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Barley is one of the most important cereal crops in the world, and its value as a food is constantly being revealed, so the research into and the use of barley germplasm are very important for global food security. Although a large number of barley germplasm samples have been collected globally, their specific genetic compositions are not well understood, and in many cases their origins are even disputed. In this study, 183 barley germplasm samples from the Shanghai Agricultural Gene Bank were genotyped using genotyping-by-sequencing (GBS) technology, SNPs were identified and their genetic parameters were estimated, principal component analysis (PCA) was preformed, and the phylogenetic tree and population structure of the samples were also analyzed. In addition, a genome-wide association study (GWAS) was carried out for the hulled/naked grain trait, and a KASP marker was developed using an associated SNP. The results showed that a total of 181,906 SNPs were identified, and these barley germplasm samples could be roughly divided into three categories according to the phylogenetic analysis, which was generally consistent with the classification of the traits of row type and hulled/naked grain. Population structure analysis showed that the whole barley population could be divided into four sub-populations (SPs), the main difference from previous classifications being that the two-rowed and the hulled genotypes were sub-divided into two SPs. The GWAS analysis of the hulled/naked trait showed that many associated loci were unrelated to the Nud/nud locus, indicating that there might be new loci controlling the trait. A KASP marker was developed for one exon-type SNP on chromosome 7. Genotyping based on the KASP assay was consistent with that based on SNPs, indicating that the gene of this locus might be associated with the hulled/naked trait. The above work not only lays a good foundation for the future utilization of this barley germplasm population but it provides new loci and candidate genes for the hulled/naked trait. [ABSTRACT FROM AUTHOR]

As the global human population continues to increase, the use of saline–alkali land for food production is an important consideration for food security. In addition to breeding or cultivating salt-tolerant crop varieties, microorganisms are increasingly being evaluated for their ability to improve plant salt tolerance. Barley is one of the most important and salt-tolerant cereal crops and is a model system for investigating the roles of microorganisms in improving plant salt tolerance. However, a comprehensive review of the mechanisms by which microorganisms improve barley salt tolerance remains lacking. In this review, the mechanisms of barley salt tolerance improvement by microorganisms are summarized, along with a discussion of existing problems in current research and areas of future research directions. In particular, with the development of sequencing technology and the great reduction of prices, the use of omics can not only comprehensively evaluate the role of microorganisms but also evaluate the impact of the microbiome on plants, which will provide us with many opportunities and challenges in this research area. [ABSTRACT FROM AUTHOR]

Obtaining homozygous lines from transgenic plants is an important step for phenotypic evaluations, but the selection of homozygous plants is time-consuming and laborious. The process would be significantly shortened if anther or microspore culture could be completed in one generation. In this study, we obtained 24 homozygous doubled haploid (DH) transgenic plants entirely by microspore culture from one T0 transgenic plant overexpressing the gene HvPR1 (pathogenesis-related-1). Nine of the doubled haploids grew to maturity and produced seeds. qRCR (quantitative real-time PCR) validation showed that the HvPR1 gene was expressed differentially even among different DH1 plants (T2) from the same DH0 line (T1). Phenotyping analysis suggested that the overexpression of HvPR1 inhibited nitrogen use efficiency (NUE) only under low nitrogen treatment. The established method of producing homozygous transgenic lines will enable the rapid evaluation of transgenic lines for gene function studies and trait evaluation. As an example, the HvPR1 overexpression of DH lines also could be used for further analysis of NUE-related research in barley. [ABSTRACT FROM AUTHOR]

Crop breeding for high nitrogen use efficiency (NUE) or tolerance to low nitrogen fertilization is thought to be an ideal solution to reduce the cost, carbon footprint, and other environmental problems caused by the excess use of nitrogen fertilizers. As a model plant for cereal crops, barley has many advantages, including good adaptability, a short growth period, and high natural stress resistance or tolerance. Therefore, research on improving NUE in barley is not only beneficial for nitrogen-efficient barley breeding but will also inform NUE improvement in other cereal crops. In this review, recent progress in understanding barley's response to nitrogen nutrition, evaluation of NUE or low-nitrogen tolerance, quantitative trait loci (QTL) mapping and gene cloning associated with improving NUE, and breeding of nitrogen-efficient barley is summarized. Furthermore, several biotechnological tools that could be used for revealing the molecular mechanisms of NUE or breeding for improving NUE in barley are introduced, including GWAS, omics, and gene editing. The latest research ideas in unraveling the molecular mechanisms of improving NUE in other crops are also discussed. Thus, this review provides a better understanding of improving the NUE of barley and some directions for future research in this area. [ABSTRACT FROM AUTHOR]

