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Many disease resistance genes have been introgressed into wheat from its wild relatives. However, reduced recombination within the introgressed segments hinders the cloning of the introgressed genes. Here, we have cloned the powdery mildew resistance gene Pm13, which is introgressed into wheat from Aegilops longissima, using a method that combines physical mapping with radiation-induced chromosomal aberrations and transcriptome sequencing analysis of ethyl methanesulfonate (EMS)-induced loss-of-function mutants. Pm13 encodes a kinase fusion protein, designated MLKL-K, with an N-terminal domain of mixed lineage kinase domain-like protein (MLKL_NTD domain) and a C-terminal serine/threonine kinase domain bridged by a brace. The resistance function of Pm13 is validated through transient and stable transgenic complementation assays. Transient over-expression analyses in Nicotiana benthamiana leaves and wheat protoplasts reveal that the fragment Brace-Kinase122-476 of MLKL-K is capable of inducing cell death, which is dependent on a functional kinase domain and the three α-helices in the brace region close to the N-terminus of the kinase domain.
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Aegilops , Ascomicetos , Resistencia a la Enfermedad , Enfermedades de las Plantas , Proteínas de Plantas , Triticum , Triticum/microbiología , Triticum/genética , Enfermedades de las Plantas/microbiología , Enfermedades de las Plantas/inmunología , Enfermedades de las Plantas/genética , Proteínas de Plantas/genética , Proteínas de Plantas/metabolismo , Resistencia a la Enfermedad/genética , Aegilops/genética , Aegilops/metabolismo , Plantas Modificadas Genéticamente , Proteínas Quinasas/metabolismo , Proteínas Quinasas/genética , Proteínas Recombinantes de Fusión/metabolismo , Proteínas Recombinantes de Fusión/genética , Nicotiana/genética , Nicotiana/microbiología , Hojas de la Planta/microbiología , Hojas de la Planta/genética , Hojas de la Planta/metabolismo , Regulación de la Expresión Génica de las PlantasRESUMEN
This article investigates the control problem of bearing-based formation tracking for underactuated autonomous underwater vehicles (AUVs) considering actuator constraints and unknown disturbances. A leader-follower structure is adopted, where the leaders move with an unknown reference velocity. For the followers, an integrated strategy is proposed, which includes i) a bearing-based control method composed of a reference velocity estimator, a virtual velocity for achieving the desired formation, and an adaptive robust formation controller to track the virtual velocity under disturbances; and ii) a parameter tuning method based on control parameterization approaches and heuristic algorithms. By employing the cascade system theory, asymptotic convergence of errors in the overall system is proved in the presence of unknown disturbances. The tuning method optimizes controller gains to ensure, all while preserving the convergence properties of the closed-loop error system constraint feasibility and performance optimality. As a result, convergence, robustness, feasibility, and optimality are all achieved. Extension to the case where AUVs have sideslip motions in 3-D space is also discussed. Simulation results are presented to demonstrate the effectiveness of the proposed strategy.
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Rye (Secale cereale), a valuable relative of wheat, contains abundant powdery mildew resistance (Pm) genes. Using physical mapping, transcriptome sequencing, barley stripe mosaic virus-induced gene silencing, ethyl methane sulfonate mutagenesis, and stable transformation, we isolated and validated two coiled-coil, nucleotide-binding site and leucine-rich repeat (CC-NBS-LRR) alleles, PmTR1 and PmTR3, located on rye chromosome 6RS from different triticale lines. PmTR1 confers age-related resistance starting from the three-leaf stage, whereas its allele, PmTR3, confers typical all-stage resistance, which may be associated with their differential gene expression patterns. Overexpression in Nicotiana benthamiana showed that the CC, CC-NBS, and CC-LRR fragments of PMTR1 induce cell death, whereas in PMTR3 the CC and full-length fragments perform this function. Luciferase complementation imaging and pull-down assays revealed distinct interaction activities between the CC and NBS fragments. Our study elucidates two novel rye-derived Pm genes and their derivative germplasm resources and provides novel insights into the mechanism of age-related resistance, which can aid the improvement of resistance against wheat powdery mildew.
