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Domestication syndrome, selection pressure, and modern plant breeding programs have reduced the genetic diversity of the wheat germplasm. For the genetic gains of breeding programs to be sustainable, plant breeders require a diverse gene pool to select genes for resistance to biotic stress factors, tolerance to abiotic stress factors, and improved quality and yield components. Thus, old landraces, subspecies and wild ancestors are rich sources of genetic diversity that have not yet been fully exploited, and it is possible to utilize this diversity. Compared with durum wheat, tetraploid wheat subspecies have retained much greater genetic diversity despite genetic drift and various environmental influences, and the identification and utilization of this diversity can make a greater contribution to the genetic enrichment of wheat. In addition, using the pre-breeding method, the valuable left-behind alleles in the wheat gene pool can be re-introduced through hybridization and introgressive gene flow to create a sustainable opportunity for the genetic gain of wheat. This review provides some insights about the potential of tetraploid wheats in plant breeding and the genetic gains made by them in plant breeding across past decades, and gathers the known functional information on genes/QTLs, metabolites, traits and their direct involvement in wheat resistance/tolerance to biotic/abiotic stresses. • Genetic erosion during domestication and modern breeding programs has led to reduce the genetic diversity of wheat. • Genetic structure of wheat can be improved through genetic enrichment by diversifying allelic combinations. • Tetraploid wheat subspecies are an excellent gene pool for improving tolerance to biotic and abiotic stresses. • Utilizing genes from tetraploid wheat subspecies provide great potential for increasing grain yield and quality of durum wheat. • Synthetic hexaploid wheat is an effective genetic resource to transfer important genes from wild relatives to common wheat. [ABSTRACT FROM AUTHOR]

Clustered Regularly Interspaced Short Palindromic Repeats (CRISPR) technologies have been implemented in recent years in the genome editing of eukaryotes, including plants. The original system of knocking out a single gene by causing a double-strand break (DSB), followed by non-homologous end joining (NHEJ) or Homology-directed repair (HDR) has undergone many adaptations. These adaptations include employing CRISPR/Cas9 to upregulate gene expression or to cause specific small changes to the DNA sequence of the gene-of-interest. In plants, multiplexing, i.e. , inducing multiple changes by CRISPR/Cas9, is extremely relevant due to the redundancy of many plant genes, and the time- and labor-consuming generation of stable transgenic plant lines via crossing. Here we discuss relevant examples of various traits, such as yield, biofortification, gluten content, abiotic stress tolerance, and biotic stress resistance, which have been successfully manipulated using CRISPR/Cas9 in plants. While existing studies have primarily focused on proving the impact of CRISPR/Cas9 on a single trait, there is a growing interest among researchers in creating a multi-stress tolerant wheat cultivar 'super wheat', to commercially and sustainably enhance wheat yields under climate change. Due to the complexity of the technical difficulties in generating multi-target CRISPR/Cas9 lines and of the interactions between stress responses, we propose enhancing already commercial local landraces with higher yield traits along with stress tolerances specific to the respective localities, instead of generating a general 'super wheat'. We hope this will serve as the sustainable solution to commercially enhancing crop yields under both stable and challenging environmental conditions. • In the past decade, CRISPR/Cas9 systems have been developed extensively, for eukaryotic genome editing. • CRISPR/Cas9 has been utilized for trait adaptation in Triticum aestivum mainly by knocking out a single gene. • It is necessary to begin implementing CRISPR/Cas9 in T. aestivum in a multi-trait approach. • We suggest enhancing local cultivars for specific traits rather than pursuing a 'super wheat' tolerant to all stresses. [ABSTRACT FROM AUTHOR]

The transition of food crops from the vegetative to reproductive stages is an important process that affects the final yield. Despite extensive characterization of E3 ligases in model plants, their roles in wheat development remain unknown. In this study, we elucidated the molecular function of wheat TaATL1 (Arabidopsis thaliana Toxicos EN Levadura), which acts as a negative regulator of flowering time and cell division. TaATL1 amino acid residues contain a RING domain and exist mainly in a beta-turn form. The expression level of TaATL1 was highly reduced during the transition from vegetative to reproductive stages. TaATL1 is localized in the nucleus and exhibits E3 ligase activity. Transgenic Arabidopsis plants, in which the TaATL1 gene is constitutively overexpressed under the control of the cauliflower mosaic virus 35 S promoter, exhibited regulation of cell numbers, thereby influencing both leaf and root growth. Moreover, TaATL1 overexpression plants showed a late-flowering phenotype compared to wild-type (WT) plants. Following transcriptome analysis, it was discovered that 1661 and 901 differentially expressed genes were down- or up- regulated, respectively, in seedling stages between WT and TaATL1 overexpression. TaATL1 transcripts are involved in cell division, flowering, and signaling. Overall, our findings demonstrated that the regulatory mechanism of wheat TaATL1 gene plays a significant role in cell division-mediated flowering in Arabidopsis. • TaATL1 was significantly down-regulated in early reproductive phases. • TaATL1 possessed E3 ligase activity and transiently expressed in the nucleus. • TaATL1 overexpression plants exhibited late-flowering phenotype and negatively regulated cell division. [ABSTRACT FROM AUTHOR]

Monodehydroascorbate reductase (MDHAR) is a crucial enzymatic antioxidant of the ascorbate-glutathione pathway involved in reactive oxygen species scavenging. Herein, we identified 15 TaMDHAR genes in bread wheat. Phylogenetic analysis revealed their clustering into three groups, which are also related to the subcellular localization in the peroxisome matrix, peroxisome membrane, and chloroplast. Each TaMDHAR protein consisted of two conserved domains; Pyr_redox and Pyr_redox_2 of the pyridine nucleotide disulfide oxidoreductase family. The occurrence of diverse groups of cis -regulatory elements in the promoter region and their interaction with numerous transcription factors suggest assorted functions of TaMDHARs in growth and development and in light, phytohormones, and stress responses. Expression analysis in various tissues further revealed their importance in vegetative and reproductive development. In addition, the differential gene expression and enhanced enzyme activity during drought, heat, and salt treatments exposed their role in abiotic stress response. Interaction of MDHARs with various antioxidant enzymes and biochemicals related to the ascorbate-glutathione cycle exposed their synchronized functioning. Interaction with auxin indicated the probability of cross-talk between antioxidants and auxin signaling. The miR168a, miR169, miR172 and others interaction with various TaMDHARs further directed their association with developmental processes and stress responses. The current study provides extensive information about the importance of TaMDHARs , moreover, the precise role of each gene needs to be established in future studies. • Monodehydroascorbate reductase (MDHAR) is an enzymatic antioxidant involved in ROS scavenging during stress conditions. • The phylogenetic analysis, gene structure, and physicochemical properties of TaMDHARs revealed their conserved nature. • The expression profiling and enzyme assay suggested their involvement in stress response. • The miRNA, TF interaction and cis- regulatory elements analyses revealed their involvement in various developmental processes and stress response. [ABSTRACT FROM AUTHOR]

Recent progress in the characterization of two groups of genes responsible for natural differences in frost tolerance in wheat and barley is reviewed here. The first group includes the vernalization genes that delay flowering until the end of the winter and protect sensitive floral primordia. This process is regulated mainly by differences in the regulatory regions of VRN1 and VRN3 genes or in the coding regions of VRN2. The second group includes a set of tandemly duplicated CBF (C-repeat Binding Factors) transcription factors at the FR2 (Frost Resistance 2) locus. CBF transcription factors are known regulators of the COR genes (COld Regulated genes) which are induced by cold and confer tolerance to subsequent freezing temperatures (acclimation). Natural differences in frost tolerance in both wheat and barley have been mapped to the FR2 locus, and are associated with differences in threshold induction temperatures and/or transcript levels of several CBF genes. Higher threshold induction temperatures result in earlier up-regulation of COR genes during the fall, whereas higher induction levels by cold are associated with faster cold acclimation rates. Both processes result in longer acclimation periods and improved frost tolerance. Increases in VRN1 transcript levels in the leaves are associated with reduced responsiveness of CBF and COR genes to cold and with the end of the acclimation period. Therefore, delays in the induction of VRN1 and in the transition to the reproductive stage can extend the acclimation period and improve frost tolerance. These observations suggest that the vernalization and cold acclimation regulatory gene networks are interconnected.

