regulation network of key regulators and downstream pathway in seedling morphogenesis of Brassica napus under different light quality. Plants undergo photomorphogenesis upon the presence of light, mediated by different light (e.g., blue, red, and far-red) signaling pathways. Although the light signaling pathway has been well documented in Arabidopsis, the underlying mechanisms were studied to a less extent in other plant species including Brassica napus. In this study, we investigated the effect of different light qualities (white, blue, red, and far-red light) on the hypocotyl elongation in B. napus, and performed the transcriptomic analysis of seedlings in response to different light qualities. The results showed that hypocotyl elongation was slightly inhibited by red light, while it was strongly inhibited by blue/far-red light. Transcriptome analysis identified 9748 differentially expressed genes (DEGs) among treatments. Gene ontology (GO) enrichment analysis of DEGs showed that light-responsive and photosynthesis-related genes were highly expressed in response to blue/far-red light rather than in red light. Furthermore, the key genes in light signaling (i.e., PHYB, HY5, HYH, HFR1, and PIF3) exhibited distinct expression patterns between blue/far-red and red light treatments. In addition, subgenome dominant expression of homoeologous genes were observed for some genes, such as PHYA, PHYB, HFR1, and BBXs. The current study displayed a comprehensive dissection of light-mediated transcriptional regulation network, including light signaling, phytohormone, and cell elongation/modification, which improved the understanding on the underlying mechanism of light-regulated hypocotyl growth in B. napus.

Ferns appear in the fossil record some 200 Myr before angiosperms. However, as angiosperm‐dominated forest canopies emerged in the Cretaceous period there was an explosive diversification of modern (leptosporangiate) ferns, which thrived in low, blue‐enhanced light beneath angiosperm canopies. A mechanistic explanation for this transformative event in the diversification of ferns has remained elusive. We used physiological assays, transcriptome analysis and evolutionary bioinformatics to investigate a potential connection between the evolution of enhanced stomatal sensitivity to blue light in modern ferns and the rise of angiosperm‐dominated forests in the geological record. We demonstrate that members of the largest subclade of leptosporangiate ferns, Polypodiales, have significantly faster stomatal response to blue light than more ancient fern lineages and a representative angiosperm. We link this higher sensitivity to levels of differentially expressed genes in blue‐light signaling, particularly in the cryptochrome (CRY) signaling pathway. Moreover, CRYs of the Polypodiales examined show gene duplication events between 212.9–196.9 and 164.4–151.8 Ma, when angiosperms were emerging, which are lacking in other major clades of extant land plants. These findings suggest that evolution of stomatal blue‐light sensitivity helped modern ferns exploit the shady habitat beneath angiosperm forest canopies, fueling their Cretaceous hyperdiversification.

The large size and complexity of most fern genomes have hampered efforts to elucidate fundamental aspects of fern biology and land plant evolution through genome-enabled research. Here we present a chromosomal genome assembly and associated methylome, transcriptome and metabolome analyses for the model fern species Ceratopteris richardii. The assembly reveals a history of remarkably dynamic genome evolution including rapid changes in genome content and structure following the most recent whole-genome duplication approximately 60 million years ago. These changes include massive gene loss, rampant tandem duplications and multiple horizontal gene transfers from bacteria, contributing to the diversification of defence-related gene families. The insertion of transposable elements into introns has led to the large size of the Ceratopteris genome and to exceptionally long genes relative to other plants. Gene family analyses indicate that genes directing seed development were co-opted from those controlling the development of fern sporangia, providing insights into seed plant evolution. Our findings and annotated genome assembly extend the utility of Ceratopteris as a model for investigating and teaching plant biology.