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Ascomicetos , Secale , Secale/genética , Resistencia a la Enfermedad/genética , Triticum/genética , Proteínas Repetidas Ricas en Leucina , Ascomicetos/fisiología , Nucleótidos , Cromosomas de las Plantas/genética , Sitios de Unión , Enfermedades de las Plantas/genéticaRESUMEN
This article studies the bearing-based formation tracking control problem of multiple double-integrator agents. A leader-following structure, where the leader moves with the reference dynamics, is adopted. Different from the existing methods, which require complete information of the time-varying reference velocity, in this article, only the time-varying reference orientation information is known by part of the followers and the amplitude of the reference velocity is unknown. To solve the problem, this article proposes a velocity-estimation-based control scheme, which consists of an estimator for estimating the varying rate of the reference orientation, an adaptation law for estimating the amplitude of the reference velocity, and bearing-based control inputs for tracking the leader and achieving the bearing-based formation based on the estimations. Moreover, the scaling formation maneuver can be achieved by using an auxiliary distance measurement. It shows that both the estimation errors and control errors converge to zero under the connectivity of the topology and properties of bearing rigidity. The closed-loop system is analyzed to be semiglobally uniformly asymptotically stable based on the cascaded system theory. Numerical simulations are presented to demonstrate the effectiveness of our method.
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Wheat powdery mildew, caused by Blumeria graminis f. sp. tritici (Bgt), is a devastating disease that threatens wheat production worldwide. Pm12, which originated from Aegilops speltoides, a wild relative of wheat, confers strong resistance to powdery mildew and therefore has potential use in wheat breeding. Using susceptible mutants induced by gamma irradiation, we physically mapped and isolated Pm12 and showed it to be orthologous to Pm21 from Dasypyrum villosum, also a wild relative of wheat. The resistance function of Pm12 was validated via ethyl methanesulfonate mutagenesis, virus-induced gene silencing, and stable genetic transformation. Evolutionary analysis indicates that the Pm12/Pm21 loci in wheat species are relatively conserved but dynamic. Here, we demonstrated that the two orthologous genes, Pm12 and Pm21, possess differential resistance against the same set of Bgt isolates. Overexpression of the coiled-coil domains of both PM12 and PM21 induces cell death in Nicotiana benthamiana leaves. However, their full-length forms display different cell death-inducing activities caused by their distinct intramolecular interactions. Cloning of Pm12 will facilitate its application in wheat breeding programs. This study also gives new insight into two orthologous resistance genes, Pm12 and Pm21, which show different race specificities and intramolecular interaction patterns.
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Fitomejoramiento , Triticum , Triticum/genética , Genes de Plantas , Poaceae/genéticaRESUMEN
In this article, we consider the power scheduling problem of the multihop transmission with limited power resources. For a discrete-time linear time-invariant process, we consider a more practical scenario where the forward-error-correcting (FEC) coding scheme is utilized. An approximate communication model is introduced to formulate the nonanalytical relationship between the consumption of power and the successful-decoding-probability. For the single-hop transmission, we propose an analytical method to figure out the optimal offline scheduling for the finite-time case and the optimal periodic schedule for the infinite-time case. We consider the process and terminal errors simultaneously, and explicitly discuss how different values of parameters affect the optimality. Moreover, we extend our conclusions to the multihop case. In order to deal with the difficulty and complexity brought by the multihop scenario, a novel method based on the equivalent-scheduling matrix (ESM) is proposed to describe the accumulated effects through the multihop transmission. Meanwhile, explicit solutions of the multihop case are provided for finite- and infinite-time cases, respectively. Numerical examples are provided to demonstrate the effectiveness of the proposed methods.
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Wheat powdery mildew is a devastating disease leading to severe yield loss. The powdery mildew resistance gene Pm21, encoding a nucleotide-binding leucine-rich repeat receptor (NLR) protein, confers broad-spectrum resistance to powdery mildew and has great potential for controlling this disease. In this study, a large-scale mutagenesis was conducted on wheat cultivar (cv.) Yangmai 18 carrying Pm21. As a result, a total of 113 independent mutant lines susceptible to powdery mildew were obtained, among which, only one lost the whole Pm21 locus and the other 112 harbored one- (107) or two-base (5) mutations in the encoding region of Pm21. From the 107 susceptible mutants containing one-base change, we found that 25 resulted in premature stop codons leading to truncated proteins and 82 led to amino acid changes involving in 59 functional sites. We determined the mutations per one hundred amino acids (MPHA) indexes of different domains, motifs, and non-domain and non-motif regions of PM21 protein and found that the loss-of-function mutations occurred in a tendentious means. We also observed a new mutation hotspot that was closely linked to RNBS-D motif of the NB-ARC domain and relatively conserved in different NLRs of wheat crops. In addition, we crossed all the susceptible mutants with Yangmai 18 carrying wild-type Pm21, subsequently phenotyped their F1 plants and revealed that the variant E44K in the coiled-coil (CC) domain could lead to dominant-negative effect. This study revealed key functional sites of PM21 and their distribution characteristics, which would contribute to understanding the relationship of resistance and structure of Pm21-encoded NLR.