Glutathione peroxidases (GPXs) are known to play an essential role in guarding cells against oxidative stress by catalyzing the reduction of hydrogen peroxide and organic hydroperoxides. The current study aims functional characterization of the TaGPX1-D gene of bread wheat (Triticum aestivum) for salinity and osmotic stress tolerance. To achieve this, we initially performed the spot assays of TaGPX1-D expressing yeast cells. The growth of recombinant TaGPX1-D expressing yeast cells was notably higher than the control cells under stress conditions. Later, we generated transgenic Arabidopsis plants expressing the TaGPX1-D gene and investigated their tolerance to various stress conditions. The transgenic plants exhibited improved tolerance to both salinity and osmotic stresses compared to the wild-type plants. The higher germination rates, increased antioxidant enzymes activities, improved chlorophyll, carotenoid, proline and relative water contents, and reduced hydrogen peroxide and MDA levels in the transgenic lines supported the stress tolerance mechanism. Overall, this study demonstrated the role of TaGPX1-D in abiotic stress tolerance, and it can be used for improving the tolerance of crops to environmental stressors, such as salinity and osmotic stress in future research. • TaGPX1-D expressing yeast cells exhibited higher growth than the control cells under stress conditions. • TaGPX1-D enhances salinity and osmotic stress tolerance in transgenic Arabidopsis. • Transgenic lines exhibited higher germination rates, improved chlorophyll and carotenoid levels. • Overexpression of TaGPX1-D enhanced antioxidant activity and proline content and reduced hydrogen peroxide and MDA accumulation during stress. [ABSTRACT FROM AUTHOR]

Iron (Fe), an essential micronutrient, participates in photosynthesis, respiration, and many other enzymatic reactions. Cadmium (Cd), by contrast, is a toxic element to virtually all living organisms. Both Fe deficiency and Cd toxicity severally impair crop growth and productivity, finally leading to human health issues. Understanding how plants control the uptake and homeostasis of Fe and combat Cd toxicity thus is mandatory to develop Fe-enriched but Cd-cleaned germplasms for human beings. Recent studies in Arabidopsis and rice have revealed that IRON MAN (IMA) peptides stand out as a key regulator to respond to Fe deficiency by competitively interacting with a ubiquitin E3 ligase, thus inhibiting the degradation of IVc subgroup bHLH transcription factors (TFs), mediated by 26 S proteasome. Elevated expression of IMA confers tolerance to Cd stress in both Arabidopsis and wheat by activating the iron deficiency response. Here, we discuss recent breakthroughs that IMA peptides function in the Fe-deficiency response to attain Fe homeostasis and combat Cd toxicity as a potential candidate for phytoremediation. • The function of IMAs might be conserved across land species. • IMA peptides win the competitive interaction with BTS/HRZ ubiquitin E3 ligase thus alleviating the degradation of IVc bHLH TFs by these ligases. • Elevated expression of IMA peptides activates Fe deficiency response. • High Fe contents will decrease Cd absorption but increase Cd sequestration. • Recent findings pave the way for use of IMA peptides to design gold crops of a concurrent biofortification of essential microelements but toxic Cd mitigation. [ABSTRACT FROM AUTHOR]

Homeodomain proteins encoded by BEL1 - and KNAT1 -type genes are ubiquitously distributed across plant species and play important roles in growth and development, whereby a comprehensive investigation of their molecular interactions and potential functions in wheat is of great significance. In this study, we systematically investigated the phylogenetic relationships, gene structures, conserved domains, and cis -acting elements of 34 TaBEL and 34 TaKNAT genes in the wheat genome. Our analysis revealed these genes evolved under different selective pressures and showed variable transcript levels in different wheat tissues. Subcellular localization analysis further indicated the proteins encoded by these genes were either exclusively located in the nucleus or both in the nucleus and the cytoplasm. Additionally, a comprehensive protein-protein interaction network was constructed with representative genes in which each TaBEL or TaKNAT proteins interact with at least two partners. The evaluation of wheat mutants identified key genes, including TaBEL-5B , TaBEL-4A.4 , and TaKNAT6 , which are involved in grain-related traits. Finally, haplotype analysis suggests TaKNAT-6B is associated with grain-related traits and is preferentially selected among a large set of wheat accessions. Our study provides important information on BEL1 - and KNAT1 -type gene families in wheat, and lays the foundation for functional research in the future. • Selective pressures shaped composition and structure of TaBELs and TaKNATs. • Extensive protein interactions between wheat BEL1 - and KNAT1 -type members. • Mutant analysis associates key homeobox genes with wheat kernel-related traits. • Haplotypes of TaKNAT-6B are preferentially selected in wheat accessions. [ABSTRACT FROM AUTHOR]

Unpredicted variability in climate change on the planet is associated with frequent extreme high-temperature events impacting crop yield globally. Wheat is an economically and nutritionally important crop that fulfils global food requirements and each degree rise in temperature results in ∼6% of its yield reduction. Thus, understanding the impact of climate change, especially the terminal heat stress on global wheat production, becomes critically important for policymakers, crop breeders, researchers and scientists to ensure global food security. This review describes how wheat perceives heat stress and induces stress adaptation events by its morpho-physiological, phenological, molecular, and biochemical makeup. Temperature above a threshold level in crop vicinity leads to irreversible injuries, viz. destruction of cellular membranes and enzymes, generation of active oxygen species, redox imbalance, etc. To cope with these changes, wheat activates its heat tolerance mechanisms characterized by hoarding up soluble carbohydrates, signalling molecules, and heat tolerance gene expressions. Being vulnerable to heat stress, increasing wheat production without delay seeks strategies to mitigate the detrimental effects and provoke the methods for its sustainable development. Thus, to ensure the crop's resilience to stress and increasing food demand, this article circumscribes the integrated management approaches to enhance wheat's performance and adaptive capacity besides its alleviating risks of increasing temperature anticipated with climate change. Implementing these integrated strategies in the face of risks from rising temperatures will assist us in producing sustainable wheat with improved yield. • The rising temperature caused by climate change leads to abiotic stress harms crop growth and productivity. • Heat stress (HS) is limiting factor that declines and affects wheat production globally. • Perception and adaptation mechanism induced in response to stress explains the impact of climate change on crop "wheat". • Need of crop resilience to sustain the productivity matching the rising food demand. • Recently and successfully used management strategies improve stress tolerance and yield to cope with the existing/upcoming environmental changes. [ABSTRACT FROM AUTHOR]

Application of the mlo -based resistance in barley against powdery mildew attacks is a major success in crop breeding, since it confers durable disease resistance. Resistance caused by mutations in the Mlo gene seems to be ubiquitous across a range of species. This work addresses the introduction of mlo -based resistance into hexaploid wheat, which is complicated by the occurrence of three homoeologous genes: Mlo-A1, Mlo-B1 and Mlo-D1. EMS-generated mutant plants were screened for mutations in the three homoeologues. We selected and combined 6, 8, and 4 mutations, respectively, to obtain triple homozygous mlo mutant lines. Twenty-four mutant lines showed highly effective resistance towards attack by the powdery mildew pathogen under field conditions. All 18 mutations appeared to contribute to resistance; however, they had different effects on the occurrence of symptoms such as chlorotic and necrotic spots, which are pleiotropic to the mlo -based powdery mildew resistance. We conclude that to obtain highly effective powdery mildew resistance in wheat and to avoid detrimental pleiotropic effects, all three Mlo homoeologues should be mutated; however, at least one of the mutations should be of the weaker type in order to alleviate strong pleiotropic effects from the other mutations. • A range of different mutations in the three Mlo homoeologues in wheat can contribute to powdery mildew resistance. • Simultaneous mutations in all three Mlo homoeologues provide highly effective powdery mildew resistance in the field. • Strong mutations in all three Mlo homoeologues lead to severe pleiotropic effects, i.e. necrosis/chlorosis. • Weak mutations in one Mlo homoeologue can alleviate the pleiotropic effects from strong mutations in other homoeologues. • mlo mutations can be combined in wheat to obtain highly effective powdery mildew resistance without pleiotropic effects. [ABSTRACT FROM AUTHOR]

• Premature termination mutation of COBRA-like gene TmBr1 leads to tissue brittleness. • TmBr1 modulates cell cellulose content of cell walls. • TmBr1 probably modulates synthesis of noncellulosic polysaccharide pectin. • MapRseq cloning approach is optimal for cloning genes in complicated genomes. Plant tissue brittleness is related to cellular structure and lodging. MED0031 is a mutant identified previously from ethyl methane sulfonate treatment of diploid wheat accession TA2726, showing brittleness in both stem and leaf. In microscopic and histological observations, the mutant was found to have less large vascular bundles per unit area, a thinner sclerenchyma cell wall, and a broader parenchyma, compared with the wild type. The mutated gene, TmBr1 , was mapped to a 0.056 cM interval on chromosome 5Am. This gene was cloned using a MapRseq approach that searched the candidate gene through combination of the prior target gene mapping information with SNP calling and discovery of differentially expressed genes from RNA_seq data of the wild type and a BC 3 F 2 bulk showing the mutant phenotype. TmBr1 encodes a COBL protein and a nonsense mutation within the region coding for the conserved COBRA domain caused premature translation termination. Introduction of TmBr1 to Arabidopsis AtCOBL4 mutant rescued the phenotype, demonstrating their functional conservation. Apart from the effect on cellulose content, the TmBr1 mutation might modulate synthesis of noncellulosic polysaccharide pectin as well. Application of the MapRseq approach to isolation of genes present in recombination cold spots and complicated genomes was discussed. [ABSTRACT FROM AUTHOR]

• Wheat carry mutations in the BTR1-A and BTR1-B genes affecting spike shattering. • We probed the geographical provenances of these mutations via haplotype analyses. • The progenitor haplotypes were identified in a collection of wild wheat accessions. • A single Southern Levant accession harbored both BTR1 progenitor haplotypes. • This imply that Southern Levant played an important role in wheat domestication. Wheat domestication was a milestone in the rise of agrarian societies in the Fertile Crescent. As opposed to the freely dispersing seeds of its tetraploid progenitor wild emmer, the hallmark trait of domesticated wheat is intact, harvestable spikes. During domestication, wheat acquired recessive loss-of-function mutations in the Brittle Rachis 1 genes, both in the A genome (BTR1-A) and B genome (BTR1-B). In this study, we probe the geographical provenances of these mutations via haplotype analyses of a collection of wild and domesticated accessions. Our results show that the precursor of the domesticated haplotype of BTR1-A was detected in 32% of the wild accessions gathered throughout the Levant, from central Israel to central Turkey. In contrast, the precursor of the domesticated haplotype of BTR1-B , which carries a distinct 11 bp deletion in the promoter region, was found in only 10% of the tested wild accessions, all from the Southern Levant. Moreover, we identified of a single wild emmer accession in Southern Levant that carries the progenitor haplotypes for both BTR1-A and BTR1-B genes. These observations suggest that at least part of the emmer domestication process occurred in Southern Levant, contrary to the widely held view that the northern part of the Fertile Crescent was the center of wheat domestication. [ABSTRACT FROM AUTHOR]

