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SUMMARY: Strigolactones (SLs) are key regulators of shoot growth and responses to environmental stimuli. Numerous studies have indicated that nitrogen (N) limitation induces SL biosynthesis, suggesting that SLs may play a pivotal role in coordinating systemic responses to N availability, but this idea has not been clearly demonstrated. Here, we generated triple knockout mutants in the SL synthesis gene TaDWARF17 (TaD17) in bread wheat and investigated their phenotypic and transcriptional responses under N limitation, aiming to elucidate the role of SLs in the adaptation to N limitation. Tad17 mutants display typical SL mutant phenotypes, and fail to adapt their shoot growth appropriately to N. Despite exhibiting an increased tillering phenotype, Tad17 mutants continued to respond to N limitation by reducing tiller number, suggesting that SLs are not the sole regulators of tillering in response to N availability. RNA‐seq analysis of basal nodes revealed that the loss of D17 significantly altered the transcriptional response of N‐responsive genes, including changes in the expression profiles of key N response master regulators. Crucially, our findings suggest that SLs are required for the transcriptional downregulation of cytokinin (CK) synthesis and signalling in response to N limitation. Collectively, our results suggest that SLs are essential for the appropriate morphological and transcriptional adaptation to N limitation in wheat, and that the repressive effect of SLs on shoot growth is partly mediated by their repression of CK synthesis. Significance Statement: This research investigates the key role of strigolactones (SLs) in bread wheat tillering and adaptation to nitrogen limitation. Our findings using TaDWARF17 mutants reveal that SLs, while not the sole regulators of tillering under nitrogen limitation, crucially mediate the transcriptional downregulation of cytokinin synthesis and signalling, underscoring the importance of SLs in coordinating morphological and transcriptional responses to nitrogen availability in wheat. [ABSTRACT FROM AUTHOR]

The importance of monitoring and management of yellow rust, caused by Puccinia striiformis f. sp. tritici (Pst), has markedly increased in Russia over recent years because of intensifying negative effects of this destructive disease on wheat production. The Pst virulence survey was conducted in 2019–2021 in the European (North‐West, Volga, North Caucasus) and Asian (Siberia) parts of Russia, and the same 109 isolates have been genotyped here with 20 SSR markers. Fifty‐three multilocus genotypes (MLGs) were detected, seven of which occurred in two or more regions. The most frequent MLG was found in three European Pst collections (except Dagestan) and Siberia. SSR‐based variation within the regional collections was low because many MLGs were closely related. Nevertheless, each collection contained relatively different small genotype groups. No clear geographic structure of the Russian Pst population was observed because the five studied regional collections shared groups of similar genotypes. The most singular SSR genotype was identified in the North‐West collection, but genotypes of four invasive isolates belonging to the PstS2 temperature‐adapted aggressive strain did not considerably differ from others. The observed heterozygosity at all SSR loci exceeded the expected one in all regional Pst collections, which could imply asexual reproduction of the pathogen. Both the SSR and virulence analyses demonstrated similar patterns of relationships between the regional Pst collections, although the magnitude of differences between the collections was larger for virulence pathotypes. However, no significant association was established between the virulence pathotypes and SSR genotypes because multiple pathotypes had the same genotype and vice versa. [ABSTRACT FROM AUTHOR]

Summary: High‐throughput genotyping arrays have provided a cost‐effective, reliable and interoperable system for genotyping hexaploid wheat and its relatives. Existing, highly cited arrays including our 35K Wheat Breeder's array and the Illumina 90K array were designed based on a limited amount of varietal sequence diversity and with imperfect knowledge of SNP positions. Recent progress in wheat sequencing has given us access to a vast pool of SNP diversity, whilst technological improvements have allowed us to fit significantly more probes onto a 384‐well format Axiom array than previously possible. Here we describe a novel Axiom genotyping array, the 'Triticum aestivum Next Generation' array (TaNG), largely derived from whole genome skim sequencing of 204 elite wheat lines and 111 wheat landraces taken from the Watkins 'Core Collection'. We used a novel haplotype optimization approach to select SNPs with the highest combined varietal discrimination and a design iteration step to test and replace SNPs which failed to convert to reliable markers. The final design with 43 372 SNPs contains a combination of haplotype‐optimized novel SNPs and legacy cross‐platform markers. We show that this design has an improved distribution of SNPs compared to previous arrays and can be used to generate genetic maps with a significantly higher number of distinct bins than our previous array. We also demonstrate the improved performance of TaNGv1.1 for Genome‐wide association studies (GWAS) and its utility for Copy Number Variation (CNV) analysis. The array is commercially available with supporting marker annotations and initial genotyping results freely available. [ABSTRACT FROM AUTHOR]

The genus Clavibacter (family Microbacteriaceae) includes gram‐positive actinobacterial species infecting diverse agricultural crops such as alfalfa, maize, pepper, potato, tomato and wheat. Clavibacter tessellarius, causing bacterial mosaic of wheat, was reported for the first time in 1976 in Nebraska (United States). Since then, the pathogen was thought to be restricted to North America (Canada and the United States). While bacterial strains suspected to be C. tessellarius have occasionally been isolated from wheat samples outside North America, occurrence of the pathogen in the Old World has not yet been confirmed by the plant protection authorities. In spring 2020 (January to May) dozens of wheat fields with young plants showing leaf chlorosis and mild mosaic symptoms were surveyed and sampled in southern Iran. Among 192 surveyed wheat fields, gram‐positive, orange‐pigmented actinobacterial strains were isolated from samples of 53 fields. Altogether, 61 bacterial strains were identified as C. tessellarius based on the phenotypic features, PCR‐based molecular tests and pathogenicity assays. Multilocus sequence analysis of five housekeeping genes (atpD, dnaK, gyrB, ppk and rpoB) using 10 representative strains from diverse geographic areas in southern Iran confirmed the phylogenetic status of the strains within C. tessellarius. Our data indicate that the bacterial mosaic pathogen occurs in four southern Iranian provinces including Bushehr, Fars, Kerman and Khuzestan. Thus, occurrence of C. tessellarius outside North America is confirmed. Potential risks associated with the emergence of the pathogen in the Old World, and the need for strict quarantine measures in the area, is further discussed. [ABSTRACT FROM AUTHOR]

SUMMARY Plant fungal parasites manipulate host metabolism to support their own survival. Among the many central metabolic pathways altered during infection, the glyoxylate cycle is frequently upregulated in both fungi and their host plants. Here, we examined the response of the glyoxylate cycle in bread wheat (Triticum aestivum) to infection by the obligate biotrophic fungal pathogen Puccinia striiformis f. sp. tritici (Pst). Gene expression analysis revealed that wheat genes encoding the two unique enzymes of the glyoxylate cycle, isocitrate lyase (TaICL) and malate synthase, diverged in their expression between susceptible and resistant Pst interactions. Focusing on TaICL, we determined that the TaICL B homoeolog is specifically upregulated during early stages of a successful Pst infection. Furthermore, disruption of the B homoeolog alone was sufficient to significantly perturb Pst disease progression. Indeed, Pst infection of the TaICL‐B disruption mutant (TaICL‐BY400*) was inhibited early during initial penetration, with the TaICL‐BY400* line also accumulating high levels of malic acid, citric acid, and aconitic acid. Exogenous application of malic acid or aconitic acid also suppressed Pst infection, with trans‐aconitic acid treatment having the most pronounced effect by decreasing fungal biomass 15‐fold. Thus, enhanced TaICL‐B expression during Pst infection may lower accumulation of malic acid and aconitic acid to promote Pst proliferation. As exogenous application of aconitic acid and malic acid has previously been shown to inhibit other critical pests and pathogens, we propose TaICL as a potential target for disruption in resistance breeding that could have wide‐reaching protective benefits for wheat and beyond. [ABSTRACT FROM AUTHOR]

Efficient use of nitrogen (N) and phosphorus (P) is essential to reduce fertilizer costs and nutrient pollution and to lower the carbon footprint of agriculture. This requires a better understanding of N and P limitations on photosynthesis and biomass generation in one of the world's most important crops, wheat (Triticum aestivum). In a fully factorial nutrient experiment, we find that the limiting nutrient, N or P, at a Redfield‐like N:P ratio (21N:1P molar) produces the 'effective dose of fertilizer' for the generation of above‐ground biomass (seed or vegetative). Best‐fit structural equation models showed that neither N nor P applied in excess of 21N:1P led to any increase in biomass. The light (qP, NPQ) and dark (Amax, Vcmax, Jmax) reactions of photosynthesis are also constrained by the limiting nutrient at the 21N:1P Redfield‐like ratio. Beyond this ratio, 'excess N' or 'excess P' had no effects. The direct effects of the 'effective dose of fertilizer' on the accumulation of biomass were stronger than its indirect effects via photosynthesis, likely driven by the N and P costs in building nucleic acids needed for cell division, cell maintenance and RNA transcription. Modern composite fertilizers are typically provided at a much higher P content than a 21N:1P ratio, potentially resulting in a huge global wastage of P, a finite resource, with all the concomitant costs to the farmer, consumer and the environment. [ABSTRACT FROM AUTHOR]