Background: Sucrose nonfermenting-1 (SNF1)-related protein kinases (SnRKs) play important roles in regulating metabolism and stress responses in plants, providing a conduit for crosstalk between metabolic and stress signalling, in some cases involving the stress hormone, abscisic acid (ABA). The burgeoning and divergence of the plant gene family has led to the evolution of three subfamilies, SnRK1, SnRK2 and SnRK3, of which SnRK2 and SnRK3 are unique to plants. Therefore, the study of SnRKs in crops may lead to the development of strategies for breeding crop varieties that are more resilient under stress conditions. In the present study, we describe the SnRK gene family of barley (Hordeum vulgare), the widespread cultivation of which can be attributed to its good adaptation to different environments. Results: The barley HvSnRK gene family was elucidated in its entirety from publicly-available genome data and found to comprise 50 genes. Phylogenetic analyses assigned six of the genes to the HvSnRK1 subfamily, 10 to HvSnRK2 and 34 to HvSnRK3. The search was validated by applying it to Arabidopsis (Arabidopsis thaliana) and rice (Oryza sativa) genome data, identifying 50 SnRK genes in rice (four OsSnRK1, 11 OsSnRK2 and 35 OsSnRK3) and 39 in Arabidopsis (three AtSnRK1, 10 AtSnRK2 and 26 AtSnRK3). Specific motifs were identified in the encoded barley proteins, and multiple putative regulatory elements were found in the gene promoters, with light-regulated elements (LRE), ABA response elements (ABRE) and methyl jasmonate response elements (MeJa) the most common. RNA-seq analysis showed that many of the HvSnRK genes responded to ABA, some positively, some negatively and some with complex time-dependent responses. Conclusions: The barley HvSnRK gene family is large, comprising 50 members, subdivided into HvSnRK1 (6 members), HvSnRK2 (10 members) and HvSnRK3 (34 members), showing differential positive and negative responses to ABA. [ABSTRACT FROM AUTHOR]

Background: Reducing the dependence of crop production on chemical fertilizer with its associated costs, carbon footprint and other environmental problems is a challenge for agriculture. New solutions are required to solve this problem, and crop breeding for high nitrogen use efficiency or tolerance of low nitrogen availability has been widely considered to be a promising approach. However, the molecular mechanisms of high nitrogen use efficiency or low-nitrogen tolerance in crop plants are still to be elucidated, including the role of long non-coding RNAs (lncRNAs). Results: In this study, we identified 498 lncRNAs in barley (Hordeum vulgare) landrace B968 (Liuzhutouzidamai), of which 487 were novel, and characterised 56 that were responsive to low-nitrogen stress. For functional analysis of differentially-expressed lncRNAs, the gene ontology (GO) and Kyoto Encyclopedia of Genes and Genomes (KEGG) enrichment of co-expressed and co-located protein-coding genes were analyzed, and interactions with annotated co-expressed protein coding genes or micro RNAs (miRNAs) were further predicted. Target mimicry prediction between differentially-expressed lncRNAs and miRNAs identified 40 putative target mimics of lncRNAs and 58 target miRNAs. Six differentially-expressed lncRNAs were further validated by qPCR, and one in particular showed consistent differential expression using both techniques. Expression levels of most of the lncRNAs were found to be very low, and this may be the reason for the apparent inconsistency between RNA-seq and qPCR data. Conclusions: The analysis of lncRNAs that are differentially-expressed under low-nitrogen stress, as well as their co-expressed or co-located protein coding genes and target mimics, could elucidate complex and hitherto uncharacterised mechanisms involved in the adaptation to low-nitrogen stress in barley and other crop plants. [ABSTRACT FROM AUTHOR]

The expression of genes encoding components of ABA and metabolic signalling pathways in developing barley endosperm and embryo was investigated. The genes included HvRCAR35_47387 and HvRCAR35_2538 (encoding ABA receptors), HvABI1d (protein phosphatase 2C), HvSnRK2.4, HvSnRK2.6 and HvPKABA1 (SnRK2-type protein kinases) and HvABI5 (ABA response element binding protein; AREBP), as well as two genes encoding SnRK1-type protein kinases. Both SnRK1 and SnRK2 phosphorylate AREBPs, but SnRK2 is activated by ABA whereas SnRK1 may be broken down. Multiple cereal AREBPs with two conserved SnRK1/2 target sites and another class of BZIP transcription factors with SnRK1/2 binding sites, including HvBLZ1, were identified. Barley grain (cv. Triumph) was sampled at 15, 20, 25 and 30 days post-anthesis (dpa). HvRCAR35_47387, HvABI1d, HvSnRK2.4 and HvABI5 were expressed highly in the endosperm but at much lower levels in the embryo. Conversely, HvPKABA1 and HvRCAR35_2538 were expressed at higher levels in the embryo than the endosperm, while HvSnRK2.6 was expressed at similar levels in both. HvRCAR35_47387, HvABI1d, HvSnRK2.4 and HvABI5 all peaked in expression in the endosperm at 20 dpa. A model is proposed in which ABA brings about a transition from a SnRK1-dominated state in the endosperm during grain filling to a SnRK2-dominated state during maturation. [ABSTRACT FROM AUTHOR]