Highlights • An unmanned aerial vehicle (UAV) was optimized and used for non-destructive high-throughput phenotyping of NDVI. • UAV-NDVI measurements were highly consistent with ground data captured by a handheld Greenseeker. • UAV-NDVI explained significant variations in biomass and grain yield. • UAV-NDVI was accurate and can be used for selection of high yielding genotypes during grain-filling stages in large breeding programs. Abstract Wheat improvement programs require rapid assessment of large numbers of individual plots across multiple environments. Vegetation indices (VIs) that are mainly associated with yield and yield-related physiological traits, and rapid evaluation of canopy normalized difference vegetation index (NDVI) can assist in-season selection. Multi-spectral imagery using unmanned aerial vehicles (UAV) can readily assess the VIs traits at various crop growth stages. Thirty-two wheat cultivars and breeding lines grown in limited irrigation and full irrigation treatments were investigated to monitor NDVI across the growth cycle using a Sequoia sensor mounted on a UAV. Significant correlations ranging from R2 = 0.38 to 0.90 were observed between NDVI detected from UAV and Greenseeker (GS) during stem elongation (SE) to late grain gilling (LGF) across the treatments. UAV-NDVI also had high heritabilities at SE (h2 = 0.91), flowering (F)(h2 = 0.95), EGF (h2 = 0.79) and mid grain filling (MGF) (h2 = 0.71) under the full irrigation treatment, and at booting (B) (h2 = 0.89), EGF (h2 = 0.75) in the limited irrigation treatment. UAV-NDVI explained significant variation in grain yield (GY) at EGF (R2 = 0.86), MGF (R2 = 0.83) and LGF (R2 = 0.89) stages, and results were consistent with GS-NDVI. Higher correlations between UAV-NDVI and GY were observed under full irrigation at three different grain-filling stages (R2 = 0.40, 0.49 and 0.45) than the limited irrigation treatment (R2 = 0.08, 0.12 and 0.14) and GY was calculated to be 24.4% lower under limited irrigation conditions. Pearson correlations between UAV-NDVI and GY were also low ranging from r = 0.29 to 0.37 during grain-filling under limited irrigation but higher than GS-NDVI data. A similar pattern was observed for normalized difference red-edge (NDRE) and normalized green red difference index (NGRDI) when correlated with GY. Fresh biomass estimated at late flowering stage had significant correlations of r = 0.30 to 0.51 with UAV-NDVI at EGF. Some genotypes Nongda 211, Nongda 5181, Zhongmai 175 and Zhongmai 12 were identified as high yielding genotypes using NDVI during grain-filling. In conclusion, a multispectral sensor mounted on a UAV is a reliable high-throughput platform for NDVI measurement to predict biomass and GY and grain-filling stage seems the best period for selection. [ABSTRACT FROM AUTHOR]

Highlights • Multiple genes have been associated with pre-harvest sprouting (PHS). • Development of perfect markers underpinning PHS QTLs is necessary for efficient selection. • TaMFT-3A allelic variation drastically impacts PHS. • The ability to manipulate PHS will be enhanced by understanding how all genes that impact PHS together impact wheat dormancy. Abstract Pre-harvest sprouting (PHS) of wheat (Triticum aestivum L.) is an important phenomenon that results in weather dependent reductions in grain yield and quality across the globe. Due to the large annual losses, breeding PHS resistant varieties is of great importance. Many quantitative trait loci have been associated with PHS and a number of specific genes have been proven to impact PHS. TaPHS1 , TaMKK3, Tamyb10, and TaVp1 have been shown to have a large impact on PHS susceptibility while many other genes such as TaSdr , TaQSd , and TaDOG1 have been shown to account for smaller, but significant, proportions of variation. These advances in understanding the genetics behind PHS are making molecular selection and loci stacking viable methods for affecting this quantitative trait. The current review article serves to provide a brief synthesis of recent advances regarding PHS, as well as provide unique insight into the genetic mechanisms governing PHS in bread wheat. [ABSTRACT FROM AUTHOR]

Highlights • We developed an improved, robust method to eliminate repetitive genomic elements. • A > 50% increase in gene-derived reads was obtained in a diploid wheat cultivar. • Our method, applicable to any species, is superior to target-enrichment sequencing. Abstract Genome size and complexity often present major challenges to genome-based approaches in crop plants and other agricultural species. For instance, repetitive sequences comprise 80% to 90% of the genome of hexaploid wheat, which has a haploid genome size of approximately 17 Gb. In this study, we developed an improved design and procedure for short-read library preparation that uses a modified adaptor and duplex-specific nuclease (DSN) for the efficient elimination of highly repeated sequence elements within genomes. The improved adapter, which has a hairpin-like form for stability, was constructed from truncated sequences adjacent to the original Illumina TruSeq adapter and can be converted to a full-length adapter structure during PCR amplification. Using the hairpin-structured adaptor, we prepared randomly sheared genomic libraries from rice and diploid, tetraploid, and hexaploid wheat cultivars and evaluated the efficiency of DSN for the enzymatic depletion of repetitive elements. According to real-time quantitative PCR analysis, the relative abundances of 18S and 25S ribosomal DNA decreased respectively to 1.15% and 3.54% in rice and 1.70%–1.95% and 14.71%–20.01% in the three wheat cultivars. Whole-genome sequencing analysis of a diploid wheat cultivar, KU104-1, indicated that DSN treatment with the designed hairpin-structured adapter dramatically reduced highly repetitive elements, such as Ty1-Copia and Ty3-Gypsy retrotransposons and DNA transposons, within the genome, while sequencing reads derived from low-copy genes and protein coding sequences increased more than 50%. Our new procedure should be useful not only for wheat genomes but also for other agricultural plant species with relatively large and complex genomes. [ABSTRACT FROM AUTHOR]

Highlights • Cytokinin metabolism was involved in delayed flag leaf senescence in a wheat stay-green mutant, tasg1. • N metabolism was involved in delayed flag leaf senescence in a wheat stay-green mutant, tasg1. • The cisZOGT1 gene has an important role in regulating wheat leaf senescence by regulating cytokinin and N metabolism. Abstract In the present study on a wheat stay-green mutant, tasg1 , we found that its delayed senescence at the late filling stage was related to the high cytokinin (CK) and N contents. RNA sequencing suggested that several genes may be responsible for the different senescence processes between wild-type (WT) and tasg1 plants. WT and tasg1 seedlings were treated with NH 4 NO 3 , lovastatin, and 6-benzylaminopurine (BAP), and the results suggested that the feedback of CK with N content regulated the leaf senescence in the tasg1 plants. Furthermore, a knock-out of the candidate gene cisZOGT1 (catalytic O-glucosylation in cis -zeatin) in the wheat mutant pool 'Kronos' exhibited delayed senescence at the late grain filling stage. Overall, our results suggested the cisZOGT1 gene has an important role in regulating wheat leaf senescence by regulating CK and N metabolism. At the same time, CK and N metabolism involved in delayed flag leaf senescence of tasg1 may be by a feedback pattern. [ABSTRACT FROM AUTHOR]

Highlights • Wheat genotypes vary in their seedling emergence force. • Seedling emergence force of wheat genotypes reduces with increasing sodicity. • Seedling emergence force is correlated to the cross sectional area of the hypocotyl. Abstract Surface crusting of sodic soils is a major problem in the semi-arid tropics when rapid drying after sowing follows light showers, leading to reduced seedling emergence and grain yield. The magnitude of the force exerted by germinating seeds affects the ability of the seedlings to rupture the crust and emerge. This study aimed to determine whether the seed germination and seedling emergence force of wheat (Triticum aestivum L.) seedlings vary among different genotypes at different sodicity levels. Germination and emergence force of seedlings of four wheat genotypes was determined in assays using four solutions with sodium adsorption ratio (SAR) values ranging from 0 to 60. Seed germination and seedling emergence force varied between genotypes at different sodicity levels, with the emergence force of the coleoptile correlated to the cross sectional area of the hypocotyl. The results suggest that the selection of wheat genotypes with rapid germination, higher seedling emergence force and larger hypocotyl cross sectional area, offers a strategy to improve seedling emergence in crusted sodic soils. [ABSTRACT FROM AUTHOR]