SUMMARY Septoria nodorum blotch (SNB), caused by Parastagonospora nodorum, is a disease of durum and common wheat initiated by the recognition of pathogen‐produced necrotrophic effectors (NEs) by specific wheat genes. The wheat gene Snn1 was previously cloned, and it encodes a wall‐associated kinase that directly interacts with the NE SnTox1 leading to programmed cell death and ultimately the development of SNB. Here, sequence analysis of Snn1 from 114 accessions including diploid, tetraploid, and hexaploid wheat species revealed that some wheat lines possess two copies of Snn1 (designated Snn1‐B1 and Snn1‐B2) approximately 120 kb apart. Snn1‐B2 evolved relatively recently as a paralog of Snn1‐B1, and both genes have undergone diversifying selection. Three point mutations associated with the formation of the first SnTox1‐sensitive Snn1‐B1 allele from a primitive wild wheat were identified. Four subsequent and independent SNPs, three in Snn1‐B1 and one in Snn1‐B2, converted the sensitive alleles to insensitive forms. Protein modeling indicated these four mutations could abolish Snn1–SnTox1 compatibility either through destabilization of the Snn1 protein or direct disruption of the protein–protein interaction. A high‐throughput marker was developed for the absent allele of Snn1, and it was 100% accurate at predicting SnTox1‐insensitive lines in both durum and spring wheat. Results of this study increase our understanding of the evolution, diversity, and function of Snn1‐B1 and Snn1‐B2 genes and will be useful for marker‐assisted elimination of these genes for better host resistance. [ABSTRACT FROM AUTHOR]

Leaf senescence is an essential physiological process related to grain yield potential and nutritional quality. Green leaf duration (GLD) after anthesis directly reflects the leaf senescence process and exhibits large genotypic differences in common wheat; however, the underlying gene regulatory mechanism is still lacking. Here, we identified TaNAM‐A1 as the causal gene of the major loci qGLD‐6A for GLD during grain filling by map‐based cloning. Transgenic assays and TILLING mutant analyses demonstrated that TaNAM‐A1 played a critical role in regulating leaf senescence, and also affected spike length and grain size. Furthermore, the functional divergences among the three haplotypes of TaNAM‐A1 were systematically evaluated. Wheat varieties with TaNAM‐A1d (containing two mutations in the coding DNA sequence of TaNAM‐A1) exhibited a longer GLD and superior yield‐related traits compared to those with the wild type TaNAM‐A1a. All three haplotypes were functional in activating the expression of genes involved in macromolecule degradation and mineral nutrient remobilization, with TaNAM‐A1a showing the strongest activity and TaNAM‐A1d the weakest. TaNAM‐A1 also modulated the expression of the senescence‐related transcription factors TaNAC‐S‐7A and TaNAC016‐3A. TaNAC016‐3A enhanced the transcriptional activation ability of TaNAM‐A1a by protein–protein interaction, thereby promoting the senescence process. Our study offers new insights into the fine‐tuning of the leaf functional period and grain yield formation for wheat breeding under various geographical climatic conditions. [ABSTRACT FROM AUTHOR]

Wheat stripe mosaic virus (WhSMV) is the causal agent of soilborne wheat mosaic disease (SBWMD) in Brazil, which is a serious threat to wheat production in the southern part of the country. WhSMV also occurs in Paraguay and South Africa. The virus is soilborne, transmitted by Polymyxa graminis, and management strategies are mainly based on genetic resistance. Variation in the reaction (type and severity of symptoms) of cultivars has been reported depending on the region and/or year of cultivation, leading wheat breeding programmes to test their materials in several locations, which increases costs and is time‐consuming. One hypothesis is that this variation in symptoms is a result of the genetic variability of WhSMV. We assessed the genetic variability and population structure of WhSMV infecting wheat in Brazil and South Africa. In field experiments conducted in different locations in southern Brazil, a consistent reproducibility of symptoms was observed in a set of cultivars, and molecular analysis showed a low degree of genetic variability of the Brazilian viral population. The hypothesis that the variation in SBWMD symptoms in Brazil is due to genetic variability of the virus was thus rejected. Comparison of the Brazilian and South African WhSMV isolates indicated that they comprise distinct subpopulations and that the Brazilian subpopulation is more variable than the South African subpopulation. The lower genetic variability of the South African subpopulation suggests genetic stability or a recent emergence of WhSMV in that region. [ABSTRACT FROM AUTHOR]

Summary: Wheat needs different durations of vernalization, which accelerates flowering by exposure to cold temperature, to ensure reproductive development at the optimum time, as that is critical for adaptability and high yield. TaVRN1 is the central flowering regulator in the vernalization pathway and encodes a MADS‐box transcription factor (TF) that usually works by forming hetero‐ or homo‐dimers. We previously identified that TaVRN1 bound to an MADS‐box TF TaSOC1 whose orthologues are flowering activators in other plants. The specific function of TaSOC1 and the biological implication of its interaction with TaVRN1 remained unknown. Here, we demonstrated that TaSOC1 was a flowering repressor in the vernalization and photoperiod pathways by overexpression and knockout assays. We confirmed the physical interaction between TaSOC1 and TaVRN1 in wheat protoplasts and in planta, and further validated their genetic interplay. A Flowering Promoting Factor 1‐like gene TaFPF1‐2B was identified as a common downstream target of TaSOC1 and TaVRN1 through transcriptome and chromatin immunoprecipitation analyses. TaSOC1 competed with TaVRT2, another MADS‐box flowering regulator, to bind to TaVRN1; their coding genes synergistically control TaFPF1‐2B expression and flowering initiation in response to photoperiod and low temperature. We identified major haplotypes of TaSOC1 and found that TaSOC1‐Hap1 conferred earlier flowering than TaSOC1‐Hap2 and had been subjected to positive selection in wheat breeding. We also revealed that wheat SOC1 family members were important domestication loci and expanded by tandem and segmental duplication events. These findings offer new insights into the regulatory mechanism underlying flowering control along with useful genetic resources for wheat improvement. [ABSTRACT FROM AUTHOR]

The numbers per tiller (abundance) and per unit area of ground (population density) of the dominant cereal aphids in winter wheat crops, Sitobion avenae (Fabricius) and Metopolophium dirhodum (Walker) (Hemiptera: Aphididae) were recorded from 2000–2015. The results were used to test the hypothesis that at the population peak and 4 weeks previously, the population density of aphids in winter wheat crops (Triticum aestivum L., Poaceae) is uniformly distributed with respect to surface area and, as a result, their abundance is high in sparse stands (SPA) where the colonizing aphids are scattered among a small number of tillers (ca. 80 m−2), and low in dense stands (DEN) where they are scattered among a larger number of tillers (ca. 440 m−2). This prediction was verified for S. avenae, the population density of which did not significantly differ between SPA and DEN stands throughout this period. This species feeds mainly on the ears, whose nutritional quality for aphids varies little with crop density and during the course of ripening. In contrast and contrary to our prediction, the population density of M. dirhodum increased throughout this period more rapidly in SPA than in DEN stands. This species lives on leaves whose quality is better and remain present for longer during the course of ripening of the crop in SPA than DEN stands. The abundance of both species was higher in SPA than in DEN stands. Greater direct damage to cereal crops can be expected when using technologies and cultivars of winter wheat that need to be planted sparsely. [ABSTRACT FROM AUTHOR]

Summary: High‐molecular‐weight glutenin subunits (HMW‐GS), a major component of seed storage proteins (SSP) in wheat, largely determine processing quality. HMW‐GS encoded by GLU‐1 loci are mainly controlled at the transcriptional level by interactions between cis‐elements and transcription factors (TFs). We previously identified a conserved cis‐regulatory module CCRM1‐1 as the most essential cis‐element for Glu‐1 endosperm‐specific high expression. However, the TFs targeting CCRM1‐1 remained unknown. Here, we built the first DNA pull‐down plus liquid chromatography‐mass spectrometry platform in wheat and identified 31 TFs interacting with CCRM1‐1. TaB3‐2A1 as proof of concept was confirmed to bind to CCRM1‐1 by yeast one hybrid and electrophoretic mobility shift assays. Transactivation experiments demonstrated that TaB3‐2A1 repressed CCRM1‐1‐driven transcription activity. TaB3‐2A1 overexpression significantly reduced HMW‐GS and other SSP, but enhanced starch content. Transcriptome analyses confirmed that enhanced expression of TaB3‐2A1 down‐regulated SSP genes and up‐regulated starch synthesis‐related genes, such as TaAGPL3, TaAGPS2, TaGBSSI, TaSUS1 and TaSUS5, suggesting that it is an integrator modulating the balance of carbon and nitrogen metabolism. TaB3‐2A1 also had significant effects on agronomic traits, including heading date, plant height and grain weight. We identified two major haplotypes of TaB3‐2A1 and found that TaB3‐2A1‐Hap1 conferred lower seed protein content, but higher starch content, plant height and grain weight than TaB3‐2A1‐Hap2 and was subjected to positive selection in a panel of elite wheat cultivars. These findings provide a high‐efficiency tool to detect TFs binding to targeted promoters, considerable gene resources for dissecting regulatory mechanisms underlying Glu‐1 expression, and a useful gene for wheat improvement. [ABSTRACT FROM AUTHOR]