At the microsite “Evolution Slope”, Tabigha, Israel, wild barley (Hordeum spontaneum) populations adapted to dry Terra Rossa soil, and its derivative abutting wild barley population adapted to moist and fungi‐rich Basalt soil. However, the mechanisms underlying the edaphic adaptation remain elusive. Accordingly, whole genome bisulfite sequencing, RNA‐sequencing, and metabolome analysis are performed on ten wild barley accessions inhabiting Terra Rossa and Basalt soil. A total of 121 433 differentially methylated regions (DMRs) and 10 478 DMR‐genes are identified between the two wild barley populations. DMR‐genes in CG context (CG‐DMR‐genes) are enriched in the pathways related with the fundamental processes, and DMR‐genes in CHH context (CHH‐DMR‐genes) are mainly associated with defense response. Transcriptome and metabolome analysis reveal that the primary and secondary metabolisms are more active in Terra Rossa and Basalt wild barley populations, respectively. Multi‐omics analysis indicate that sugar metabolism facilitates the adaptation of wild barley to dry Terra Rossa soil, whereas the enhancement of phenylpropanoid/phenolamide biosynthesis is beneficial for wild barley to inhabit moist and fungi pathogen‐rich Basalt soil. The current results make a deep insight into edaphic adaptation of wild barley and provide elite genetic and epigenetic resources for developing barley with high abiotic stress tolerance., Two wild barley populations adapt to hot‐dry Terra Rossa soil, and abutting moist fungi‐rich Basalt soil at the microsite “Evolution Slope”, Tabigha, Israel. Sugar metabolism facilitates edaphic adaptation of wild barley to dry Terra Rossa soil, and phenylpropanoid/phenolamide biosynthesis enables the edaphic adaptation of wild barley to moist and fungi‐rich Basalt soil at epigenomic, genomic, transcriptomic, and metabolomic levels.

Background Endo-β-1,4-xylanase1 (EA), the key endoxylanase in plants, is involved in the degradation of arabinoxylan during grain germination. In barley (Hordeum vulgare L.), one gene (HvXYN-1) that encode a endo-beta-1,4-xylanase, has been cloned. However, the single nucleotide polymorphisms (SNPs) that affect the endoxylanase activity and total arabinoxylan (TAX) content have yet to be characterized. The investigation of genetic variation in HvXYN1 may facilitate a better understanding of the relationship between TAX content and EA activity in barley. Results In the current study, 56 polymorphisms were detected in HvXYN1 among 210 barley accessions collected from 34 countries, with 10 distinct haplotypes identified. The SNPs at positions 110, 305, 1045, 1417, 1504, 1597, 1880 bp in the genomic region of HvXYN1 were significantly associated with EA activity (P

Phosphite (PHI) has been used in the management of Phytophthora diseases since the 1970s.We assessed the effect of PHI on controlling the incidence of Xanthomonas oryzae pv.oryzae and Pyricularia grisea. As a result, PHI application significantly inhibited the incidence of the diseases. To clarify the molecular mechanism underlying this, a transcriptome study was employed. In total, 2064 differentially expressed genes (DEGs) were identified between control and PHI treatment. The key DEGs could be classified into phenylpropanoid biosynthesis (ko00940), starch and sucrose metabolism (ko00500), and plant hormone signal transduction (ko04075). The expressions of defense-related genes had a higher expression lever upon PHI treatment. This study provides new insights into the mechanism of protection effect of PHI against pathogens.

Background: Plants undergo skotomorphogenesis or photomorphogenesis upon the absence or presence of light. Although the light signaling pathway has been well documented in Arabidopsis, relatively little is known in other plant species including Brassica napus.Results: In this study, we examined the response of hypocotyl elongation to different light quality (white, blue, red and far-red light) in B. napus. The result showed each of tested light quality promoted photomorphogenesis, with less effect of red light than blue/far-red light. Subsequently, RNA-sequencing was performed to obtain the transcriptomic profile in B. napus in response to dark and different light quality. A total of 9748 genes were differentially expressed among treatments. Correlation analysis and hierarchical cluster analysis of differentially expressed genes (DEGs) indicated the transcriptomic profile in red light exhibited an analogous pattern with that in dark to some extent, while blue/far-red light induced transcriptional profiles showed distinct patterns. Moreover, we found the differential expressions of PHYB, COP1 and BBXs homologs between two subgenomes, which may affect their interactions with up/downstream pathway. Finally, we constructed a transcriptional network including light signaling, phytohormone and cell elongation/modification, which explained the observed hypocotyl phenotype under different light quality.Conclusions: We partially explained why red light has reduced effect on photomorphogenesis in B. napus by RNA-seq and displayed gene expression of the key regulators in light signaling pathway and their contribution to photomorphogenesis under different light quality in B. napus. The result could improve new sight into understanding on transcriptional regulation of photomorphogenesis of B. napus.