Drought stress is a major factor that limits the yield and quality in wheat. In this study, we identified an orthologue of the rice gene OsDIS1 ( Oryza sativa drought-induced SINA protein 1) in wheat ( Triticum aestivum L.) called TaDIS1 . TaDIS1 encodes a putative 301 amino acid protein with a C3HC4 RING finger conserved domain at the N-terminal and a SINA domain at the C-terminal. TaDIS1 contains three exons and two introns. qRT-PCR analysis showed that TaDIS1 expression was induced by PEG6000, NaCl, and abscisic acid (ABA) treatment. We generated TaDIS1 -overexpressing transgenic Arabidopsis lines. Under drought stress conditions, the transgenic Arabidopsis plants had a lower germination rate, relative water content, and proline contents, with higher water loss, chlorophyll loss, relative electrical conductivity, and malondialdehyde contents compared with the wild type. The antioxidant enzyme (superoxide dismutase, peroxidase, and catalase) activity levels were lower in the transgenic plants. The TaDIS1 -overexpressing plants had shorter roots with greater growth inhibition in response to mannitol treatment than the wild type, with increased hypersensitivity to ABA during seed germination and early seedling growth. The expression of stress-related genes in transgenic plants under drought stress suggests that TaDIS1 may function negatively in drought stress by regulating the stress response-related genes. [ABSTRACT FROM AUTHOR]

As the trace signal molecules widely existing in plants, plant hormones can regulate physiological responses of plants at low concentrations. At present, the effect of plant endogenous hormones on wheat male fertility has attracted attention, but the molecular mechanism underlying fertility regulation is unclear. Given this, the anthers of five isonuclear alloplasmic male sterile lines and their maintainer line were RNA-sequenced. A gene TaGA-6D encoding gibberellin (GA) regulated protein was isolated, which was located to the nucleus, cell wall and/or cell membrane, and predominantly highly expressed in the anther of Ju706A, a male sterile line with Aegilops juvenalis cytoplasm. By spraying assay of GA with different concentrations on fertility line Ju706R, it was found that with the increase of exogenous GA concentration, the content of endogenous GA and expression level of TaGA-6D in anther gradually increased, and the fertility decreased. However, silencing of TaGA-6D partially restore the fertility of Ju706R sprayed with 1000 ng/μl GA, and indicating that gibberellin can promote the expression of TaGA-6D and negatively regulates the fertility of wheat with Aegilops juvenalis cytoplasm, which providing new insights for understanding hormone regulation of male fertility in wheat. • A gibberellin (GA)-mediated family protein gene, TaGA-6D, negatively regulates fertility of wheat. • TaGA-6D predominantly highly expressed in the anther of Aegilops juvenalis cytoplasmic male sterile line. • Exogenous gibberellin promoted the expression of TaGA-6D. • Exogenous gibberellin induces male sterility, while silencing of TaGA-6D could partially restore the fertility of wheat. [ABSTRACT FROM AUTHOR]

Conditions that disrupt protein folding, such as heat stress, can overwhelm the capacity of cells to fold proteins, thus causing endoplasmic reticulum (ER) stress. In Arabidopsis thaliana and other plants, inositol-requiring enzyme-1 mediated unconventional splicing of bZIP60 plays a crucial role in the heat and ER stress responses. However, little is known about this pathway in wheat ( Triticum aestivum ), especially its importance in heat tolerance. Here, we found that heat stress induced upregulation and unconventional splicing of TabZIP60 occurred in wheat seedlings. Constitutive expression of the spliced form of TabZIP60 ( TabZIP60s ) enhanced heat tolerance in Arabidopsis, but overexpression of the unspliced form ( TabZIP60u ) did not. RNA-sequencing analysis revealed ER stress related genes involved in heat responses in TabZIP60s- overexpression transgenic Arabidopsis. Chromatin immunoprecipitation-qPCR showed that TabZIP60s directly binds to 17 target genes including AtbZIP60 . Also, the 26S proteasome pathway post-translationally regulates TabZIP60s levels during heat stress responses. Our findings suggest that unconventional splicing of TabZIP60 could contribute to heat tolerance in transgenic plants by modulating the expression of ER stress-related genes. [ABSTRACT FROM AUTHOR]

Water deficit is one of the major factors limiting crop productivity worldwide. Plant roots play a key role in uptaking water, perceiving and transducing of water deficit signals to shoot. Although the mechanisms of drought-tolerance have been reported recently, the transcriptional regulatory network of wheat root response to water stress has not been fully understood. In this study, drought-tolerant cultivar JM-262 and susceptible cultivar LM-2 are planted to characterize the root transcriptional changes and physiological responses to water deficit. A total of 8197 drought tolerance-associated differentially expressed genes (DEGs) are identified, these genes are mainly mapped to carbon metabolism, flavonoid biosynthesis, and phytohormone signal transduction. The number and expression level of DEGs involved in antioxidative and antiosmotic stresses are more enhanced in JM-262 under water stress. Furthermore, we find the DEGs related to root development are much more induced in JM-262 in phytohormone signal transduction and carbon metabolism pathway. In conclusion, JM-262 may alleviate the damage of drought by producing more osmoprotectants, ROS scavengers, biomass and energy. Interestingly, hormone signaling and cross-talk probably play an important role in promoting JM-262 greater root systems to take up more water, higher capabilities to induce more drought-related DEGs and higher resisitance to oxidative stresse. [ABSTRACT FROM AUTHOR]

The utilization of DNA molecular markers in plant breeding to maximize selection response via marker-assisted selection (MAS) and genomic selection (GS) has revolutionized plant breeding. A key factor affecting GS applicability is the choice of molecular marker platform. Genotyping-by-sequencing scored SNPs (GBS-scored SNPs) provides a large number of markers, albeit with high rates of missing data. Array scored SNPs are of high quality, but the cost per sample is substantially higher. The objectives of this study were 1) compare GBS-scored SNPs, and array scored SNPs for genomic selection applications, and 2) compare estimates of genomic kinship and population structure calculated using the two marker platforms. SNPs were compared in a diversity panel consisting of 299 hard winter wheat ( Triticum aestivum L.) accessions that were part of a multi-year, multi-environments association mapping study. The panel was phenotyped in Ithaca, Nebraska for heading date, plant height, days to physiological maturity and grain yield in 2012 and 2013. The panel was genotyped using GBS-scored SNPs, and array scored SNPs. Results indicate that GBS-scored SNPs is comparable to or better than Array-scored SNPs for genomic prediction application. Both platforms identified the same genetic patterns in the panel where 90% of the lines were classified to common genetic groups. Overall, we concluded that GBS-scored SNPs have the potential to be the marker platform of choice for genetic diversity and genomic selection in winter wheat. [ABSTRACT FROM AUTHOR]

Cadmium (Cd) is a severe and toxic heavy metal pollutant that affects plant growth and development. In this study, we found that the expression of an expansin gene, TaEXPA2 , was upregulated in wheat leaves under CdCl 2 toxicity. We characterized the involvement of TaEXPA2 in conferring Cd tolerance. Tobacco plants overexpressing TaEXPA2 showed higher germination rate, root elongation, and biomass accumulation compared to the wild-type (WT) plants upon CdCl 2 treatment. The improved photosynthetic parameters and lesser cellular damage in transgenic plants exposed to Cd compared to that in the WT plants suggest that TaEXPA2 overexpression improves Cd tolerance in plants. Furthermore, we noticed that Cd was efficiently effluxed out of the cytoplasm in the transgenic plants owing to the enhanced activities of H + -ATPase, V-ATPase, and PPase, which helped in conferring Cd tolerance. Moreover, Cd concentration and ROS accumulation were lower in the transgenic plants than in WT plants as a consequence of enhanced antioxidant enzyme activities in the former. In addition, atexpa2 , an Arabidopsis mutant, exhibited lower biomass and shorter primary root compared to its WT under Cd toxicity; however, the phenotype was recovered upon expression of TaEXPA2 in these mutants. Our results demonstrate that TaEXPA2 confers tolerance to Cd toxicity. The changed absorption/transportation of Cd and the antioxidative capacity may be involved in the improved tolerance of the transgenic plants with overexpression of TaEXPA2 to CdCl 2 toxicity. [ABSTRACT FROM AUTHOR]

Polyamine metabolism is in relation with several metabolic pathways and linked with plant hormones or signalling molecules; in addition polyamines may modulate the up- or down-regulation of gene expression. However the precise mechanism by which polyamines act at the transcription level is still unclear. In the present study the modifying effect of putrescine pre-treatment has been investigated using the microarray transcriptome profile analysis under the conditions where exogenous putrescine alleviated osmotic stress in wheat plants. Pre-treatment with putrescine induced the unique expression of various general stress-related genes. Although there were obvious differences between the effects of putrescine and polyethylene glycol treatments, there was also a remarkable overlap between the effects of putrescine and osmotic stress responses in wheat plants, suggesting that putrescine has already induced acclimation processes under control conditions. The fatty acid composition in certain lipid fractions and the antioxidant enzyme activities have also been specifically changed under osmotic stress conditions or after treatment with putrescine. [ABSTRACT FROM AUTHOR]

Gibberellins (GA) are involved in seed development and E3 ubiquitin-ligases actively participate in GA perception and signal transduction. TaGW2-6A encodes a RING E3 ubiquitin-ligase that negatively regulates grain size. Therefore, Chinese Spring (CS) and its TaGW2-6A allelic variants (NIL31) were investigated to elucidate the relative contribution of GA to the regulation of seed development in wheat. The expression levels of GA biosynthesis and response genes were higher in NIL31 than CS, especially those of GA 3-oxidase and GASA4 . The expression of TaGW2-6A exhibited the opposite pattern compared with those of the GA biosynthesis and response genes in CS and NIL31. The results showed that the GA content of NIL31 was significantly higher than that of CS. Thus, TaGW2-6A had a negative relationship on GA synthesis and response genes. Moreover, after GA treatment, CS and NIL31 exhibited the opposite phenotypes and GA contents. These results demonstrate that allelic variation in TaGW2-6A increases the seed size via the GA hormone pathway. Transcriptional analysis and cytological analysis showed that TaGW2-6A allelic variants regulated GA synthesis via GA 3-oxidases , thereby leading to the higher expression of GASA4 to control endosperm cell elongation and division during grain filling. Finally, germination experiments were performed to elucidate the relationships between TaGW2-6A and GA synthesis and response genes in wheat with full fertility. These results provide new insights into the effects of the ubiquitination system mediated by TaGW2-6A on the GA hormone signaling pathway, thereby improving our understanding of the role of TaGW2-6A in seed development in wheat. [ABSTRACT FROM AUTHOR]