BACKGROUND: Climate changes associated with global warming are increasingly affecting the quality of cultivated crops. Cultivation at different altitudes and similar latitudes may offer an extraordinarily useful opportunity to obtain a diversificated dataset of climate variables and to further investigate their effect on crop quality. This study evaluated the effect of climate indices – temperature, rainfall precipitation and solar radiation data – on commercial parameters and the volatile organic compound (VOC) profile of wheat at harvest. RESULTS: Three common and durum wheat varieties, including two heritage wheats, were sown in experimental fields sited at three altitudes for 3 years consecutively, and they were analyzed for their yield, grading parameters, and VOC profiles. The datasets were processed by partial least squares regression (PLS‐R) and the results indicate that summer days (SU25) and diurnal temperature range (ΔT) are the climate indices mainly responsible for the VOC profile changes in both common and durum wheat. Accumulated growth degree days (GDD), consecutive dry days (CDD), and accumulated solar radiation (ASR) induced species‐specific responses. Terpenes represented the chemical class of VOCs most affected by stresses, followed by ketones and alcohols, which were affected by CDD, GDD, and ASR. CONCLUSION: This study showed a selective response of wheat to abiotic stresses associated with climate variables in terms of VOC synthesis. Its findings may be relevant in several fields, from plant ecology to agronomy and food quality, with implications for local economic strategies. © 2023 Society of Chemical Industry. [ABSTRACT FROM AUTHOR]

Drought is a major environmental stress limiting global wheat (Triticum aestivum) production. Exploring drought tolerance genes is important for improving drought adaptation in this crop. Here, we cloned and characterized TaTIP41, a novel drought tolerance gene in wheat. TaTIP41 is a putative conserved component of target of rapamycin (TOR) signaling, and the TaTIP41 homoeologs were expressed in response to drought stress and abscisic acid (ABA). The overexpression of TaTIP41 enhanced drought tolerance and the ABA response, including ABA‐induced stomatal closure, while its downregulation using RNA interference (RNAi) had the opposite effect. Furthermore, TaTIP41 physically interacted with TaTAP46, another conserved component of TOR signaling. Like TaTIP41, TaTAP46 positively regulated drought tolerance. Furthermore, TaTIP41 and TaTAP46 interacted with type‐2A protein phosphatase (PP2A) catalytic subunits, such as TaPP2A‐2, and inhibited their enzymatic activities. Silencing TaPP2A‐2 improved drought tolerance in wheat. Together, our findings provide new insights into the roles of TaTIP41 and TaTAP46 in the drought tolerance and ABA response in wheat, and their potential application in improving wheat environmental adaptability. [ABSTRACT FROM AUTHOR]

SUMMARY: Triticum aestivum L. (bread wheat) is a crop relied upon by billions of people around the world, as a major source of both income and calories. Rising global temperatures, however, pose a genuine threat to the livelihood of these people, as wheat growth and yields are extremely vulnerable to damage by heat stress. Here we present the YoGI wheat landrace panel, comprising 342 accessions that show remarkable phenotypic and genetic diversity thanks to their adaptation to different climates. We quantified the abundance of 110 790 transcripts from the panel and used these data to conduct weighted co‐expression network analysis and to identify hub genes in modules associated with abiotic stress tolerance. We found that the expression of three hub genes, all heat‐shock proteins (HSPs), were significantly correlated with early thermotolerance in a validation panel of landraces. These hub genes belong to the same module, with one (TraesCS4D01G207500.1) being a candidate master‐regulator potentially controlling the expression of the other two hub genes, as well as a suite of other HSPs and heat‐stress transcription factors (HSFs). In this work, therefore, we identify three validated hub genes, the expression of which can serve as markers of thermotolerance during early development, and suggest that TraesCS4D01G207500.1 is a potential master regulator of HSP and HSF expression – presenting the YoGI landrace panel as an invaluable tool for breeders wishing to determine and introduce novel alleles into modern varieties, for the production of climate‐resilient crops. Significance Statement: Global wheat yields and crop growth are vulnerable to damage by heat stress, a threat that is likely to become more common globally in the coming years through climate change. Conducting weighted co‐expression network analysis on 337 diverse landrace accessions, we identified and validated TraesCS4D01G207500.1 as a hub gene that can be used as a marker of early thermotolerance, seemingly via the regulation of the expression of a suite of HSP and HSF genes. [ABSTRACT FROM AUTHOR]

PHOSPHORUS‐STARVATION TOLERANCE 1 (OsPSTOL1) is a variably present gene that benefits crown root growth and phosphorus (P) sufficiency in rice (Oryza sativa). To explore the ecophysiological importance of this gene, we performed a biogeographic survey of landraces and cultivars, confirming that functional OsPSTOL1 alleles prevail in low nutrient and drought‐prone rainfed ecosystems, whereas loss‐of‐function and absence haplotypes predominate in control‐irrigated paddy varieties of east Asia. An evolutionary history analysis of OsPSTOL1 and related genes in cereal, determined it and other genes are kinase‐only domain derivatives of membrane‐associated receptor like kinases. Finally, to evaluate the potential value of this kinase of unknown function in another Gramineae, wheat (Triticum aestivum) lines overexpressing OsPSTOL1 were evaluated under field and controlled low P conditions. OsPSTOL1 enhances growth, crown root number, and overall root plasticity under low P in wheat. Survey of root and shoot crown transcriptomes at two developmental stages identifies transcription factors that are differentially regulated in OsPSTOL1 wheat that are similarly controlled by the gene in rice. In wheat, OsPSTOL1 alters the timing and amplitude of regulators of root development in dry soils and hastens induction of the core P‐starvation response. OsPSTOL1 and related genes may aid more sustainable cultivation of cereal crops. Summary statement: Might a rice gene that controls root plasticity confer a similar benefit in another grain crop? Here, we evaluate the genetic variation and evolutionary history of OsPSTOL1 and demonstrate its impact in wheat. [ABSTRACT FROM AUTHOR]

Summary: Blumeria graminis f. sp. tritici (Bgt) is a globally important fungal pathogen of wheat that can rapidly evolve to defeat wheat powdery mildew (Pm) resistance genes. Despite periodic regional deployment of the Pm1a resistance gene in US wheat production, Bgt strains that overcome Pm1a have been notably nonpersistent in the United States, while on other continents, they are more widely established.A genome‐wide association study (GWAS) was conducted to map sequence variants associated with Pm1a virulence in 216 Bgt isolates from six countries, including the United States. A virulence variant apparently unique to Bgt isolates from the United States was detected in the previously mapped gene AvrPm1a (BgtE‐5612) on Bgt chromosome 6; an in vitro growth assay suggested no fitness reduction associated with this variant.A gene on Bgt chromosome 8, Bgt‐51526, was shown to function as a second determinant of Pm1a virulence, and despite < 30% amino acid identity, BGT‐51526 and BGTE‐5612 were predicted to share > 85% of their secondary structure. A co‐expression study in Nicotiana benthamiana showed that BGTE‐5612 and BGT‐51526 each produce a PM1A‐dependent hypersensitive response.More than one member of a B. graminis effector family can be recognized by a single wheat immune receptor, and a two‐gene model is necessary to explain virulence to Pm1a. [ABSTRACT FROM AUTHOR]

SUMMARY: Because allohexaploid wheat genome contains ABD subgenomes, how the expression of homoeologous genes is coordinated remains largely unknown, particularly at the co‐transcriptional level. Alternative polyadenylation (APA) is an important part of co‐transcriptional regulation, which is crucial in developmental processes and stress responses. Drought stress is a major threat to the stable yield of wheat. Focusing on APA, we used poly(A) tag sequencing to track poly(A) site dynamics in wheat under drought stress. The results showed that drought stress led to extensive APA involving 37–47% of differentially expressed genes in wheat. Significant poly(A) site switching was found in stress‐responsive genes. Interestingly, homoeologous genes exhibit unequal numbers of poly(A) sites, divergent APA patterns with tissue specificity and time‐course dynamics, and distinct 3′‐UTR length changes. Moreover, differentially expressed transcripts in leaves and roots used different poly(A) signals, the up‐ and downregulated isoforms had distinct preferences for non‐canonical poly(A) sites. Genes that encode key polyadenylation factors showed differential expression patterns under drought stress. In summary, poly(A) signals and the changes in core poly(A) factors may widely affect the selection of poly(A) sites and gene expression levels during the response to drought stress, and divergent APA patterns among homoeologous genes add extensive plasticity to this responsive network. These results not only reveal the significant role of APA in drought stress response, but also provide a fresh perspective on how homoeologous genes contribute to adaptability through transcriptome diversity. In addition, this work provides information about the ends of transcripts for a better annotation of the wheat genome. Significance Statement: The functional conservation and divergence in a polyploidy genome across the subgenomes of wheat have always been intriguing. From the angle of APA, this work not only reveals the widespread occurrence of APA in the allohexaploid wheat and the important role it plays, but also broadens the knowledge of how homoeologous genes contribute to the adaptability and transcriptional plasticity in a polyploidy genome through RNA processing. [ABSTRACT FROM AUTHOR]

Summary: Phased, small interfering RNAs (phasiRNAs) are important for plant anther development, especially for male sterility. PhasiRNA biogenesis is dependent on genes like RNA polymerase 6 (RDR6), DICER‐LIKE 4 (DCL4), or DCL5 to produce 21‐ or 24 nucleotide (nt) double‐strand small RNAs. Here, we generated mutants of DCL4, DCL5 and RDR6 using CRISPR/Cas9 system and studied their effects on plant reproductive development and phasiRNA production in wheat. We found that RDR6 mutation caused sever consequence throughout plant development starting from seed germination and the dcl4 mutants grew weaker with thorough male sterility, while dcl5 plants developed normally but exhibited male sterility. Correspondingly, DCL4 and DCL5, respectively, specified 21‐ and 24‐nt phasiRNA biogenesis, while RDR6 contributed to both. Also, the three key genes evolved differently in wheat, with TaDCL5‐A/B becoming non‐functioning and TaRDR6‐A being lost after polyploidization. Furthermore, we found that PHAS genes (phasiRNA precursors) identified via phasiRNAs diverged rapidly among sub‐genomes of polyploid wheat. Despite no similarity being found among phasiRNAs of grasses, their targets were enriched for similar biological functions. In light of the important roles of phasiRNA pathways in gametophyte development, genetic dissection of the function of key genes may help generate male sterile lines suitable for hybrid wheat breeding. [ABSTRACT FROM AUTHOR]