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In recent decades, although the research on gait recognition of lower limb exoskeleton robot has been widely developed, there are still limitations in rehabilitation training and clinical practice. The emergence of interactive information fusion technology provides a new research idea for the solution of this problem, and it is also the development trend in the future. In order to better explore the issue, this paper summarizes gait recognition based on interactive information fusion of lower limb exoskeleton robots. This review introduces the current research status, methods, and directions for information acquisition, interaction, fusion, and gait recognition of exoskeleton robots. The content involves the research progress of information acquisition methods, sensor placements, target groups, lower limb sports biomechanics, interactive information fusion, and gait recognition model. Finally, the current challenges, possible solutions, and promising prospects are analysed and discussed, which provides a useful reference resource for the study of interactive information fusion and gait recognition of rehabilitation exoskeleton robots.
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Powdery mildew is one of the most destructive diseases in wheat production. Identifying novel resistance genes and deploying them in new cultivars is the most effective approach to minimize wheat losses caused by powdery mildew. In this study, wheat breeding line PBDH1607 showed high resistance to powdery mildew at both the seedling and adult plant stages. Genetic analysis of the seedling data demonstrated that the resistance was controlled by a single dominant gene, tentatively designated PmPBDH. The ΔSNP index based on bulked segregant RNA sequencing indicated that PmPBDH was associated with an interval of about 30.8 Mb (713.5 to 744.3 Mb) on chromosome arm 4AL. Using newly developed markers, we mapped PmPBDH to a 3.2-cM interval covering 7.1 Mb (719,055,516 to 726,215,121 bp). This interval differed from those of Pm61 (717,963,176 to 719,260,469 bp), MlIW30 (732,769,506 to 732,790,522 bp), and MlNSF10 (729,275,816 to 731,365,462 bp) reported on the same chromosome arm. PmPBDH also differed from Pm61, MlIW30, and MlNSF10 by its response spectrum, origin, or inheritance mode, suggesting that PmPBDH should be a new Pm gene. In the candidate interval, five genes were found to be associated with PmPBDH via time course gene expression analysis, and thus they are candidate genes of PmPBDH. Six closely linked markers, including two kompetitive allele-specific PCR markers, were confirmed to be applicable for tracking PmPBDH in marker-assisted breeding.
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Ascomicetos , Triticum , Ascomicetos/fisiología , Mapeo Cromosómico , Resistencia a la Enfermedad/genética , Genes de Plantas/genética , Marcadores Genéticos , Fitomejoramiento , Enfermedades de las Plantas/genética , Triticum/genéticaRESUMEN
The new generation of the industrial cyber-physical system (ICPS) supported by the edge computing technology facilitates the deep integration of sensing and control. System observability is the key factor to characterize the internal relationship of them. In most existing works, the observability is regarded as the assumption for subsequent sensing and control. But, in fact, with the gradually expanded network scale, this assumption is more difficult to directly satisfy sensing design. For this problem, we propose the observability guaranteed method (OGM) for edge sensing and control co-design. Specifically, the nonconvex observability condition is transformed into the convex range of key parameters of the sensing strategy based on the graph signal processing (GSP) technology. Then, we establish the relationship between these parameters and control performance. In OGM, except the previous design from sensing to control, we reversely adjust the sensing design for control demands to satisfy observability. Finally, our algorithm is applied into the hot rolling laminar cooling process based on the semiphysical evaluation. The effectiveness is verified by the results.
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Wheat powdery mildew is a devastating disease that seriously threatens yield worldwide. Utilization of host resistance is considered an effective strategy to minimize powdery mildew damage. Pm21, PmV, and Pm12 confer broad-spectrum resistance to wheat powdery mildew in China, of which Pm21 and PmV are allelic genes derived from the 6VS chromosome of Dasypyrum villosum, and Pm12 is derived from the 6SS chromosome of Aegilops speltoides and most likely orthologous to the former two genes. To accurately and efficiently transfer and pyramid these genes using marker-assisted selection (MAS), distinctive single-nucleotide polymorphisms (SNPs) among the exon sequences of Pm21, PmV, and Pm12 and their homologous sequences in the common wheat genome were identified and then used for developing diagnostic Kompetitive Allele-Specific PCR (KASP) markers. The markers were validated in different genotypes including transgenic vectors, transgenic lines, translocation lines, resistance stocks with documented Pm genes, and in multiple susceptible cultivars without Pm genes. As a result, we initially developed a KASP marker that can simultaneously diagnose Pm21, Pm12, and PmV. Subsequently, we obtained a highly diagnostic KASP marker for each of the three genes that could distinguish among the three genes and also accurately distinguish them from other resistant stocks with documented Pm genes and from multiple susceptible genotypes. Compared with previously reported markers, the highly diagnostic KASP markers developed in this study have the advantages of low cost, easy assay, accuracy, and potentially high throughput for MAS.