Heat stress is a limiting factor in wheat production along with global warming. Development of heat-tolerant wheat varieties and generation of suitable pre-breeding materials are the major goals in current wheat breeding programs. Our understanding on the genetic basis of thermotolerance remains sparse. In this study, we genotyped a collection of 211 core spring wheat accessions and conducted field trials to evaluate the grain-related traits under heat stress and non-stress conditions in two different locations for three consecutive years. Based on SNP datasets and grain-related traits, we performed genome-wide association study (GWAS) to detect stable loci related to thermotolerance. Thirty-three quantitative trait loci (QTL) were identified, nine of them are the same loci as previous studies, and 24 are potentially novel loci. Functional candidate genes at these QTL are predicted and proved to be relevant to heat stress and grain-related traits such as TaELF3-A1 (1A) for earliness per se (Eps), TaHSFA1-B1 (5B) influencing heat tolerance and TaVIN2-A1 (6A) for grain size. Functional markers of TaELF3-A1 were detected and converted to KASP markers, with their function and genetic diversity being analyzed in the natural populations. In addition, our results unveiled favor alleles controlling agronomic traits and/or heat stress tolerance. In summary, we provide insights into heritable correlation between yield and heat stress tolerance, which will accelerate the development of new cultivars with high and stable yield of wheat in the future. • Grain-related traits of spring wheat were subjected to GWAS under heat stress and yield potential conditions. • Thirty-three high-confidence stable QTL were identified. • Three genes were functionally annotated, possibly involve in grain size and heat tolerance. • Haplotype analysis revealed that TaELF3-A1 enhanced the adaptability of accessions to heat stress. [ABSTRACT FROM AUTHOR]

Glucuronokinase (GlcAK) converts glucuronic acid into glucuronic acid-1-phosphate, which is then converted into UDP-glucuronic acid (UDP-GlcA) via myo- inositol oxygenase (MIOX) pathway. UDP-GlcA acts as a precursor in the synthesis of nucleotide-sugar moieties forming cell wall biomass. GlcAK being present at the bifurcation point between UDP-GlcA and ascorbic acid (AsA) biosyntheses, makes it necessary to study its role in plants. In this study, the three homoeologs of GlcAK gene from hexaploid wheat were overexpressed in Arabidopsis thaliana. The GlcAK overexpressing transgenic lines showed decreased contents of AsA and phytic acid (PA) as compared to control plants. Root length and seed germination analyses under abiotic stress (drought and abscisic acid) conditions revealed enhanced root length in transgenic lines as compared to control plants. These results indicate that the MIOX pathway might be contributing towards AsA biosynthesis as evident by the decreased AsA content in the GlcAK overexpressing transgenic Arabidopsis thaliana plants. Findings of the present study will enhance the understanding of the involvement of GlcAK gene in MIOX pathway and subsequent physiological effects in plants. • Wheat GlcAK revealed differential expression pattern under heat, drought and salt stress conditions. • Overexpression of wheat GlcAK leads to decrease in phytic acid and ascorbic acid content in Arabidopsis thaliana. • Transgenic lines showed enhanced root length against drought and abscisic acid stress conditions. [ABSTRACT FROM AUTHOR]

Histone methylation is actively involved in plant flowering time and is regulated by a myriad of genetic pathways that integrate endogenous and exogenous signals. We identified an F-box gene from wheat (Triticum aestivum L.) and named it TaF-box3. Transcript expression analysis showed that TaF-box3 expression was gradually induced during the floret development and anthesis stages (WS2.5–10). Furthermore, ubiquitination assays have shown that TaF-box3 is a key component of the SCF ubiquitin ligase complex. TaF-box3 overexpression in Arabidopsis resulted in an early flowering phenotype and different cell sizes in leaves compared to the WT. Furthermore, the transcript level of a flowering time-related gene was significantly reduced in TaF-box3 overexpressing plants, which was linked with lower histone H3 Lys4 trimethylation (H3K4me3) and H3 Lys36 trimethylation (H3K36me3). Overexpression of TaF-box3 in Arabidopsis was shown to be involved in the regulation of flowering time by demethylating FLC chromatin, according to ChIP experiments. Protein analysis confirmed that TaMETS interacts with TaF-box3 and is ubiquitinated and degraded in a TaF-box3-dependnent manner. Based on these findings, we propose that TaF-box3 has a positive role in flowering time, which leads to a better understanding of TaF-box3 physiological mechanism in Arabidopsis. • TaF-box3 related to floret developmental stages in wheat, was cloned. • Overexpression of TaF-box3 in Arabidopsis promotes early flowering and leaf size. • TaF-box3 causes demethylation of histone H3 at FLC locus. • TaF-box3 interacts with TaMETS1, thereby mediating the 26 S proteasome pathway. [ABSTRACT FROM AUTHOR]

Few available leaf color mutants in crops have greatly limited the understanding of photosynthesis mechanisms, leading to few accomplishments in crop yield improvement via enhanced photosynthetic efficiency. Here, a noticeable albino mutant, CN19M06, was identified. A comparison between CN19M06 and the wild type CN19 at different temperatures showed that the albino mutant was temperature-sensitive and produced leaves with a decreased chlorophyll content at temperatures below 10 °C. Genetic analysis suggested that the albinism was controlled by one recessive nuclear gene named TSCA1 , which was putatively assigned to the region of 718.1–729.8 Mb on chromosome 2AL using bulked-segregant analysis and double-digest restriction site-associated DNA. Finally, molecular linkage analysis physically anchored TSCA1 to a narrowed region of 718.8–725.3 Mb with a 6.5 Mb length on 2AL flanked by InDel 18 and InDel 25 with 0.7 cM genetic interval. Among the 111 annotated functional genes in the corresponding chromosomal region, only TraesCS2A01G487900 of the PAP fibrillin family was both related to chlorophyll metabolism and temperature sensitivity; therefore, it was considered the putative candidate gene of TSCA1. Overall, CN19M06 has great potential for exploring the molecular mechanism of photosynthesis and monitoring temperature changes in wheat production. • A temperature sensitivity of wheat albino mutant was identified in the field and growth chamber. • Genetic analysis showed that inheritance of albino phenotype was controlled by one recessive nuclear locus TSCA1. • The TSCA1 gene was mapped in a narrow region of wheat chromosome 2AL by reduced-representation sequencing. [ABSTRACT FROM AUTHOR]

In the context of a continuously increasing human population that needs to be fed, with environmental protection in mind, nitrogen use efficiency (NUE) improvement is becoming very important. To understand the natural variation of traits linked to nitrogen uptake efficiency (UPE), one component of NUE, the multiparent advanced generation intercross (MAGIC) winter wheat population WM-800 was phenotyped under two contrasting nitrogen (N) levels in a high-throughput phenotyping facility for six weeks. Three biomass-related, three root-related, and two reflectance-related traits were measured weekly under each treatment. Subsequently, the population was genetically analysed using a total of 13,060 polymorphic haplotypes and singular SNPs for a genome-wide association study (GWAS). In total, we detected 543 quantitative trait loci (QTL) across all time points and traits, which were pooled into 42 stable QTL (sQTL; present in at least three of the six weeks). Besides Rht-B1 and Rht-D1 , candidate genes playing a role in gibberellic acid-regulated growth and nitrate transporter genes from the NPF gene family, like NRT 1.1 , were linked to sQTL. Two novel sQTL on chromosomes 5 A and 6D showed pleiotropic effects on several traits. The high number of N-specific sQTL indicates that selection for UPE is useful specifically under N-limited conditions. • Breeding for nitrogen uptake efficiency (UPE) is essential. • Wheat varieties differ significantly in their ability to cope with N deficiency. • N-dependent QTL suggest that selection for UPE under low N is preferable. • In total, 543 QTL were detected by GWAS, pooled into 42 time stable QTL. • Multiple detected QTL showed pleiotropic effects. [ABSTRACT FROM AUTHOR]

The wheat pericarp is the main component of the caryopsis at the early development stage and ultimately converts into a tissue that covers the mature caryopsis. A large number of starch granules are accumulated in the pericarp, but the production of and the role of starch granules in caryopsis development remain- elusive. In the present study, the relationship between accumulated starch granules and starch metabolism-related genes in wheat pericarp was investigated using paraffin section observations, expression analysis, and mutant analysis. Starch synthesis is initiated before anthesis and is dependent on a sucrose uptake and conversion system similar to that in the endosperm. TaPTST2 is required to initiate the production of pericarp starch granules. Pericarp starch granules gradually disappeared at the filling stage with high expression levels of genes encoding β-amylase, sucrose-phosphate synthase, and sucrose-phosphate phosphatase. As a maternal tissue adjacent to the endosperm and embryo, the pericarp plays a temporary reservoir for excess nutrients delivered into the caryopsis during the early development stage and exported at the filling stage. The pericarp contributes to the development of the endosperm and embryo as well as the accumulation of endosperm starch. The metabolism of pericarp starch may affect the weight of the wheat caryopsis. • Pericarp cell possesses a complete starch metabolism system. • TaPTST2 promotes the initiation of pericarp starch granule. • Pericarp acts as a temporary reservoir for nutrients delivered into the young caryopsis. [ABSTRACT FROM AUTHOR]