Fungicides play an important role in wheat blast management in Brazil. To determine whether the reported patterns of fungicide resistance are present in Pyricularia oryzae populations from wheat and signal grass grown in Minas Gerais state, Brazil, we characterized the sensitivity of a contemporary collection (2018 and 2019) of 64 fungal isolates to seven fungicides belonging to DMI (tebuconazole and epoxiconazole), QoI (azoxystrobin and pyraclostrobin) and SDHI (bixafen, fluxapyroxad and benzovindiflupyr) groups. We further assessed if the differential sensitivity of selected isolates affected disease control using commercial doses of the fungicides applied protectively (prior to inoculation) on the heads of potted plants. Despite finding relatively low levels of in vitro sensitivity to some of the fungicides (e.g., azoxystrobin, fluxapyroxad and bixafen), control efficacies (percentage reductions in head blast severity) were significantly higher than the untreated plants and not affected by the sensitivity of the strain to all fungicides but azoxystrobin. The levels of head blast control for some of the fungicides (>70%) in the greenhouse were similar to or higher than previously reported in the field. The molecular data for a set of isolates representing different fungicide sensitivity levels were generally not predictive of the phenotype. The wheat‐infecting isolates were less sensitive than signal grass‐infecting isolates to all fungicides including azoxystrobin. Our study not only confirms previous reports of low sensitivity to some fungicides, but it also suggests options among site‐specific fungicides, most notably tebuconazole and benzovindiflupyr, that can still be used under an antiresistance strategy such as coformulation with multisite fungicides. [ABSTRACT FROM AUTHOR]

Summary: During stem elongation, wheat (Triticum aestivum) increases its stem carbohydrate content before anthesis as a reserve for grain filling. Hydraulic functioning during this mobilization process is not well understood, and contradictory results exist on the direct effect of drought on carbohydrate mobilization.In a dedicated experiment, wheat plants were subjected to drought stress during carbohydrate mobilization. Measurements, important to better understand stem physiology, showed some unexpected patterns that could not be explained by our current knowledge on water transport. Traditional water flow and storage models failed to properly describe the drought response in wheat stems during carbohydrate mobilization. To explain the measured patterns, hypotheses were formulated and integrated in a dedicated model for wheat.The new mechanistic model simulates two hypothetical water storage compartments: one where water is quickly exchanged with the xylem and one that contains the carbohydrate storage. Water exchange between these compartments is turgor‐driven. The model was able to simulate the measured increase in stored carbohydrate concentrations with a decrease in water content and stem diameter.Calibration of the model showed the importance of turgor‐driven apoplastic water flow during carbohydrate mobilization. This resulted in an increase in stem hydraulic capacitance, which became more important under drought stress. [ABSTRACT FROM AUTHOR]

Summary: Aegilops tauschii is the diploid progenitor of the wheat D subgenome and a valuable resource for wheat breeding, yet, genetic analysis of resistance against Fusarium head blight (FHB) and the major Fusarium mycotoxin deoxynivalenol (DON) is lacking. We treated a panel of 147 Ae. tauschii accessions with either Fusarium graminearum spores or DON solution and recorded the associated disease spread or toxin‐induced bleaching. A k‐mer‐based association mapping pipeline dissected the genetic basis of resistance and identified candidate genes. After DON infiltration nine accessions revealed severe bleaching symptoms concomitant with lower conversion rates of DON into the non‐toxic DON‐3‐O‐glucoside. We identified the gene AET5Gv20385300 on chromosome 5D encoding a uridine diphosphate (UDP)‐glucosyltransferase (UGT) as the causal variant and the mutant allele resulting in a truncated protein was only found in the nine susceptible accessions. This UGT is also polymorphic in hexaploid wheat and when expressed in Saccharomyces cerevisiae only the full‐length gene conferred resistance against DON. Analysing the D subgenome helped to elucidate the genetic control of FHB resistance and identified a UGT involved in DON detoxification in Ae. tauschii and hexaploid wheat. This resistance mechanism is highly conserved since the UGT is orthologous to the barley UGT HvUGT13248 indicating descent from a common ancestor of wheat and barley. [ABSTRACT FROM AUTHOR]

SUMMARY: Puccinia striiformis f. sp. tritici (Pst), the causal agent of stripe rust, is a destructive pathogen of Triticum aestivum (wheat), threatening wheat production worldwide. Pst delivers hundreds of effectors to manipulate processes in its hosts during infection. The SGT1 (suppressor of the G2 allele of skp1), RAR1 (required for Mla12 resistance) and HSP90 (heat‐shock protein 90) proteins form a chaperone complex that acts as a core modulator in plant immunity. However, little is known about how Pst effectors target this immune component to suppress plant immunity. Here, we identified a Pst effector PstSIE1 that interacts with TaSGT1 in wheat and is upregulated during the early infection stage. Transient expression of PstSIE1 suppressed cell death in Nicotiana benthamiana induced by VmE02 and PcNLP2. Transgenic expression of PstSIE1‐RNAi constructs in wheat significantly reduced the virulence of Pst. Overexpression of PstSIE1 in wheat increased the number of rust pustules and reduced the accumulation of reactive oxygen species (ROS), indicating that PstSIE1 functions as an important pathogenicity factor in Pst. PstSIE1 was found to compete with TaRAR1 to bind TaSGT1, thus disrupting the formation of the TaRAR1–TaSGT1 subcomplex. Taken together, PstSIE1 is an important Pst effector targeting the immune component TaSGT1 and involved in suppressing wheat defense. Significance Statement: PstSIE1 is an important Pst effector targeting the immune component TaSGT1 and involved in suppressing wheat defense. This work broadens our understanding of how the molecular mechanisms of fungal effectors suppress plant immunity. [ABSTRACT FROM AUTHOR]

Spikelets compactness (SC) plays an important role in the yield improvement and Fusarium head blight (FHB) resistance of wheat. In this study, 151 recombinant inbred lines (RIL) derived from a cross between Yangmai 4 (YM4) and Yanzhan 1 (YZ1) and 143 RIL derived from a cross between Yangmai 5 (YM5) and YZ1 were used to construct two wheat hexaploidy genetic linkage maps, evaluate SC across three years and identify quantitative trait loci (QTL) for SC. A total of seven QTL were identified in two populations and explained the phenotypic variation ranging from 6.27% to 22.49%. Four QTL were co‐located in two genomic regions on chromosomes 2D and 6B, namely, QSc‐2D and QSc‐6B, respectively. Comparison analysis showed that QSc‐6B is a novel QTL, whereas QSc‐2D was located in the interval of Rht8. Significant additive effects of QSc‐2D and QSc‐6B on SC were observed in two RIL populations. Moreover, the positive alleles of these two QTL extremely significantly increased grain number per spike (GNS) but did not have a significantly negative effect on thousand‐grain weight (TGW), kernel length (KL) and kernel weight (KW). Whether in spray or point inoculation, QSc‐6B did not significantly influence FHB resistance in two populations. However, QSc‐2D significantly affected FHB resistance in spray inoculation in YM4/YZ1 population. A breeder‐friendly Kompetitive Allele‐Specific PCR (KASP) marker KASP_6B for QSc‐6B has been developed and verified in a set of 181 wheat cultivars/lines. Furthermore, we preliminary analysed the candidate genes within the physical interval of QSc‐6B by sequence similarity, spatial expression patterns and collinearity analysis. TraesCS6B03G1206900, TraesCS6B03G1214800 and TraesCS6B03G1218300 show specific expression in spikelet. These results provide valuable information for map‐based cloning of QSc‐6B and a useful tool for wheat breeding. [ABSTRACT FROM AUTHOR]

Root‐lesion nematodes (RLNs) Pratylenchus thornei and P. neglectus are globally important pathogens of cereal and pulse crops. These RLNs can occur together in farming systems and must be managed concurrently to minimize substantial yield losses in intolerant crop cultivars. Australian wheat cultivars with resistance to P. neglectus, have either the Rlnn1 resistance gene, which provides a high level of resistance but is linked with yellow flour colour that reduces cultivar marketability for bread production, or QRlnn.lrc‐2B, which provides moderate resistance. We evaluated a collection of 91 P. thornei‐resistant Iranian landrace wheats (ILWs) for their resistance to P. neglectus in four glasshouse experiments to (a) identify genotypes with resistance to both RLNs, (b) determine if any genotypes carried Rlnn1 and/or QRlnn.lrc‐2B and (c) develop ILW‐derived advanced breeding lines (ABLs) with resistance to both RLNs. A factor analytic linear mixed model (FA‐1) that explained 70% of the genetic variation, where the genetic correlations between the experiments ranged from 0.54 to 0.77, was used for the combined analysis of all experiments. Seven P. neglectus‐resistant genotypes were identified, with five that had potentially novel resistance. Subsequently, six breeding lines that were resistant to both RLNs were developed by crossing six ILWs with Australian cultivars and selecting for resistance in each generation. Both the ILWs and ABLs will be valuable genetic resources for wheat breeders to develop cultivars with dual resistance, enabling better management of mixed RLN populations with novel P. neglectus resistance that potentially is not linked with yellow flour colour. [ABSTRACT FROM AUTHOR]