Plants will meet various abiotic stresses during their growth and development. One of the important strategies for plants to deal with the stress is involved in metabolic regulation, causing the dramatic changes of metabolite profiles. Metabolomic studies have been intensively conducted to reveal the responses of plants to abiotic stress, but most of them were limited to one or at most two abiotic stresses in a single experiment. In this study, we compared the metabolite profiles of barley seedlings exposed to seven abiotic stresses, including drought, salt stress, aluminum (Al), cadmium (Cd), deficiency of nitrogen (N), phosphorus (P) and potassium (K). The results showed that metabolite profiles of barley under these stresses could be classified into three groups: osmotic stresses (drought and salt); metal stresses (Al and Cd) and nutrient deficiencies (N, P and K deficiencies). Compared with the control, some metabolites (including polyamines, raffinose and pipecolic acid) in plants exposed to all abiotic stresses changed significantly, while some other metabolites showed the specific change only under a certain abiotic stress, such as proline being largely increased by osmotic stress (drought and salinity), the P-containing metabolites being largely decreased under P deficiency, some amino acids (lysine, tyrosine, threonine, ornithine, glutamine and so on) showing the dramatic reduction in the plants exposed to N deficiencies, respectively. The current meta-analysis obtained a comprehensive view on the metabolic responses to various abiotic stress, and improved the understanding of the mechanisms for tolerance of barley to abiotic stress.

Aluminum (Al) toxicity is a major abiotic stress that restricts crop production in acid soils. Plants have evolved internal and external mechanisms of tolerance, and among them it is well known that AtSTOP1 and OsART1 are key transcription factors involved in tolerance through regulation of multiple downstream genes. Here, we identified the closest homolog of these two proteins in barley, namely HvATF1, Al-tolerance Transcription Factor 1, and determined its potential function in Al stress. HvATF1 is expressed in the nucleus, and functions in transcriptional activation. The transcription of HvATF1 was found to be constitutive in different tissues, and was little affected by Al stress. Knockdown of HvATF1 by RNAi resulted in increased Al sensitivity. Transcriptomics analysis identified 64 differently expressed genes in the RNAi lines compared to the wild-type, and these were considered as candidate downstream genes regulated by HvATF1. This study provides insights into the different molecular mechanisms of Al tolerance in barley and other plants.

Stronger Na+ extrusion and vacuolar sequestration are essential to confer better salt tolerance in bread wheat than in durum wheat. Removal of the root meristems increased salt sensitivity in wheat., The progress in plant breeding for salinity stress tolerance is handicapped by the lack of understanding of the specificity of salt stress signalling and adaptation at the cellular and tissue levels. In this study, we used electrophysiological, fluorescence imaging, and real-time quantitative PCR tools to elucidate the essentiality of the cytosolic Na+ extrusion in functionally different root zones (elongation, meristem, and mature) in a large number of bread and durum wheat accessions. We show that the difference in the root’s ability for vacuolar Na+ sequestration in the mature zone may explain differential salinity stress tolerance between salt-sensitive durum and salt-tolerant bread wheat species. Bread wheat genotypes also had on average 30% higher capacity for net Na+ efflux from the root elongation zone, providing the first direct evidence for the essentiality of the root salt exclusion trait at the cellular level. At the same time, cytosolic Na+ accumulation in the root meristem was significantly higher in bread wheat, leading to the suggestion that this tissue may harbour a putative salt sensor. This hypothesis was then tested by investigating patterns of Na+ distribution and the relative expression level of several key genes related to Na+ transport in leaves in plants with intact roots and in those in which the root meristems were removed. We show that tampering with this sensing mechanism has resulted in a salt-sensitive phenotype, largely due to compromising the plant’s ability to sequester Na+ in mesophyll cell vacuoles. The implications of these findings for plant breeding for salinity stress tolerance are discussed.