Mitogen-activated protein kinases (MAPKs) play important roles in the stress response of plants. However, the function of MPK proteins in freeze-resistance in wheat remains unclear. Dongnongdongmai No.1 (Dn1) is a winter wheat variety with a strong freezing resistance at extremely low temperature. In this study, we demonstrated that TaMPK6 is induced by JA signaling and is involved in the modulation of Dn1 freeze resistance. Overexpression of TaMPK6 in Arabidopsis increased the survival rate of plant at −10 ℃. The scavenging ability of reactive oxygen species (ROS) and the expression of cold-responsive genes CBFs and CORs were significantly enhanced in TaMPK6 -overexpressed Arabidopsis, suggesting a role of TaMPK6 in activating the ICE-CBF-COR module and antioxidant enzyme system to resist freezing stress. Furthermore, TaMPK6 is localized in the nucleus and TaMPK6 interacts with TaICE41, TaCBF14, and TaMYC2 proteins, the key components in JA signaling and the ICE-CBF-COR pathway. These results suggest that JA-induced TaMPK6 may regulate freezing-resistance in wheat by interacting with the TaICE41, TaCBF14, and TaMYC2 proteins, which in turn enhances the ICE-CBF-COR pathway. Our study revealed the molecular mechanism of TaMPK6 involvement in the cold resistance pathway in winter wheat under cold stress, which provides a basis for enriching the theory of wheat cold resistance. • TaMPK6 is induced by JA signaling. • TaMPK6 is involved in the modulation of winter wheat Dn1 freeze resistance. • Overexpression of TaMPK6 gene enhances freeze resistance in Arabidopsis plants. • TaMPK6 is localized in the nucleus. • TaMPK6 interacts with TaICE1, TaCBF14, and TaMYC2 proteins. [ABSTRACT FROM AUTHOR]

Accelerated soil-nitrifier activity and rapid nitrification are the cause of declining nitrogen-use efficiency (NUE) and enhanced nitrous oxide (N 2 O) emissions from farming. Biological nitrification inhibition (BNI) is the ability of certain plant roots to suppress soil-nitrifier activity, through production and release of nitrification inhibitors. The power of phytochemicals with BNI-function needs to be harnessed to control soil-nitrifier activity and improve nitrogen-cycling in agricultural systems. Transformative biological technologies designed for genetic mitigation are needed, so that BNI-enabled crop-livestock and cropping systems can rein in soil-nitrifier activity, to help reduce greenhouse gas (GHG) emissions and globally make farming nitrogen efficient and less harmful to environment. This will reinforce the adaptation or mitigation impact of other climate-smart agriculture technologies. [ABSTRACT FROM AUTHOR]

ROP is a subfamily of small GTP-binding proteins that uniquely exist in plants. It acts as versatile molecular switches that regulate various developmental processes. Some ROP proteins are also reported to affect defense responses, although their exact mechanism is not fully understood. Herein, ROP members in wheat were mined; the functions of three wheat ROP proteins were studied. RT-PCR results showed that the expression of TaRac1 was rapidly and strongly induced after leaf rust infection. TaRac1 interacted with TaCCR in yeast-hybridization assay. The overexpression of TaRac1 in tobacco promoted CCR and CAD gene expression, increased the total lignin content and sinapyl lignin proportion, and then enhanced resistance to tobacco black shank and bacterial wilt diseases. In contrast, TaRac3 and TaRac4 did not show to interact with TaCCR. Furthermore, the overexpression of TaRac3 and TaRac4 did not increase lignin gene expression and lignin accumulation either. Unlike TaRac1, the overexpression of TaRac3 increased susceptibility to both black shank and bacterial wilt pathogens, while overexpression of TaRac4 showed no effect on disease resistance but promoted the root growth in tobacco seedling. These data collectively suggest that TaRac1 in Group II is mainly involved in regulating lignin metabolism which, in turn, responsible for the observed roles in pathogen resistance. TaRac3 and TaRac4 have the minor roles in defense response but may act on regulation in plant developmental processes. These results shed light on the complexity and diverse function of ROP in plant defense pathway. [ABSTRACT FROM AUTHOR]

Abiotic stresses, such as heat and drought, are major environmental factors restricting crop productivity and quality worldwide. A plastid outer envelope protein gene, TaOEP16-2 , was identified from our previous transcriptome analysis [1,2] . In this study, the isolation and functional characterization of the TaOEP16-2 gene was reported. Three homoeologous sequences of TaOEP16-2 were isolated from hexaploid wheat, which were localized on the chromosomes 5A, 5B and 5D, respectively. These three homoeologues exhibited different expression patterns under heat stress conditions, TaOEP16-2 - 5B was the dominant one, and TaOEP16-2 - 5B was selected for further analysis. Compared with wild type (WT) plants, transgenic Arabidopsis plants overexpressing the TaOEP16-2 - 5B gene exhibited enhanced tolerance to heat stress, which was supported by improved survival rate, strengthened cell membrane stability and increased sucrose content. It was also found that TaOEP16-2 was induced by drought stress and involved in drought stress tolerance. TaOEP16-2-5B has the same function in ABA-controlled seed germination as AtOEP16-2 . Our results suggest that TaOEP16-2-5B plays an important role in heat and drought stress tolerance, and could be utilized in transgenic breeding of wheat and other crop plants. [ABSTRACT FROM AUTHOR]

Plant MAP kinase phosphatases (MKPs) are major regulators of MAPK signaling pathways and play crucial roles in controlling growth, development and stress responses. The presence of several functional domains in plant MKPs such as a dual specificity phosphatase catalytic domain, gelsolin, calmodulin-binding and serine-rich domains, suggests that MKPs can interact with distinct cellular partners, others than MAPKs. In this report, we identified a canonical mode I 14-3-3-binding motif ( 574 KLPSLP 579 ) located at the carboxy-terminal region of the wheat MKP, TMKP1. We found that this motif is well-conserved among other MKPs from monocots including Hordeum vulgare , Brachypodium distachyon and Aegilops taushii . Using co-immunoprecipitation assays, we provide evidence for interaction between TMKP1 and 14-3-3 proteins in wheat. Moreover, the phosphatase activity of TMKP1 is increased in a phospho-dependent manner by either Arabidopsis or yeast 14-3-3 isoforms. TMKP1 activation by 14-3-3 proteins is enhanced by Mn 2+ , whereas in the presence of Ca 2+ ions, TMKP1 activation was limited to Arabidopsis 14-3-3φ (phi), an isoform harboring an EF-hand motif. Such findings strongly suggest that 14-3-3 proteins, in conjunction with specific divalent cations, may stimulate TMKP1 activity and point-out that 14-3-3 proteins bind and regulate the activity of a MKP in eukaryotes. [ABSTRACT FROM AUTHOR]

There will be longer and more intense periods of heat and drought stress in the future for terrestrial ecosystems. Although the responses of wheat plants to heat and drought stress alone have been extensively investigated, little is known about the extent to which their recovery can be assured after stress relief. In this study, a winter wheat pot experiment was conducted to investigate the changes in photosynthetic performance, antioxidant activity, osmoregulation, and membrane lipid peroxidation under heat stress (36 °C), drought (45–55% of soil water holding capacity), and combined stress conditions. The results showed that heat and drought stress significantly reduced the photosynthetic rate and the contents of chlorophyll and carotenoid. Superoxide dismutase (SOD), peroxidase (POD), catalase (CAT), and glutathione reductase (GR) activities were greatly activated by heat and drought stress to scavenge overproduced superoxide anion (O 2 -). Plants exhibited positive osmoregulation through the synthesis of soluble protein (SP), soluble sugar (SS), and proline (Pro) to improve membrane stability. Within a range of stress, combined heat and drought stress exhibited significant interactive effects in the above mentioned indicators. After stress relief, the majority of physiological processes were reversible, as indicated by the effective recovery of pigment contents, photosynthetic rate, antioxidant enzyme activities, osmoregulatory substance contents, and O 2 - production. Antioxidant enzyme activities tended to increase after recovering from 12 days of combined stress, whereas they were still not effective in mitigating oxidative damage. High levels of O 2 - and malondialdehyde (MDA) and a low relative growth rate during the recovery confirmed the irreversible damage caused by combined heat and drought stress. ROC (receiver operating characteristic) analysis indicated that GR and SS could accurately detect individual heat and drought stress that wheat plants were suffering or had suffered (AUC = 0.812–0.965), while POD and Pro had greater potential for diagnosing combined heat and drought stress (AUC = 0.871–0.958). Physiological indicators of stress tolerance were closely related to the photosynthetic rate during the stress, particularly Pro and GR. Collectively, the physiological processes of plants are reversible within a certain range of stress. POD, GR, Pro, and SS play vital roles in identifying and resisting heat, drought, and combined stress, and the recovery of these indicators contributed to improving photosynthesis and thereby increasing wheat growth. Our research contributes to the understanding of the underlying physiological mechanisms of plants in response to combined heat and drought stress and after stress relief. [Display omitted] • The responses of physiological activity to stress and recovery periods were studied. • Physiological processes of plants are reversible within a range of stresses. • POD, GR, Pro, and SS play vital roles in identifying heat and drought stress. • Physiological traits of stress resistance are closely related to photosynthesis. [ABSTRACT FROM AUTHOR]