Summary: Abscisic acid (ABA) receptors are considered as the targeted manipulation of ABA sensitivity and water productivity in plants. Regulation of their stability or activity will directly affect ABA signalling. Mitogen‐activated protein kinase (MAPK) cascades link multiple environmental and plant developmental cues. However, the molecular mechanism of ABA signalling and MAPK cascade interaction remains largely elusive.TaMPK3 overexpression decreases drought tolerance and wheat sensitivity to ABA, significantly weakening ABA's inhibitory effects on growth. Under drought stress, overexpression lines show lower survival rates, shoot fresh weight and proline content, but higher malondialdehyde levels at seedling stage, as well as decreased grain width and 1000 grain weight in both glasshouse and field conditions at the adult stage. TaMPK3‐RNAi increases drought tolerance.TaMPK3 interaction with TaPYL4 leads to decreased TaPYL4 levels by promoting its ubiquitin‐mediated degradation, whereas ABA treatment diminishes TaMPK3–TaPYL interactions. In addition, the expression of ABA signalling proteins is impaired in TaMPK3‐overexpressing wheat plants under ABA treatment. The MPK3‐PYL interaction module was found to be conserved across monocots and dicots.Our results suggest that the MPK3‐PYL module could serve as a negative regulatory mechanism for balancing appropriate drought stress response with normal plant growth signalling in wheat. [ABSTRACT FROM AUTHOR]

Promoting seed germination after short episodes of heat stress during the wheat grain filling stage is a serious problem that results in pre-harvest sprouting. The plant hormones abscisic acid (ABA), gibberellins (GAs), and ethylene (ETH) are well known to be involved in germination control. However, the genes associated with the metabolism and responsiveness of these hormones to heat stress during wheat grain filling are not well understood. Transcriptomic analysis was carried out to explore the mechanisms controlling seed germination under five days (15-20 days after flowering, DAF) of heat stress (20, 24, 28, and 32°C) in wheat grains at 15-30 DAF using comparative RNA sequencing. A dataset of 2073 differentially regulated genes was used to help elucidate the molecular mechanisms that respond to heat stress and affect seed germination in wheat. Some genes related to ABA, GA, and ETH biosynthesis, transport, and signaling had significantly different expression levels under heat stress. Among these genes, the transcriptional alterations of plant hormone-related genes, such as NCED9, AAO3, CYP707A2, GA20ox, and SAM1 uncovered here, provide a foundation for identifying key players involved in determining seed dormancy and germination. The expression levels of many germination-related genes did not linearly increase with increasing temperature. In this study, 28°C is a threshold of temperature tolerance during the grain filling stage. Heat stress, especially extremely high temperature (>28°C), represses ABA-related gene expression and promotes seed germination. [ABSTRACT FROM AUTHOR]

The worldwide spread of wheat yellow rust lineage PstS1/S2 adapted to higher temperatures prompted us to investigate how diverse temperature responses of this lineage are in the Middle East, where diversity was previously observed within this lineage for pathotypes and genotypes. Here we highlight the diversity of response to temperature within a PstS1/S2 population. Twenty‐six isolates from eight countries and different altitudes, which were tested under four combinations of cold and warm incubation and postincubation temperature conditions, showed diversity for infection efficiency (IE) and latency period (LP). IE of the various isolates ranged from 5.8% to 13.7% under cold (5°C) and 0.04% to 1% under warm (20°C) incubation temperatures. LP varied from 10.2 days under warm to 4.43 days under cold incubation. LP of isolates from the same country could differ by 2 days. Significant differences in thermal aptitudes of the isolates were observed between and within countries. IE and LP diversity was not related to altitude origin of the isolates on the whole; however, a trade‐off between IE and LP was observed for isolates from low altitude (<400 m) under a warm regime. We showed diversity for thermal aptitude for IE and LP of isolates belonging to the same PstS1/S2 lineage. Understanding Pst temperature aptitude among geographically distant isolates of the same clonal lineage may help to identify the geographic range of pathogens and also to improve forecast models or breeding programmes. [ABSTRACT FROM AUTHOR]

SUMMARY: High temperature stress inhibits photosynthesis and threatens wheat production. One measure of photosynthetic heat tolerance is Tcrit – the critical temperature at which incipient damage to photosystem II (PSII) occurs. This trait could be improved in wheat by exploiting genetic variation and genotype‐by‐environment interactions (GEI). Flag leaf Tcrit of 54 wheat genotypes was evaluated in 12 thermal environments over 3 years in Australia, and analysed using linear mixed models to assess GEI effects. Nine of the 12 environments had significant genetic effects and highly variable broad‐sense heritability (H2 ranged from 0.15 to 0.75). Tcrit GEI was variable, with 55.6% of the genetic variance across environments accounted for by the factor analytic model. Mean daily growth temperature in the month preceding anthesis was the most influential environmental driver of Tcrit GEI, suggesting biochemical, physiological and structural adjustments to temperature requiring different durations to manifest. These changes help protect or repair PSII upon exposure to heat stress, and may improve carbon assimilation under high temperature. To support breeding efforts to improve wheat performance under high temperature, we identified genotypes superior to commercial cultivars commonly grown by farmers, and demonstrated potential for developing genotypes with greater photosynthetic heat tolerance. Significance Statement: Here we examined intraspecific variation in photosynthetic heat tolerance (the critical temperature at which damage to PSII occurs ‐ Tcrit); effects of genotype‐by‐environment interactions on Tcrit; and environmental drivers of Tcrit in wheat – the most widely cultivated crop. Our study provides a sound framework for identifying germplasm with superior Tcrit relative to well‐adapted commercial cultivars. This framework can be applied in agricultural and ecological settings in selection of more heat tolerant species. [ABSTRACT FROM AUTHOR]

Summary: Plant architecture is a key determinant of crop productivity and adaptation. The highly conserved microRNA319 (miR319) family functions in various biological processes, but little is known about how miR319 regulates plant architecture in wheat (Triticum aestivum).Here, we determined that the miR319/TaGAMYB3 module controls plant architecture and grain yield in common wheat. Repressing tae‐miR319 using short tandem target mimics resulted in favorable plant architecture traits, including increased plant height, reduced tiller number, enlarged spikes and flag leaves, and thicker culms, as well as enhanced grain yield in field plot tests. Overexpressing tae‐miR319 had the opposite effects on plant architecture and grain yield.Although both TaPCF8 and TaGAMYB3 were identified as miR319 target genes, genetic complementation assays demonstrated that only miR319‐resistant TaGAMYB3 (rTaGAMYB3) abolished tae‐miR319‐mediated growth inhibition of flag leaves and spikes. TaGAMYB3 functions as a transcriptional activator of downstream genes, including TaPSKR1, TaXTH23, TaMADS5 and TaMADS51, by binding to their promoters. Furthermore, TaGAMYB3 physically interacts with TaBA1, an important regulator of spike development, to additively activate the transcription of downstream genes such as TaMADS5.Our findings provide insight into how the miR319/TaGAMYB3 module regulates plant architecture and improves grain yield in common wheat. [ABSTRACT FROM AUTHOR]

Septoria tritici blotch (STB) is one of the main causes of wheat yield loss in the world. Apart from good agrotechnical practice, disease‐resistant cultivars are required to prevent yield losses. Breeding of such cultivars is time‐consuming and laborious, mainly due to the quantitative character of such resistance, but can be shortened using precise phenotyping. We show that fluorescence parameters, collected in a high‐throughput system, can be used for estimating wheat STB resistance. Using machine learning methods (deep learning network, random forest), we demonstrate that disease resistance based on the percentage of necrotic leaf area can be estimated by alterations in fluorescence parameters 8 days after inoculation, and before disease symptoms become visible. Moreover, we applied a random forest classifier to assess the significance of fluorescence parameters in an informative way (classification accuracy = 64.84%, p value [Accuracy >No information rate] <0.001). Based on this, we observed extensive alterations in parameters of sensitive cultivars, reflecting worse photosynthetic performance. The most highly altered parameters were fluorescence intensity measurements and descriptors of energy flux through photosystem II (PSII) reaction centres towards PSI such as F2, F3 and F4 fluorescence intensities at 0.1, 0.270 and 2 ms; initial to maximal fluorescence ratio (F0/Fm); the time derivative of relative variable fluorescence (dV/dT0); maximum quantum yield of PSII in the dark‐adapted state (Fv/Fm); maximum electron transport flux at PSII (ET0/RC); maximum trapped excitonic energy flux per excited PSII cross section (CS) at time T0 (TR0/CS0); and quantum efficiency of the reduction of end acceptors dR0 [dR0/(1 − dR0)]. [ABSTRACT FROM AUTHOR]