Barley is the most salt-tolerant cereal crop. However, little attention has been paid to the salt-tolerant doubled haploids of barley derived from mutagenesis combined with isolated microspore culture. In the present study, barley doubled haploid (DH) line 20, which was produced by mutagenesis combined with isolated microspore culture, showed stably and heritably better salt tolerance than the wild type H30 in terms of fresh shoot weight, dry shoot weight, K+/Na+ ratio and photosynthetic characteristics. Transcriptome and metabolome analyses were performed to compare the changes in gene expression and metabolites between DH20 and H30. A total of 462 differentially expressed genes (DEGs) and 152 differentially accumulated metabolites (DAMs) were identified in DH20 compared to H30 under salt stress. Among the DAMs, fatty acids were the most accumulated in DH20 under salt stress. The integration of transcriptome and metabolome analyses revealed that nine key biomarkers, including two metabolites and seven genes, could distinguish DH20 and H30 when exposed to high salt. The pathways of linoleic acid metabolism, alpha-linolenic acid metabolism, glycerolipid metabolism, photosynthesis, and alanine, aspartate and glutamate metabolism were significantly enriched in DH20 with DEGs and DAMs in response to salt stress. These results suggest that DH20 may enhance resilience by promoting lipid metabolism, maintaining energy metabolism and decreasing amino acids metabolism. The study provided novel insights for the rapid generation of homozygous mutant plants by mutagenesis combined with microspore culture technology and also identified candidate genes and metabolites that may enable the mutant plants to cope with salt stress. [ABSTRACT FROM AUTHOR]

Although nitrogen (N) deficiency greatly affects N absorption and metabolism in barley, the global transcriptomic changes in morphological and physiological adaptation to altered N availability remains largely unclear. We conducted a comparative transcriptome analysis of roots in A9-29 (low N tolerant line of barley) and Hua 30 (low N-sensitive variety of barley) under low N conditions to elucidate the responses and the underlying molecular mechanism. The results demonstrated that the root architecture was strongly influenced and that the root morphological indexes (total root length, total root area surface, and root volume) were remarkably promoted in A9-29 compared to Hua30 under low N stress. The transcriptome analysis of roots identified 1779 upregulated differentially expressed genes (DEGs) and 1487 downregulated DEGs specifically expressed in A9-29 under low N stress. Specific DEGs in A9-29 were largely enriched in energy metabolism, lipid metabolism, and the metabolism of other amino acids. In addition, transcription factor genes ERFs and IAA-related genes were specifically expressed in A9-29. To conclude, this study could provide a foundation for improving low N tolerance in barley. [ABSTRACT FROM AUTHOR]

Somatic embryogenesis receptor-like kinases (SERKs), a subfamily of receptor-like kinases, showed important roles in plant response to abiotic and biotic stresses in addition to embryogenesis and organogenesis in numerous plant species. In the present study, three orthologs ofSERKgenes (HvSERK1/2/3) were isolated from barley, and their expression patterns duringin vitroculture of microspores as well as their responses to different stresses including salinity and powdery mildew were characterized. Sequence analysis suggested that threeHvSERKgenes were highly conserved in the grass family. Subcellular localization showed the HvSERK1 protein located on the plasma membrane. TheHvSERK1transcript was up-regulated during the microspore culture period, suggesting its roles in microspore embryogenesis.HvSERK1andHvSERK3showed the highest expression level in the leaves; however no difference was detected forHvSERK2expression in different plants’ tissues. Under salt stress, all threeHvSERKgenes were quickly induced in microspore-derived embryogenic calli, whereas onlyHvSERK1was up-regulated in the roots of barley seedlings. Moreover, onlyHvSERK2was induced in the barley leaves upon powdery mildew inoculation. These results suggest that barleySERKgenes may participate in barley microspores’ development and plant response toward salt and fungal stress, and the function of them has some evolutionary changes. [ABSTRACT FROM PUBLISHER]