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Resistencia a la Enfermedad , Triticum , Alelos , Resistencia a la Enfermedad/genética , Genes de Plantas/genética , Marcadores Genéticos/genética , Enfermedades de las Plantas/genética , Reacción en Cadena de la Polimerasa , Triticum/genéticaRESUMEN
KEY MESSAGE: PmSESY, a new wheat powdery mildew resistance gene was characterized and genetically mapped to the terminal region of chromosome 1RL of wild rye Secale sylvestre. The genus Secale is an important resource for wheat improvement. The Secale species are usually considered as non-adapted hosts of Blumeria graminis f. sp. tritici (Bgt) that causes wheat powdery mildew. However, as a wild species of cultivated rye, S. sylvestre is rarely studied. Here, we reported that 25 S. sylvestre accessions were susceptible to isolate BgtYZ01, whereas the other five confer effective resistance to all the tested isolates of Bgt. A population was then constructed by crossing the resistant accession SESY-01 with the susceptible accession SESY-11. Genetic analysis showed that the resistance in SESY-01 was controlled by a single dominant gene, temporarily designated as PmSESY. Subsequently, combining bulked segregant RNA-Seq (BSR-Seq) analysis with molecular analysis, PmSESY was mapped into a 1.88 cM genetic interval in the terminus of the long arm of 1R, which was closely flanked by markers Xss06 and Xss09 with genetic distances of 0.87 cM and 1.01 cM, respectively. Comparative mapping demonstrated that the corresponding physical region of the PmSESY locus was about 3.81 Mb in rye cv. Lo7 genome, where 30 disease resistance-related genes were annotated, including five NLR-type disease resistance genes, three kinase family protein genes, three leucine-rich repeat receptor-like protein kinase genes and so on. This study gives a new insight into S. sylvestre that shows divergence in response to Bgt and reports a new powdery mildew resistance gene that has potential to be used for resistance improvement in wheat.
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Ascomicetos/fisiología , Resistencia a la Enfermedad/genética , Regulación de la Expresión Génica de las Plantas , Enfermedades de las Plantas/genética , Proteínas de Plantas/genética , Secale/genética , Mapeo Cromosómico/métodos , Cromosomas de las Plantas/genética , Resistencia a la Enfermedad/inmunología , Ligamiento Genético , Marcadores Genéticos , Enfermedades de las Plantas/microbiología , Secale/inmunología , Secale/microbiologíaRESUMEN
KEY MESSAGE: New powdery mildew resistance gene Pm68 was found in the terminal region of chromosome 2BS of Greek durum wheat TRI 1796. The co-segregated molecular markers could be used for MAS. Durum wheat (Triticum turgidum L. var. durum Desf.) is not only an important cereal crop for pasta making, but also a genetic resource for common wheat improvement. In the present study, a Greek durum wheat TRI 1796 was found to confer high resistance to all 22 tested isolates of Blumeria graminis f. sp. tritici (Bgt). Inheritance study on the F1 plants and the F2 population derived from the cross TRI 1796/PI 584832 revealed that the resistance in TRI 1796 was controlled by a single dominant gene, herein designated Pm68. Using the bulked segregant RNA-Seq (BSR-Seq) analysis combined with molecular analysis, Pm68 was mapped to the terminal part of the short arm of chromosome 2B and flanked by markers Xdw04 and Xdw12/Xdw13 with genetic distances of 0.22 cM each. According to the reference genome of durum wheat cv. Svevo, the corresponding physical region spanned the Pm68 locus was about 1.78-Mb, in which a number of disease resistance-related genes were annotated. This study reports the new powdery mildew resistance gene Pm68 that would be a valuable resource for improvement of both common wheat and durum wheat. The co-segregated markers (Xdw05-Xdw11) developed here would be useful tools for marker-assisted selection (MAS) in breeding.