SUMMARY: Cultivated wheat is continually exposed to various pathogens. Blumeria graminis f. sp. tritici (Bgt) causes powdery mildew disease and significant yield loss. Pm60 was cloned from Triticum urartu and confers race‐specific powdery mildew resistance in wheat. Pm60a and Pm60b are allelic variants of Pm60 and have two leucine‐rich repeat motifs deletions and insertions, respectively, which were detected in other T. urartu accessions. Through map‐based cloning, virus‐induced gene silencing, and stable transformation assays, we demonstrated that Pm60a and Pm60b conferred Bgt E09 resistance resembling that provided by Pm60. However, the homozygous Pm60a (but not Pm60 or Pm60b) transformants driven by the native promoters lacked race‐specific resistance when they were inoculated with Bgt E18. As all three T. urartu accessions contained the three foregoing alleles, they had high resistance to Bgt E18. Pyramiding Pm60a with either of the allelic genes in F1 plants did not cause mutual allele suppression or interference with Bgt E18 resistance. Deletion (but not insertion) of the two leucine‐rich repeat motifs in Pm60a substantially narrowed the resistance spectrum. In T. urartu accession PI428210, we identified another locus adjacent to Pm60a and resistant to Bgt E18. Characterization of the alleles at the Pm60 locus revealed their diversity and similarity and may facilitate wheat breeding for resistance to powdery mildew disease caused by B. graminis f. sp. tritici. Significance Statement: In the present study, we identified Pm60a and Pm60b, which are allelic variants at the Pm60 locus. The three foregoing alleles confer resistance to Bgt E09, which causes powdery mildew disease in wheat. Bgt E18 resistance was only absent in Pm60a transgenic plants as this allele has two leucine‐rich repeat (LRR) motif deletions. Pyramiding Pm60a with Pm60 or Pm60b did not hinder Bgt E18 resistance in wheat. Assays involving 54 Bgt isolates confirmed that deletion of the two LRR motifs in Pm60a markedly narrows the Bgt resistance spectrum. In contrast, the insertion of additional LRR motifs in Pm60b did not enhance Bgt resistance. The foregoing findings demonstrated diversity and similarity among alleles at the Pm60 locus and may support wheat breeding for powdery mildew disease resistance. [ABSTRACT FROM AUTHOR]

Wheat farmers rely on fungicides to protect fields against several foliar and flowering diseases, including Fusarium head blight (FHB). A range of active ingredients is used in isolation or in dual premixes that include a dimethylation inhibitor (DMI) or a quinone outside inhibitor (QoI) fungicide. Comprehensive information about fungicide resistance in F. graminearum is available for DMIs, while for QoIs the data are scarce. We characterized 225 strains obtained from two states in southern Brazil, Rio Grande do Sul (RS) and Paraná (PR), in relation to their response to two QoIs. The median EC50 (effective concentration leading to 50% inhibition of conidial germination) value for azoxystrobin (n = 25 isolates) was 2.20 μg/ml in the PR population and 4.04 μg/ml in the RS population. For pyraclostrobin (n = 50), the median EC50 was 0.28 μg/ml in the PR population and 0.24 μg/ml in the RS population. Evidence of cross‐resistance could not be detected. Screening using a discriminatory dose (DD) for azoxystrobin in a larger number of isolates from PR (n = 75) and RS (n = 100) states allowed the detection of 50% and 28% sensitive strains, respectively. Using the DD for pyraclostrobin, 33% and 18.8% were classified as less sensitive in the PR and RS isolates, respectively. In RS, the frequency of less‐sensitive isolates increased over time (2007–2011). No point mutation at any of the target spots (F129L, G137R, G143A) was detected. Our results represent an important step towards the establishment of a sensitivity profile for two of the most commonly used QoIs in commercial premixes targeting FHB control. [ABSTRACT FROM AUTHOR]

Summary: STAUROSPORINE AND TEMPERATURE SENSITIVE3 (STT3) is a catalytic subunit of oligosaccharyltransferase, which is important for asparagine‐linked glycosylation. Sharp eyespot, caused by the necrotrophic fungal pathogen Rhizoctonia cerealis, is a devastating disease of bread wheat. However, the molecular mechanisms underlying wheat defense against R. cerealis are still largely unclear. In this study, we identified TaSTT3a and TaSTT3b, two STT3 subunit genes from wheat and reported their functional roles in wheat defense against R. cerealis and increasing grain weight. The transcript abundance of TaSTT3b‐2B was associated with the degree of wheat resistance to R. cerealis and induced by both R. cerealis and exogenous jasmonic acid (JA). Overexpression of TaSTT3b‐2B significantly enhanced resistance to R. cerealis, grain weight, and JA content in transgenic wheat subjected to R. cerealis stress, while silencing of TaSTT3b‐2B compromised resistance of wheat to R. cerealis. Transcriptomic analysis showed that TaSTT3b‐2B affected the expression of a series of defense‐related genes and JA biosynthesis–related genes, as well as genes coding starch synthase and sucrose synthase. Application of exogenous JA elevated expression levels of the abovementioned defense‐ and grain weight–related genes, and rescuing the resistance of TaSTT3b‐2B–silenced wheat to R. cerealis, while pretreatment with sodium diethyldithiocarbamate, an inhibitor of JA synthesis, attenuated the TaSTT3b‐2B–mediated resistance to R. cerealis, suggesting that TaSTT3b‐2B played critical roles in regulating R. cerealis resistance and grain weight via JA biosynthesis. Altogether, this study reveals new functional roles of TaSTT3b‐2B in regulating plant innate immunity and grain weight, and illustrates its potential application value for wheat molecular breeding. [ABSTRACT FROM AUTHOR]

Summary: Only a few transcriptional regulators of seed storage protein (SSP) genes have been identified in common wheat (Triticum aestivum L.). Coexpression analysis could be an efficient approach to characterize novel transcriptional regulators at the genome‐scale considering the correlated expression between transcriptional regulators and target genes. As the A genome donor of common wheat, Triticum urartu is more suitable for coexpression analysis than common wheat considering the diploid genome and single gene copy. In this work, the transcriptome dynamics in endosperm of T. urartu throughout grain filling were revealed by RNA‐Seq analysis. In the coexpression analysis, a total of 71 transcription factors (TFs) from 23 families were found to be coexpressed with SSP genes. Among these TFs, TuNAC77 enhanced the transcription of SSP genes by binding to cis‐elements distributed in promoters. The homolog of TuNAC77 in common wheat, TaNAC77, shared an identical function, and the total SSPs were reduced by about 24% in common wheat when TaNAC77 was knocked down. This is the first genome‐wide identification of transcriptional regulators of SSP genes in wheat, and the newly characterized transcriptional regulators will undoubtedly expand our knowledge of the transcriptional regulation of SSP synthesis. Significance Statement: This genome‐wide analysis followed by experimental validation identified novel candidate regulators of protein storage in seeds of common wheat. [ABSTRACT FROM AUTHOR]

Summary: Nocturnal stomatal conductance (gsn) represents a significant source of water loss, with implications for metabolism, thermal regulation and water‐use efficiency. With increasing nocturnal temperatures due to climate change, it is vital to identify and understand variation in the magnitude and responses of gsn in major crops.We assessed interspecific variation in gsn and daytime stomatal conductance (gs) in a wild relative and modern spring wheat genotype. To investigate intraspecific variation, we grew six modern wheat genotypes and two landraces under well watered, simulated field conditions.For the diurnal data, higher gsn in the wild relative was associated with significantly lower nocturnal respiration and higher daytime CO2 assimilation while both species exhibited declines in gsn post‐dusk and pre‐dawn. Lifetime gsn achieved rates of 5.7–18.9% of gs. Magnitude of gsn was genotype specific 'and positively correlated with gs. gsn and gs were significantly higher on the adaxial surface. No relationship was determined between harvest characteristics, stomatal morphology and gsn, while cuticular conductance was genotype specific. Finally, for the majority of genotypes, gsn declined with age.Here we present the discovery that variation in gsn occurs across developmental, morphological and temporal scales in nonstressed wheat, presenting opportunities for exploiting intrinsic variation under heat or water stressed conditions. [ABSTRACT FROM AUTHOR]

Aegilops tauschii, the progenitor of the wheat D genome, contains extensive diversity for biotic and abiotic resistance. Lr21 is a leaf rust resistance gene, which did not enter the initial gene flow from Ae. tauschii into hexaploid wheat due to restrictive hybridization events. Here, we used population genetics and high‐resolution comparative genomics to study evolutionary and functional divergence of Lr21 in diploid and hexaploid wheats. Population genetics identified the original Lr21, lr21‐1 and lr21‐2 alleles and their evolutionary history among Ae. tauschii accessions. Comparative genetics of Lr21 variants between Ae. tauschii and cultivated genotypes suggested at least two independent polyploidization events in bread wheat evolution. Further, a recent re‐birth of a unique Lr21‐tbk allele and its neofunctionalization was discovered in the hexaploid wheat cv. Tobak. Altogether, four independent alleles were investigated and validated for leaf rust resistance in diploid, synthetic hexaploid and cultivated wheat backgrounds. Besides seedling resistance, we uncover a new role of the Lr21 gene in conferring an adult plant field resistance. Seedling and adult plant resistance turned out to be correlated with developmentally dependent variation in Lr21 expression. Our results contribute to understand Lr21 evolution and its role in establishing a broad‐spectrum leaf rust resistance in wheat. The present study uncovers evolutionary divergence of Lr21 in Aegilops tauschii accessions and its transfer to hexaploid wheat as well as its new role in conferring broad‐spectrum resistance against leaf rust in bread wheat. [ABSTRACT FROM AUTHOR]