Background The interest has been increasing on the phenolic compounds in plants because of their nutritive function as food and the roles regulating plant growth. However, their underlying genetic mechanism in barley is still not clear. Results A genome-wide association study (GWAS) was conducted for total phenolic content (TPC), total flavonoid content (FLC) and antioxidant activity (AOA) in 67 cultivated and 156 Tibetan wild barley genotypes. Most markers associated with phenolic content were different in cultivated and wild barleys. The markers bPb-0572 and bPb-4531 were identified as the major QTLs controlling phenolic compounds in Tibetan wild barley. Moreover, the marker bPb-4531 was co-located with the UDP- glycosyltransferase gene (HvUGT), which is a homolog to Arabidopsis UGTs and involved in biosynthesis of flavonoid glycosides . Conclusions GWAS is an efficient tool for exploring the genetic architecture of phenolic compounds in the cultivated and Tibetan wild barleys. The DArT markers applied in this study can be used in barley breeding for developing new barley cultivars with higher phenolics content. The candidate gene (HvUGT) provides a potential route for deep understanding of the molecular mechanism of flavonoid synthesis. Electronic supplementary material The online version of this article (10.1186/s12864-018-4483-6) contains supplementary material, which is available to authorized users.

While the role of the vacuolar NHX Na+/H+ exchangers in plant salt tolerance has been demonstrated on numerous occasions, their control over cytosolic ionic relations has never been functionally analysed in the context of subcellular Na+ and K+ homeostasis. In this work, PutNHX1 and SeNHX1 were cloned from halophytes Puccinellia tenuiflora and Salicornia europaea and transiently expressed in Arabidopsis wild type Col-0 and the nhx1 mutant. Phylogentic analysis, topological prediction, analysis of evolutionary conservation, the topology structure and analysis of hydrophobic or polar regions of PutNHX1 and SeNHX1 indicated that they are unique tonoplast Na+/H+ antiporters with characteristics for salt tolerance. As a part of the functional assessment, cytosolic and vacuolar Na+ and K+ in different root tissues and ion fluxes from root mature zone of Col-0, nhx1 and their transgenic lines were measured. Transgenic lines sequestered large quantity of Na+ into root cell vacuoles and also promoted high cytosolic and vacuolar K+ accumulation. Expression of PutNHX1 and SeNHX1 led to significant transient root Na+ uptake in the four transgenic lines upon recovery from salt treatment. In contrast, the nhx1 mutant maintained a prolonged Na+ efflux and the nhx1:PutNHX1 and nhx1:SeNHX1 lines started to actively pump Na+ out of the cell. Overall, our findings suggest that PutNHX1 and SeNHX1 improve Na+ sequestration in the vacuole and K+ retention in the cytosol and vacuole of root cells of Arabidopsis, and that they interact with other regulatory mechanisms to provide a highly orchestrated regulation of ionic relations among intracellular cell compartments.