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Ascomicetos/patogenicidad , Resistencia a la Enfermedad/genética , Enfermedades de las Plantas/genética , Mapeo Cromosómico , Cromosomas de las Plantas , Hibridación Genómica Comparativa , Cruzamientos Genéticos , Genes Dominantes , Genes de Plantas , Marcadores Genéticos , Grecia , Enfermedades de las Plantas/microbiología , RNA-Seq , Triticum/genética , Triticum/microbiologíaRESUMEN
The Yellow River Estuary (YRE) and adjacent Laizhou Bay (LB) encounter eco-environmental risks caused by heavy metals (HMs) pollution. Here magnetic measurements were performed on 239 surface sediment samples from the YRE and LB to establish a rapid and effective method for detecting HMs. Magnetite, maghemite, and hematite coexist in the sediments. The distributions of magnetic minerals are dominated by sediment sources (Yellow River in northern and western LB, and rivers in southern and eastern coastal LB), and the anticlockwise water current. Compared to the background values, Cd content is enriched for all samples, while Co, Cr, Ni, Cu, Zn, and Pb contents are lower for most samples. The low pollution load indexes (PLI) of HMs (< 1-1.56) indicate the unpolluted to moderately polluted status, while the muddy area is the most polluted. The principal component analysis indicates that Co, Cr, Ni, Cu, and Zn are mainly from natural weathering substances, while Cd and Pb are anthropogenic. Contents of fine-grained sediments and magnetic particles are positively correlated to Co, Ni, Cu, Zn, and PLI. The high-risk Co, Ni, Cu, and Zn regions can be quickly delineated with the frequency-dependent susceptibility.
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Wheat powdery mildew caused by Blumeria graminis f. sp. tritici (Bgt) is a devastating disease that threatens wheat production and yield worldwide. The powdery mildew resistance gene Pm21, originating from wheat wild relative Dasypyrum villosum, encodes a coiled-coil, nucleotide-binding site, leucine-rich repeat (CC-NBS-LRR) protein and confers broad-spectrum resistance to wheat powdery mildew. In the present study, we isolated 73 Pm21 alleles from different powdery mildew-resistant D. villosum accessions, among which, 38 alleles were non-redundant. Sequence analysis identified seven minor insertion-deletion (InDel) polymorphisms and 400 single nucleotide polymorphisms (SNPs) among the 38 non-redundant Pm21 alleles. The nucleotide diversity of the LRR domain was significantly higher than those of the CC and NB-ARC domains. Further evolutionary analysis indicated that the solvent-exposed LRR residues of Pm21 alleles had undergone diversifying selection (dN/dS = 3.19734). In addition, eight LRR motifs and four amino acid sites in the LRR domain were also experienced positive selection, indicating that these motifs and sites play critical roles in resistance specificity. The phylogenetic tree showed that 38 Pm21 alleles were divided into seven classes. Classes A (including original Pm21), B and C were the major classes, including 26 alleles (68.4%). We also identified three non-functional Pm21 alleles from four susceptible homozygous D. villosum lines (DvSus-1 to DvSus-4) and two susceptible wheat-D. villosum chromosome addition lines (DA6V#1 and DA6V#3). The genetic variations of non-functional Pm21 alleles involved point mutation, deletion and insertion, respectively. The results also showed that the non-functional Pm21 alleles in the two chromosome addition lines both came from the susceptible donors of D. villosum. This study gives a new insight into the evolutionary characteristics of Pm21 alleles and discusses how to sustainably utilize Pm21 in wheat production. This study also reveals the sequence variants and origins of non-functional Pm21 alleles in D. villosum populations.
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Nucleotide-binding (NB) leucine-rich repeat (LRR) receptors (NLRs) provide resistance against several plant pathogens. We previously cloned the wheat powdery mildew resistance gene Pm21, which encodes a coiled-coil (CC) NLR that confers broad-spectrum resistance against Blumeria graminis f. sp. tritici. Here, we report comprehensive biochemical and functional analyses of Pm21 CC domain in Nicotiana benthamiana. Transient overexpression assay suggested that only the extended CC (eCC, amino acid residues 1-159) domain has cell-death-inducing activity, whereas the CC-containing truncations, including CC-NB and CC-NB-LRR, do not induce cell-death responses. Coimmunoprecipitation (Co-IP) assay showed that the eCC domain self-associates and interacts with the NB and LRR domains in planta. These results imply that the activity of the eCC domain is inhibited by the intramolecular interactions of different domains in the absence of pathogens. We found that the LRR domain plays a crucial role in D491V-mediated full-length (FL) Pm21 autoactivation. Some mutations in the CC domain leading to the loss of Pm21 resistance to powdery mildew impaired the CC activity of cell-death induction. Two mutations (R73Q and E80K) interfered with D491V-mediated Pm21 autoactivation without affecting the cell-death-inducing activity of the eCC domain. Notably, some susceptible mutants harbouring mutations in the CC domain still exhibited cell-death-inducing activity. Taken together, these results implicate the CC domain of Pm21 in cell-death signalling and disease-resistance signalling, which are potentially independent of each other.