With increasing plastic production and consumption, large amounts of polystyrene nanoplastics are accumulated in soil due to improper disposal causing pollution and deleterious effects to environment. However, little information is available about how to alleviate the adverse impacts of nanoplastics on crops. In this study, the involvement of melatonin in modulating nanoplastic uptake, translocation, and toxicity in wheat plant was investigated. The results demonstrated that exogenous melatonin application reduced the nanoplastic uptake by roots and their translocation to shoots via regulating the expression of genes associated with aquaporin, including the upregulation of the TIP2‐9, PIP2, PIP3, and PIP1.2 in leaves and TIP2‐9, PIP1‐5, PIP2, and PIP1.2 in roots. Melatonin activated the ROS scavenging system to maintain a better redox homeostasis and ameliorated the negative effects of nanoplastics on carbohydrate metabolism, hence ameliorated the plant growth and enhanced the tolerance to nanoplastics toxicity. This process was closely related to the exogenous melatonin application induced melatonin accumulation in leave. These results suggest that melatonin could alleviate the adverse effects of nanoplastics on wheat, and exogenous melatonin application might be used as a promising management strategy to sustain crop production in the nanoplastic‐polluted soils. [ABSTRACT FROM AUTHOR]

Summary: Efficient phosphate (Pi) uptake and utilisation are essential for promoting crop yield. However, the underlying molecular mechanism is still poorly understood in complex crop species such as hexaploid wheat. Here we report that TaPHT1;9‐4B and its transcriptional regulator TaMYB4‐7D function in Pi acquisition, translocation and plant growth in bread wheat.TaPHT1;9‐4B, a high‐affinity Pi transporter highly upregulated in roots by Pi deficiency, was identified using quantitative proteomics. Disruption of TaPHT1;9‐4B function by BSMV‐VIGS or CRISPR editing impaired wheat tolerance to Pi deprivation, whereas transgenic expression of TaPHT1;9‐4B in rice improved Pi uptake and plant growth. Using yeast‐one‐hybrid assay, we isolated TaMYB4‐7D, a R2R3 MYB transcription factor that could activate TaPHT1;9‐4B expression by binding to its promoter. Silencing TaMYB4‐7D decreased TaPHT1;9‐4B expression, Pi uptake and plant growth.Four promoter haplotypes were identified for TaPHT1;9‐4B, with Hap3 showing significant positive associations with TaPHT1;9‐4B transcript level, growth performance and phosphorus (P) content in wheat plants. A functional marker was therefore developed for tagging Hap3.Collectively, our data shed new light on the molecular mechanism controlling Pi acquisition and utilisation in bread wheat. TaPHT1;9‐4B and TaMYB4‐7D may aid further research towards the development of P efficient crop cultivars. [ABSTRACT FROM AUTHOR]

Septoria tritici blotch (STB), caused by the fungus Zymoseptoria tritici, is one of the most economically important diseases of wheat. Recently, both factors of a gene‐for‐gene interaction between Z. tritici and wheat, the wheat receptor‐like kinase Stb6 and the Z. tritici secreted effector protein AvrStb6, have been identified. Previous analyses revealed a high diversity of AvrStb6 haplotypes present in earlier Z. tritici isolate collections, with up to c.18% of analysed isolates possessing the avirulence isoform of AvrStb6 identical to that originally identified in the reference isolate IPO323. With Stb6 present in many commercial wheat cultivars globally, we aimed to assess potential changes in AvrStb6 genetic diversity and the incidence of haplotypes allowing evasion of Stb6‐mediated resistance in more recent Z. tritici populations. Here we show, using targeted resequencing of AvrStb6, that this gene is universally present in field isolates sampled from major wheat‐growing regions of the world in 2013–2017. However, in contrast to the data from previous AvrStb6 population studies, we report a complete absence of the originally described avirulence isoform of AvrStb6 amongst modern Z. tritici isolates. Moreover, a remarkably small number of haplotypes, each encoding AvrStb6 protein isoforms conditioning virulence on Stb6‐containing wheat, were found to predominate among modern Z. tritici isolates. A single virulence isoform of AvrStb6 was found to be particularly abundant throughout the global population. These findings indicate that, despite the ability of Z. tritici to sexually reproduce on resistant hosts, AvrStb6 avirulence haplotypes tend to be eliminated in subsequent populations. [ABSTRACT FROM AUTHOR]

Maintaining a steady increase of yields requires knowledge of plant stress physiology and modern techniques of quantitative data collection and analysis. Here, the chlorophyll fluorescence parameters are used for modelling of relative water content (RWC) in field‐grown wheat cultivars. RWC is commonly used for the detection of plant tolerance to temporary droughts, but its determination is laborious and does not meet the requirements of a mass test like fluorescence detection. The paper presents a beta generalized linear mixed model (GLMM) fitted for RWC prediction based on chlorophyll fluorescence data repeatedly measured over time. The nature of fluorescence parameters with the strong correlations between them leads to the use of a multilevel principal component analysis to overcome this issue prior to the fitting of the model. Furthermore, a beta generalized estimating equation (GEE) model is fitted for identifying population‐average effects of the parameters used. Finally, highly significant results in terms of prediction with the use of 10‐fold cross‐validation (RPearson‐CV = 0.86, MAECV = 0.0365, RMSECV = 0.048) were obtained. Moreover, the population‐average effects provide important information for the parameters used in RWC prediction. The beta GLMM can provide good predictions combined with important cultivar‐specific information. Conclusively, these implementations can be a useful tool for drought tolerance improvement. [ABSTRACT FROM AUTHOR]

SUMMARY: Grain filling and grain development are essential biological processes in the plant's life cycle, eventually contributing to the final seed yield and quality in all cereal crops. Studies of how the different wheat (Triticum aestivum L.) grain components contribute to the overall development of the seed are very scarce. We performed a proteomics and metabolomics analysis in four different developing components of the wheat grain (seed coat, embryo, endosperm, and cavity fluid) to characterize molecular processes during early and late grain development. In‐gel shotgun proteomics analysis at 12, 15, 20, and 26 days after anthesis (DAA) revealed 15 484 identified and quantified proteins, out of which 410 differentially expressed proteins were identified in the seed coat, 815 in the embryo, 372 in the endosperm, and 492 in the cavity fluid. The abundance of selected protein candidates revealed spatially and temporally resolved protein functions associated with development and grain filling. Multiple wheat protein isoforms involved in starch synthesis such as sucrose synthases, starch phosphorylase, granule‐bound and soluble starch synthase, pyruvate phosphate dikinase, 14‐3‐3 proteins as well as sugar precursors undergo a major tissue‐dependent change in abundance during wheat grain development suggesting an intimate interplay of starch biosynthesis control. Different isoforms of the protein disulfide isomerase family as well as glutamine levels, both involved in the glutenin macropolymer pattern, showed distinct spatial and temporal abundance, revealing their specific role as indicators of wheat gluten quality. Proteins binned into the functional category of cell growth/division and protein synthesis/degradation were more abundant in the early stages (12 and 15 DAA). At the metabolome level all tissues and especially the cavity fluid showed highly distinct metabolite profiles. The tissue‐specific data are integrated with biochemical networks to generate a comprehensive map of molecular processes during grain filling and developmental processes. [ABSTRACT FROM AUTHOR]

Semiochemical use is a promising way to reduce damage from pests by improving natural control in agro‐ecosystems. The aphid alarm pheromone (E)‐β‐farnesene (EβF) and herbivore‐induced methyl salicylate (MeSA) are two volatile cues to induce changes in aphid behavior with functional significance. Because of limitations related to the volatility and oxidization of EβF and MeSA under natural conditions, slow‐release and antioxidant techniques should be developed and optimized before application. Here, a slow‐release alginate bead of EβF mixed with MeSA was first designed and manufactured. We hypothesized that a mixture of these two semiochemicals could be effective in controlling Sitobion miscanthi in wheat crops. Both MeSA and EβF in alginate beads were released stably and continuously for at least 15 days in the laboratory, whereas EβF in paraffin oil and pure MeSA were released for only 2 and 7 days, respectively. In 2018 field experiments, EβF and MeSA alone or in association significantly decreased the abundance of alate and apterous aphids. An increased abundance of mummified aphids enhanced by higher parasitism rates was observed when using EβF and MeSA in association, with a significant reduction of apterous abundance, more so than EβF or MeSA alone. In 2019, plots treated with a mixture of EβF and MeSA showed significantly decreased abundance of alate and apterous aphids with higher parasitism rates compared with the control. The new slow‐release alginate bead containing a mixture of EβF with MeSA could be the most efficient formulation to control S. miscanthi population by attracting parasitoids in the wheat agro‐ecosystem. © 2021 Society of Chemical Industry. © 2021 Society of Chemical Industry. [ABSTRACT FROM AUTHOR]

SUMMARY: We expressed a bacterial glucan synthase (AgrobacteriumGlgA) in the cytosol of developing endosperm cells in wheat grains, to discover whether it could generate a glucan from cytosolic ADP‐glucose. Transgenic lines had high glucan synthase activity during grain filling, but did not accumulate glucan. Instead, grains accumulated very high concentrations of maltose. They had large volumes during development due to high water content, and very shrivelled grains at maturity. Starch synthesis was severely reduced. We propose that cytosolic glucan synthesized by the glucan synthase was immediately hydrolysed to maltose by cytosolic β‐amylase(s). Maltose accumulation resulted in a high osmotic potential in developing grain, drawing in excess water that stretched the seed coat and pericarp. Loss of water during grain maturation then led to shrinkage when the grains matured. Maltose accumulation is likely to account for the reduced starch synthesis in transgenic grains, through signalling and toxic effects. Using bioinformatics, we identify an isoform of β‐amylase likely to be responsible for maltose accumulation. Removal of this isoform through identification of TILLING mutants or genome editing, combined with co‐expression of heterologous glucan synthase and a glucan branching enzyme, may in future enable elevated yields of carbohydrate through simultaneous accumulation of starch and cytosolic glucan. Significance Statement: Grains of transgenic wheat expressing a bacterial glucan synthase in the endosperm cytosol did not accumulate cytosolic glucan but instead accumulated very high levels of maltose, presumably due to hydrolysis of nascent glucan by an endogenous β‐amylase. [ABSTRACT FROM AUTHOR]