The degradation of starch in barley grains is a primary step of beer production. The addition of an appropriate amount of gibberellin (GA) promotes the production of fermentable sugars, beneficial to the brewing industry. However, the response of proteomics in germinating barley to GA and abscisic acid (ABA) treatments is not thoroughly understood. In this study, isobaric tags for relative and absolute quantitation (iTRAQ) proteomics analysis was performed to illustrate the change of proteins in Tibetan wild barley XZ72 and XZ95 under GA and ABA treatments during germination. XZ72 had more proteins upregulated than XZ95 under GA treatment, while under ABA treatments, XZ95 had more proteins upregulated than XZ72. Concerning the proteins involved in energy metabolism under GA treatment, XZ72 had more proteins upregulated than XZ95. Among the 174 proteins related to starch metabolism, 31 proteins related to starch hydrolysis, such as α-amylase, α-glucosidase, and β-fructofuranosidase, showed higher relative abundance in control and GA treatments in XZ72 than in XZ95. Analysis of correlation between proteins and metabolites indicated that higher hydrolase activity is beneficial for the accumulation of fermentable sugars during germination. On the other hand, 26 starch-synthesis-related proteins were upregulated in XZ95 under ABA treatment. It may be suggested that GA-induced proteins act as accelerators of starch degradation, while ABA-induced proteins inhibit starch degradation. The current results showed that XZ72 is highly capable of allocating the starch-hydrolyzing enzymes, which play important roles in starch breakdown.

Table S3. EA activity and TAX content contrast in different superior alleles. (DOCX 15 kb)

Table S1. EA activity and TAX content contrast in different regions. (DOCX 15 kb)

Figure S2. Identification of TAX content by association analyses. Manhattan plots in the 210 accessions. And three SNPs (red points) about HvXYN1 identified in this study with TAX content. The X axis represents the physical position of HvXYN1, and Negative log10-transformed P values are plotted on the vertical axis. (PDF 76 kb)

Table S2. The mean value comparison of EA activity and TAX content in different haplotype. (DOCX 16 kb)

Table S5. List of overlapping primers used for the amplificati1on of 1.9â kb nucleotide region of HvXYN1 gene. (DOCX 14 kb)

Figure S3. Identification of EA activity and TAX content by association analyses. Quantile-quantile (Q-Q) plots in the 210 accessions. Red points present EA activity and blue points present TAX content. The X axis represents the expected P values and P values are plotted on the vertical axis. (PDF 67 kb)

Figure S1. Identification of EA by association analyses. Manhattan plots in the 210 accessions. and seven SNPs (red points) about HvXYN1 identified in this study with EA activity. The X axis represents the physical position of HvXYN1, and Negative log10-transformed P values are plotted on the vertical axis. (PDF 81 kb)

Limit dextrinase (LD) and limit dextrinase inhibitor (LDI) are the two important traits affecting malt quality. The genetic variation and controlling of LD activity and LDI content in barley grains and malt are not well understood. In this study, we measured LD activity and LDI content in both grains and malt of 68 cultivated barley genotypes. The results show that there is a wide difference among barley genotypes in both LD activity and LDI content. LD in malt is not correlated with LD in grains, but negatively correlated with LDI in malt. LD in malt is positively correlated with diastatic power (DP), Kolbach index (KI) and soluble nitrogen content (SN), and negatively correlated with viscosity (VC). LDI in malt is positively correlated with DP and total nitrogen (TN), and negatively correlated with KI. Association analysis identifies 5 QTLs associated with LD and 3 QTLs associated with LDI in malt. Three major QTLs controlling LD in malt account for 35.7%, 35.7% and 28.4% of phenotypic variation, respectively. A total of 17 QTLs associated with malt quality are identified. The current results address the importance of both LD and LDI in affecting malt quality and the identified QTLs could be useful in barley breeding.

Correlation analysis of TPC, FLC and AOA in Tibetan wild barley. (a) correlation for TPC between 2013 and 2014; (b) correlation in FLC between 2013 and 2014; (c) correlation in AOA between 2013 and 2014. **, represents significant correlation at Pâ

Decay of linkage disequilibrium of the population of 223 genotypes based on 801 DArT markers. The X-axis showed the genetic distance, the Y-axis showed the r2, the squared allele frequency correlations, which is a measurement of the correlation between a pair of variables. (DOCX 66Â kb)

Phylogenetic tree (UPGMA) of 223 barley genotypes based on 801 DArT markers. C, cultivated barley; X, Tibetan wild barley. (DOCX 831Â kb)

Estimation of the most probable number of clusters (k). (DOCX 27Â kb)