Wheat SOS1 (TaSOS1) activity could be relieved upon deletion of the C-terminal 168 residues (the auto-inhibitory domain). This truncated form of wheat SOS1 (TaSOS1-974) was shown to increase compensation (compared to wild-type TaSOS1) for the salt sensitivity of a yeast mutant strain, AXT3K, via increased Na + transportation out of cells during salinity stress. Expression of the plasma membrane proteins TaSOS1-974 or TaSOS1 improved the growth of transgenic tobacco plants compared with wild-type plants under normal conditions. However, plants expressing TaSOS1-974 grew better than TaSOS1 -transformed plants. Upon salinity stress, Na + efflux and K + influx rates in the roots of transgenic plants expressing TaSOS1-974 or TaSOS1 were greater than those of wild-type plants. Furthermore, compared to TaSOS1 -transgenic plants, TaSOS1-974 -expressing roots showed faster Na + efflux and K + influx, resulting in less Na + and more K + accumulation in TaSOS1-974 -transgenic plants compared to TaSOS1 -transgenic and wild-type plants. TaSOS1-974 -expressing plants had the lowest MDA content and electrolyte leakage among all tested plants, indicating that TaSOS1-974 might protect the plasma membrane against oxidative damage generated by salt stress. Overall, TaSOS1-974 conferred higher salt tolerance in transgenic plants compared to TaSOS1. Consistent with this result, transgenic plants expressing TaSOS1-974 showed a better growth performance than TaSOS1 -expressing and wild-type plants under saline conditions. [ABSTRACT FROM AUTHOR]

The effect of one-day cold-shock on the transcriptome and phytohormones (auxin, cytokinins, abscisic, jasmonic and salicylic acids) was characterised in freezing-sensitive (Chinese Spring), highly freezing-tolerant (Cheyenne) and moderately freezing-tolerant (Chinese Spring substituted with Cheyenne’s 5A chromosome) wheat genotypes. Altogether, 636 differentially expressed genes responding to cold‐shock were identified. Defence genes encoding LEA proteins, dehydrins, chaperons and other temperature-stress responsive proteins were up‐regulated in a genotype‐independent manner. Abscisic acid was up-regulated by cold accompanied by adherent expression of its metabolic genes. Data revealed the involvement of particular routes within ABA-dependent signalling in response to cold‐shock in the examined genotypes. Cold‐shock affected gene expression along carbohydrate metabolic pathways. In photosynthesis, cold‐shock changed the expression of a number of genes in the same way as it was previously reported for ABA. Overrepresentation analysis of the differentially expressed genes supported the ABA‐signalling and carbohydrate metabolism results, and revealed some pronounced biological process GO categories associated with the cold‐shock response of the genotypes. Protein network analysis indicated differences between the genotypes in the information flow along their signal perception and transduction, suggesting different biochemical and cellular strategies in their reaction to cold‐shock. [ABSTRACT FROM AUTHOR]

Q-type C 2 H 2 zinc finger proteins (ZFPs) are plant-specific DNA-binding proteins containing a conserved QALGGH motif. This study investigated the function of abiotic stress-inducible and predominantly root-expressed Triticum aestivum ZFPs (TaZFP22, TaZFP34 and TaZFP46) with a focus on TaZFP34. Expression of TaZFP34 in roots was upregulated by high salinity, dehydration, oxidative and cold stresses. Overexpression of TaZFP34 in wheat roots resulted in an increased root-to-shoot ratio, a phenomenon observed during plant adaptation to drying soil. Expression of a number of genes which are potentially involved in modulating root growth was significantly altered in the roots of TaZFP34 overexpressing lines. In particular, the transcript levels of TaRR12B , TaRR12D and TaSHY2 that are homologues of known negative regulators of root growth were significantly reduced. Expression of shoot growth-related genes, such as GA3-ox and expansins, was downregulated in the transgenic shoots. TaZFP34 bound to (C/G)AGT(G/A)-like elements in the promoters of TaZFP34 down-regulated TaRR12D and TaSHY2 and transrepressed the reporter gene expression driven by TaRR12D and TaSHY2 promoters. Expression of the above reporter genes was also repressed by TaZFP46 and TaZFP22. These data suggest that TaZFP34 is a transcriptional repressor and is involved in modulating the root-to-shoot ratio. [ABSTRACT FROM AUTHOR]

MiR396s play important roles in regulating plant growth and stress response, and great potential for crop yield promotion was anticipated. For more comprehensive and precise understanding of miR396s in Poaceae, we analyzed the phylogenetic linkage, gene expression, and chromosomal distribution of miR396s in this study. Although the mature miR396s' sequences were mostly conserved, differential expression patterns and chromosomal distribution were found among Poaceae species including the major cereal crops rice, wheat, and maize. Consistently, in comparison with rice, wheat and maize plants transformed with the target mimicry construct of miR396 (MIM396) exhibited differential effects on grain size and disease resistance. While the TaMIM396 plants showed increased grain size, panicle length and sensitivity to B. graminis , the ZmMIM396 plants didn't show obvious changes in grain size and disease resistance. In Addition, several GROWTH-REGULATING FACTOR (GRF) genes in wheat and maize were repressed by miR396s, which could be reversed by MIM396, confirming the conserved regulatory roles of miR396 on GRFs. While providing new solution to enhance grain yield in wheat and revealing potential regulatory variations of miR396s in controlling grain size and disease resistance in different crops, this study gives clues to further explore miR396s' functions in other Poaceae species. • Poaceae miR396s are divided into three groups and display both conserved and diversified evolutionary characterizations. • Inhibition of miR396s results in increased grain size, panicle length and pathogen sensitivity in wheat but not in maize. • Different GRFs were upregulated in the MIM396 transgenic plants compared to their corresponding wild type in wheat and maize. [ABSTRACT FROM AUTHOR]

Starch is a major component of cereal grains such as wheat. Physicochemical and functional properties of starch affect end-use food quality and nutrients. To improve cultivars that preserve superior starch quality, the genetic foundation of the wheat starch and amylose-lipid complex (ALc, Resistant starch type 5) gelatinization are needed. This genome-wide association (GWA) mapping used 192 wheat genotypes (previously reported) to generate SNPs using an enhanced version of sequencing termed ddRAD on the Illumina Hi-seq X platform and 3696 high-quality influential SNPs were filtered out. The heterozygosity and Fst ranges in five subpopulations were 0.31–0.40 and 0.18–0.30 respectively. Nucleotide diversity and PIC ranged from 0.21 (6A) to 0.32 (2A) and 0.29 (6A) to 0.39 (4D) respectively. The Shannon waiver index was 1.7 and the whole-genome LD decay was 22 Mb at r 2 = 0.38. Following FDR, 23 and 8 SNPs showed association with starch properties in the year 2017 and 2018, respectively while 93 and 20 SNPs were associated with ALc gelatinization in the year 2017 and 2018 respectively. The identified potential new genes (GSK3-alpha, RING-type domain-containing protein , Tetratricopeptide repeat , Hexosyltransferase, GLP, SNF1, and WRKY transcription factor) within LD range (∼16 Kb to ∼15 Mb), BLUP value, and cis and trans-position of SNPs network provide valuable information for the future wheat breeding strategy for the improvement of the starch quality trait. [Display omitted] • 192 diverse Indian wheat were used for GWAS analysis (releases year: 1910–2006). • Thermal behavior (Gelatinization) of Starch and amylose-lipid complex (RS-5) were characterized. • Advanced ddRAD Illumina High-seq genotyping was used for SNPs identification. • Discovered 144 high confidence novel SNPs controlling gelatinization in wheat. • seven new genes were functionally annotated, possibly involve in starch gelatinization. • It can be used to accelerate wheat starch and RS-5 quality by functional genomics/genome editing. [ABSTRACT FROM AUTHOR]

Male reproductive development in higher plants experienced a series of complex biological processes, which can be regulated by Gibberellins (GA). The transcriptional factor GAMYB is a crucial component of GA signaling in anther development. However, the mechanism of GAMYB in wheat male reproduction is less understood. Here, we found that the thermo-sensitive genic male sterilitywheat line YanZhan 4110S displayed delayed tapetum programmed cell death and pollen abortive under the hot temperature stress. Combined with RNA-Sequencing data analysis, TaGAMYB associated with fertility conversion was isolated, which was located in the nucleus and highly expressed in fertility anthers. The silencing of TaGAMYB in wheat displayed fertility decline, defects in tapetum, pollen and exine formation, where the abortion characteristics were the same as YanZhan 4110S. In addition, either hot temperature or GA3 treatment in YanZhan 4110S caused the downregulation of TaGAMYB at binucleate stage and trinucleate stage, as well as fertility decrease. Further, the transcription factor TaWRKY2 significantly changed under GA3-treatment and directly interacted with the TaGAMYB promoter by W-box cis -element. Therefore, we suggested that TaGAMYB may be essential for anther development and male fertility, and GA3 activates TaGAMYB by TaWRKY2 to regulate fertility in wheat. • TaGAMYB was located in the nucleus and highly expressed in anthers. • Silencing of TaGAMYB led to fertility decline and defects in the tapetum and pollen development. • GA3 can activate TaGAMYB by TaWRKY2 to regulate fertility in wheat. [ABSTRACT FROM AUTHOR]