Summary: Plant nucleotide‐binding leucine‐rich repeat receptors (NLRs) act as intracellular sensors for pathogen‐derived effector proteins and trigger an immune response, frequently resulting in the hypersensitive cell death response (HR) of the infected host cell. The wheat (Triticum aestivum) NLR Pm2 confers resistance against the fungal pathogen Blumeria graminis f. sp. tritici (Bgt) if the isolate contains the specific RNase‐like effector AvrPm2. We identified and isolated seven new Pm2 alleles (Pm2e–i) in the wheat D‐genome ancestor Aegilops tauschii and two new natural AvrPm2 haplotypes from Bgt. Upon transient co‐expression in Nicotiana benthamiana, we observed a variant‐specific HR of the Pm2 variants Pm2a and Pm2i towards AvrPm2 or its homolog from the AvrPm2 effector family, BgtE‐5843, respectively. Through the introduction of naturally occurring non‐synonymous single nucleotide polymorphisms and structure‐guided mutations, we identified single amino acids in both the wheat NLR Pm2 and the fungal effector proteins AvrPm2 and BgtE‐5843 responsible for the variant‐specific HR of the Pm2 variants. Exchanging these amino acids led to a modified HR of the Pm2–AvrPm2 interaction and allowed the identification of the effector head epitope, a 20‐amino‐acid long unit of AvrPm2 involved in the HR. Swapping of the AvrPm2 head epitope to the non‐HR‐triggering AvrPm2 family member BgtE‐5846 led to gain of a HR by Pm2a. Our study presents a molecular approach to identify crucial effector surface structures involved in the HR and demonstrates that natural and induced diversity in an immune receptor and its corresponding effectors can provide the basis for understanding and modifying NLR–effector specificity. Significance Statement: We have identified seven new alleles of the wheat nucleotide‐binding leucine‐rich repeat immune receptor (NLR) Pm2 and analyzed the molecular determinants defining the specificity towards the corresponding fungal effector proteins from wheat powdery mildew. Our study presents a molecular approach to identify crucial effector surface structures involved in immune responses and demonstrates that natural and induced diversity in an immune receptor and its corresponding effectors can provide the basis for understanding and modifying NLR–effector specificity. [ABSTRACT FROM AUTHOR]

Summary: The synthesis of seed storage protein (SSP) is mainly regulated at the transcriptional level. However, few transcriptional regulators of SSP synthesis have been characterized in common wheat (Triticum aestivum) owing to the complex genome. As the A genome donor of common wheat, Triticum urartu could be an elite model in wheat research considering its simple genome. Here, a novel NAC family transcription factor TuSPR from T. urartu was found preferentially expressed in developing endosperm during grain‐filling stages. In common wheat transgenically overexpressing TuSPR, the content of total SSPs was reduced by c. 15.97% attributed to the transcription declines of SSP genes. Both in vitro and in vivo assays showed that TuSPR bound to the cis‐element 5′‐CANNTG‐3′ distributed in SSP gene promoters and suppressed the transcription. The homolog in common wheat TaSPR shared a conserved function with TuSPR on SSP synthesis suppression. The knock‐down of TaSPR in common wheat resulted in 7.07%–20.34% increases in the total SSPs. Both TuSPR and TaSPR could be superior targets in genetic engineering to manipulate SSP content in wheat, and this work undoubtedly expands our knowledge of SSP gene regulation. [ABSTRACT FROM AUTHOR]

Background: Salinity is one of the main threats that can cause crop yield losses by limiting plant growth. Several important cultivated crops including wheat exhibit sensitivity to salt stress by severe yield reductions. Approaches to improve salt resistance are critical for improved food production in the future. Aims: Silicon (Si) is well known for its role as quasi‐essential element to enhance growth and development of monocot food crops, especially under a variety of abiotic and biotic stress conditions. However, the mechanism of Si‐induced stress resistance is still not fully understood. Here, we investigate the biochemical basis of Si‐induced salt resistance of wheat. Methods: Effect of Si treatment on wheat plants under salt stress was measured by biochemical studies of photosynthetic pigments, ions, oxidative stress indicators, enzymatic and non‐enzymatic antioxidant systems, phytohormones along with expression of two Si‐transport genes. Results: Our results suggest that Si‐induced salt resistance of wheat involves a signaling cascade that triggers the biochemical defense responses. Salinity induces production of the stress hormone abscisic acid that upregulates the expression of Si‐influx transporter gene TaLsi1 and Si‐efflux transporter gene TaLsi2, in the presence of soluble Si in cytosol, and this triggers jasmonate‐mediated activation of both enzymatic and non‐enzymatic antioxidant systems along with osmolyte production. Moreover, peculiar stoichiometric changes in photosynthetic pigments after Si application, as signatures of shaded condition, also lead us to the hypothesis that insoluble Si phytoliths in plants may aid a physical defense response by creating a "mechanically shielded leaf microenvironment" against a variety of abiotic and biotic stress conditions. Conclusion: This study demonstrates that Si activates biochemical defense responses against salt stress in wheat plants. [ABSTRACT FROM AUTHOR]

Summary: Pm1a, the first powdery mildew resistance gene described in wheat, is part of a complex resistance (R) gene cluster located in a distal region of chromosome 7AL that has suppressed genetic recombination.A nucleotide‐binding, leucine‐rich repeat (NLR) immune receptor gene was isolated using mutagenesis and R gene enrichment sequencing (MutRenSeq). Stable transformation confirmed Pm1a identity which induced a strong resistance phenotype in transgenic plants upon challenge with avirulent Blumeria graminis (wheat powdery mildew) pathogens.A high‐density genetic map of a B. graminis family segregating for Pm1a avirulence combined with pathogen genome resequencing and RNA sequencing (RNAseq) identified AvrPm1a effector gene candidates. In planta expression identified an effector, with an N terminal Y/FxC motif, that induced a strong hypersensitive response when co‐expressed with Pm1a in Nicotiana benthamiana.Single chromosome enrichment sequencing (ChromSeq) and assembly of chromosome 7A suggested that suppressed recombination around the Pm1a region was due to a rearrangement involving chromosomes 7A, 7B and 7D. The cloning of Pm1a and its identification in a highly rearranged region of chromosome 7A provides insight into the role of chromosomal rearrangements in the evolution of this complex resistance cluster. [ABSTRACT FROM AUTHOR]

Iranian wheat landrace accessions (IWAs) were collected from country‐wide farm fields and market places in 1935 by a professor at the University of Tehran and shared with University of California at Davis, California. IWAs were further submitted to the genebank of International Maize and Wheat Improvement Center (CIMMYT), Mexico. 2,403 IWAs from CIMMYT's genebank were assayed by DArT‐seq technology to assess genetic diversity. No apparent ecogeographic patterns related to genetic diversity were detected, probably due to long‐term transport and frequent interchange of landraces among farmers. A multivariate clustering procedure combining genotypic and phenotypic information was used in selecting a core‐set, which represented 15% of the hexaploid wheat accessions included in this study. This subset captured an estimated 93% of rare (frequency <0.05) alleles. Multisite phenotypic data (India, Mexico) validated the ability of the core‐set in detecting useful variants. Potential donor accessions for multiple traits (disease resistance, zinc concentration) were identified from the core‐set for wheat‐breeding. This report illustrates a breeder friendly robust core‐set formulation strategy for harnessing the useful genetic variation stored in the genebanks. [ABSTRACT FROM AUTHOR]

We conducted a five‐year survey (2011–2015) of barley and wheat fields in Paraná state, Brazil, obtaining 754 Fusarium isolates from spikes with fusarium head blight (FHB)‐symptoms. Multilocus genotyping and TEF‐1α gene sequence analyses confirmed the dominance of the F. graminearum species complex (FGSC, 75.7%), but F. poae (11.5%), as well as F. avenaceum and related members of the F. tricinctum species complex (FTSC, 8.1%) appeared as substantial contributors to FHB. Within the FGSC, F. graminearum of the 15‐ADON genotype was dominant (63%), followed by F. meridionale of the NIV genotype (23.1%), F. cortaderiae of the NIV (7%) or 3‐ADON (2.6%) genotypes, and F. austroamericanum (3.8%) of the 3‐ADON genotype. Substantial variation in pathogen composition was observed across years, with F. poae and F. meridionale frequencies significantly elevated in some years. Most F. poae strains produced DAS, diANIV, and butenolide, but not neosolaniol, T‐2, or HT‐2. All FTSC species produced moniliformin. Enniatin production was widespread among FTSC species, with the single F. acuminatum strain found to be the strongest producer of enniatins. Our findings confirm FGSC as a major contributor to FHB and expand considerably our knowledge of the presence, frequency, and conditions under which other pathogens may emerge, altering the spectrum of toxins that may accumulate in grain. [ABSTRACT FROM AUTHOR]