Quartile-quartile (Q-Q) plots of P value for 223 genotypes under three GWAS models. The black line is an expected one under the null distribution. Red symbol represents the observed P values for AOA, blue for FLC and green for TPC. (DOCX 425Â kb)

Functional domain prediction of HvUGT. a: function domain in HvUGT; b: function domain in UGT91C1. Pink color represents low complexity sequence, the number marks amino acid sequence sites. (PPTX 90 kb)

List of DArT markers (−log10(p) > 2.5) in 223 genotypes and within subgroups. These markers were identified using Q + K model with significance threshold as –log10(p) > 2.5. R2 (marker) denotes the contribution of the marker for phenotypic variation. (DOCX 35 kb)

Correlation analysis of TPC, FLC and AOA in 223 accessions. ** represent significant correlation at Pâ

Phenolic acids have been quite extensively studied in food science research because of their antioxidative effect. In this study, the genotypic difference and genetic control of phenolic acids, and their correlation with malt quality, were investigated in barley. Ferulic acid (FA) and p-coumaric acid (p-CA) were identified as two main phenolic acids, showing wide variations among 68 barley genotypes. The mean content of FA and p-CA were 2.15 μg g(-1) and 1.10 μg g(-1) in grains and 4.07 μg g(-1) and 1.44 μg g(-1) in malt, respectively. After malting, FA and p-CA were increased significantly in 55 and 37 genotypes and were reduced in 2 and 14 genotypes, respectively. Both malt FA and p-CA were positively correlated with soluble N content and Kolbach index and negatively correlated with malt extract and viscosity. The results indicated that the effect of malting on the change of an individual phenolic acid is genotype independent. Association mapping identified that 8 markers on Chromosomes 1H, 2H, 4H, and 7H are associated with grain p-CA and 4 markers on Chromosomes 3H and 7H are linked with grain FA. However, only a single marker on Chromosome 3H was found to be associated with malt FA. Moreover, a lack of overlapping markers between grain and malt indicated the genetic diversity of phenolic acids in barley grain and malt. Our results strengthen the understanding of phenolic acids in barley and their responses to the malting process.

Limit dextrinase (LD) is an essential amylolytic enzyme for the complete degradation of starch, and it is closely associated with malt quality. A survey of 51 cultivated barley and 40 Tibetan wild barley genotypes showed a wide genetic diversity of LD activity and LD thermostability. Compared with cultivated barley, Tibetan wild barley showed lower LD activity and higher LD thermostability. A doubled haploid population composed of 496 DArT and 28 microsatellite markers was used for mapping Quantitative Trait Loci (QTLs). Parental line Yerong showed low LD activity and high LD thermostability, but Franklin exhibited high LD activity and low LD thermostability. Three QTLs associated with thermostable LD were identified. The major QTL is close to the LD gene on chromosome 7H. The two minor QTLs colocalized with previously reported QTLs determining malt-extract and diastatic power on chromosomes 1H and 2H, respectively. These QTLs may be useful for a better understanding of the genetic control of LD activity and LD thermostability in barley.

The domestication of cultivated barley has been used as a model system for studying the origins and early spread of agrarian culture. Our previous results indicated that the Tibetan Plateau and its vicinity is one of the centers of domestication of cultivated barley. Here we reveal multiple origins of domesticated barley using transcriptome profiling of cultivated and wild-barley genotypes. Approximately 48-Gb of clean transcript sequences in 12 Hordeum spontaneum and 9 Hordeum vulgare accessions were generated. We reported 12,530 de novo assembled transcripts in all of the 21 samples. Population structure analysis showed that Tibetan hulless barley (qingke) might have existed in the early stage of domestication. Based on the large number of unique genomic regions showing the similarity between cultivated and wild-barley groups, we propose that the genomic origin of modern cultivated barley is derived from wild-barley genotypes in the Fertile Crescent (mainly in chromosomes 1H, 2H, and 3H) and Tibet (mainly in chromosomes 4H, 5H, 6H, and 7H). This study indicates that the domestication of barley may have occurred over time in geographically distinct regions.