Abscisic acid (ABA) regulates seed dormancy and therefore preharvest sprouting (PHS) in wheat. This study investigated the contribution of transcriptional regulation of ABA metabolism and signaling genes to genetic variation in dormancy of wheat seeds. Our results showed that genetic variation in seed dormancy is highly correlated with ABA content (r > 0.86), which, in turn, was closely associated with the expression levels of ABA biosynthesis genes, TaNCED1 (r = 0.78) and TaNCED2 (r = 0.67). A relatively lower correlation was observed between ABA content and the expression levels of ABA catabolism genes, TaCYP707A1 (r = 0.51) and TaCYP707A2 (r = 0.57). The expression level of TaABI5 exhibited strong associations with the levels of ABA (r = 0.8) and seed dormancy (r > 0.9), indicating the importance of seed ABA sensitivity in mediating genetic variation in dormancy. Furthermore, high positive correlations were prevalent between the expression patterns of TaABI5 and TaNCED1 (r = 0.91) or TaNCED2 (r = 0.82). Overall, ou r results implicated the significance of TaNCED s and TaABI5 in regulating genetic variation in ABA level and sensitivity and thereby seed dormancy, highlighting the potential use of these genes to develop molecular markers for incorporating PHS resistance in wheat. • ABA underlies genetic variation of seed dormancy in wheat. • Seed ABA levels are regulated mainly by expression of TaNCED1 and TaNCED2 genes. • Transcriptional regulation TaABI5 mediates seed ABA sensitivity. • Seed dormancy is not associated with seed coat pigmentation. [ABSTRACT FROM AUTHOR]

Low temperature is one of the main abiotic stresses that inhibit wheat growth and development. To understand the physiological mechanism of salt priming induced low temperature tolerance and its transgenerational effects, the chlorophyl b-deficient mutant (ANK) and its wild type (WT) wheat were subjected to low temperature stress after parental salt priming. Salt priming significantly decreased the levels of superoxide anions, hydrogen peroxide and malondialdehyde in both parental and offspring plants under low temperature. The catalase activity in parental wheat and activities of dehydroascorbate reductase and glutathione reductase in the offspring were significantly increased by salt priming under low temperature. Meanwhile, salt priming contributed to mantaining the integrity of chloroplast structure and relatively higher net photosynthetic rate (Pn) in both generations under low temperature. Salt priming also improved the carbohydrate metabolism enzyme activities of parental and offspring plants, such as phosphoglucomutase, fructokinase and sucrose synthase. In addition, ANK plants had significantly higher carbohydrate metabolism enzyme activities than WT plants. The differential expressed proteins (DEP) in seeds of two genotypes under salt priming were mainly related to homeostasis, electron transfer activity, photosynthesis and carbohydrate metabolism. Correlation network analysis showed that the expression of DEP under salt priming was significantly correlated to sucrose concentration and cytoplasmic peroxidase (POX) activity in WT, while that was correlated to various carbohydrate metabolism enzyme activities in ANK plants. These results indicated that the parental salt priming induced modulations of seed proteome regulated the ROS metabolism, photosynthetic carbon assimilation and carbohydrate metabolism, hence enhancing the low temperature tolerance in offspring wheat. • Salt priming decreased the levels of reactive oxygen species in seedlings under low temperature. • Salt priming improved the carbohydrate metabolism enzyme activities in seedlings. • The parental salt priming enhances low temperature tolerance in offspring wheat via modulating the seed proteome. • The differential expressed proteins were related to homeostasis, electron transfer activity, photosynthesis and carbohydrate metabolism. [ABSTRACT FROM AUTHOR]

A plethora of microtubule-associated proteins (MAPs) modulate the dynamics of microtubules (MTs) to ensure the proper elaboration of developmental programs in plants. Among the plant-specific MAPs are the IQ67 domain (IQD) family proteins. Despite the great progress in elucidating IQD protein functions, the majority of IQD proteins, especially IQDs in crop species, remain to be functionally explored. In this study, we identified 78 putative IQD family genes in the genome of hexaploid wheat (Triticum aestivum). Phylogenetic analysis of wheat and Arabidopsis IQDs supports the previous notion that the expansion of the IQD family coincides with plant terrestrialization. Further characterization of one TaIQD , TaIQD3D-6 , revealed that TaIQD3D-6 directly binds to MTs and free tubulins in vitro and is associated with cortical MTs in interphase cells in vivo. Overexpressing TaIQD3D-6 in Arabidopsis leads to a spectrum of phenotypes that are indicative of perturbed MT homeostasis, including spiral growth, hypersensitivity to MT-destabilizing drugs, defects in cell morphogenesis, and altered organization of cMT arrays. Finally, we determined that TaIQD3D-6-GFP localizes to the expanding cell plate during cytokinesis and the overexpression of TaIQD3D-6 interferes with asymmetric cell division in the stomatal lineage in Arabidopsis. In summary, our findings establish that TaIQD3D-6 is a MAP that regulates plant cell and organ morphogenesis and provide new insights into the functions of crop IQD proteins. • IQD gene family in hexaploid wheat (Triticum aestivum) contains at least 78 members. • One wheat IQD protein, TaIQD3D-6, is associated with microtubules in vitro and in vivo. • The overexpression of TaIQD3D-6 in Arabidopsis perturbs MT homeostasis and leads to left-handed helical growth. • The overexpression of TaIQD3D-6 interferes with asymmetric cell divisions in the stomatal lineage in Arabidopsis. [ABSTRACT FROM AUTHOR]

Improving yield potential is a major goal of wheat breeding that depends on identifying key genetic loci. In this study, two residual heterozygous line RHL351- and RHL78-derived populations were employed for genetic linkage map construction and QTL detection. Two genetic populations indicated a robust grain-size QTL between Marker6 and Marker10. It covered a 95.54–99.38 Mb physical interval and was named Qpleio.nwafu.3D , containing the candidate gene Tasg (TraesCS3D02G137200). Intriguingly, RNA-seq analysis and sequencing revealed two different allelic variants in Tasg , named Tasg-D1 (G>A) and Tasg-D2 (C>G), respectively. Although the relationship between Tasg-D1 and grain size had been demonstrated previously, here we provided the first genetic evidence that C/G allelic variation in Tasg-D2 was associated with grain shape and size through a newly developed dCAPS marker. In addition, transcriptome comparison indicated that Tasg-D1/2 might primarily contribute to significant expression differences in brassinolide (BR) metabolism-related genes rather than those related to BR responses in developing grains and spikes. Our study provided new evidence and a breeder-friendly dCAPS marker for improving grain size through the selection of Tasg , as well as a basis to understand Tasg function in the future. • Qpleio.nwafu.3D was a robust and stable QTL mainly responsible for grain morphology. • The C/G allelic variation in Tasg-D2 was associated with grain-size, especially for grain shape. • Tasg-D1/2 was implicated into regulate BR metabolism rather than BR signaling response. [ABSTRACT FROM AUTHOR]

The thermo-sensitive cytoplasmic male-sterility line with Aegilops kotschyi cytoplasm (K-TCMS) is completely male sterile under low temperature (< 18 ℃) during Zadoks growth stages 45–52, whereas its fertility can be restored under hot temperature (≥ 20 ℃). The K-TCMS line may facilitate hybrid breeding and hybrid wheat production. Therefore, to elucidate the molecular mechanisms of its male sterility/fertility conversion, we conducted the association analysis of proteins and transcript expression to screen fertility related genes using RNA-seq, iTRAQ, and PRM-based assay. A gene encoding expansin protein in wheat, TaEXPB5 , was isolated in K-TCMS line KTM3315A, which upregulated expression in the fertility anthers. Subcellular localization analysis suggested that TaEXPB5 protein localized to nucleus and cell wall. The silencing of TaEXPB5 displayed pollen abortion and the declination of fertility. Further, cytological investigation indicated that the silencing of TaEXPB5 induced the early degradation of tapetum and abnormal development of pollen wall. These results implied that TaEXPB5 may be essential for anther or pollen development and male fertility of KTM3315A. These findings provide a novel insight into molecular mechanism of fertility conversion for thermo-sensitive cytoplasmic male-sterility wheat, and contribute to the molecular breeding of hybrid wheat in the future. ● TaEXPB5, a gene encoding expansin protein in wheat, was upregulated expression in KTM3315A under fertility condition. ● Silencing of TaEXPB5 displayed pollen abortion and fertility decline. ● Inhibition of TaEXPB5 expression led to early programmed cell death in the tapetum. ● TaEXPB5 was involved in pollen development. [ABSTRACT FROM AUTHOR]

In the last decade the breeding technology referred to as ‘genomic selection’ (GS) has been implemented in a variety of species, with particular success in animal breeding. Recent research shows the potential of GS to reshape wheat breeding. Many authors have concluded that the estimated genetic gain per year applying GS is several times that of conventional breeding. GS is, however, a new technology for wheat breeding and many programs worldwide are still struggling to identify the best strategy for its implementation. This article provides practical guidelines on the key considerations when implementing GS. A review of the existing GS literature for a range of species is provided and used to prime breeder-oriented considerations on the practical applications of GS. Furthermore, this article discusses potential breeding schemes for GS, genotyping considerations, and methods for effective training population design. The components of selection intensity, progress toward inbreeding in half- or full-sibs recurrent schemes, and the generation of selection are also presented. [ABSTRACT FROM AUTHOR]