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Muerte Celular , Resistencia a la Enfermedad/genética , Proteínas NLR/fisiología , Enfermedades de las Plantas/genética , Dominios Proteicos/fisiología , Triticum/inmunología , Triticum/microbiología , Mutación , Proteínas NLR/química , Proteínas NLR/genética , Células Vegetales/patología , Transducción de Señal , NicotianaRESUMEN
In this paper, we investigate the transmission scheduling problem for wireless control systems (WCSs) with limited power resources. Different from the existing works, for a discrete-time linear process, we consider a more practical WCS, where a relay is introduced into the framework for remote transmission and control. To achieve the best control performance of the system, we propose a global optimal offline scheduling algorithm. Then, based on ACK-feedback framework, two different online scheduling schemes are further designed respectively under the given power resources. Theoretically, we prove the superiority of online schedule to the offline one under the same energy budget. Simulations are conducted to demonstrate and verify the effectiveness of the proposed algorithms.
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Pm21, derived from wheat wild relative Dasypyrum villosum, is one of the most effective powdery mildew resistance genes and has been widely applied in wheat breeding in China. Mapping and cloning Pm21 are of importance for understanding its resistance mechanism. In the present study, physical mapping was performed using different genetic stocks involving in structural variations of chromosome 6VS carrying Pm21. The data showed that 6VS could be divided into eight distinguishable chromosomal bins, and Pm21 was mapped to the bin FLb4-b5/b6 closely flanked by the markers 6VS-08.6 and 6VS-10.2. Comparative genomic mapping indicated that the orthologous regions of FLb4-b5/b6 carrying Pm21 were narrowed to a 117.7 kb genomic region harboring 19 genes in Brachypodium and a 37.7 kb region harboring 5 genes in rice, respectively. The result was consistent with that given by recent genetic mapping in diploid D. villosum. In conclusion, this study demonstrated that physical mapping based on chromosomal structural variations is an efficient method for locating alien genes in wheat background.
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Cromosomas de las Plantas , Resistencia a la Enfermedad/genética , Genes de Plantas , Variación Genética , Interacciones Huésped-Patógeno/efectos de los fármacos , Enfermedades de las Plantas/genética , Triticum/genética , Triticum/microbiología , Mapeo Cromosómico , Biología Computacional/métodos , Evolución Molecular , Marcadores Genéticos , Genoma de Planta , Genómica/métodos , Enfermedades de las Plantas/microbiología , Polimorfismo GenéticoAsunto(s)
Resistencia a la Enfermedad , Enfermedades de las Plantas/microbiología , Proteínas de Plantas/genética , Proteínas de Plantas/metabolismo , Triticum/inmunología , Triticum/microbiología , Ascomicetos/fisiología , Enfermedades de las Plantas/inmunología , Triticum/genética , Triticum/metabolismoRESUMEN
Pm21, originating from wheat wild relative Dasypyrum villosum, confers immunity to all known races of Blumeria graminis f. sp. tritici (Bgt) and has been widely utilized in wheat breeding. However, little is known on the genetic basis of the Pm21 locus. In the present study, four seedling-susceptible D. villosum lines (DvSus-1 â¼ DvSus-4) were identified from different natural populations. Based on the collinearity among genomes of Brachypodium distachyon, Oryza, and Triticeae, a set of 25 gene-derived markers were developed declaring the polymorphisms between DvRes-1 carrying Pm21 and DvSus-1. Fine genetic mapping of Pm21 was conducted by using an extremely large F2 segregation population derived from the cross DvSus-1/DvRes-1. Then Pm21 was narrowed to a 0.01-cM genetic interval defined by the markers 6VS-08.4b and 6VS-10b. Three DNA markers, including a resistance gene analog marker, were confirmed to co-segregate with Pm21. Moreover, based on the susceptible deletion line Y18-S6 induced by ethyl methanesulfonate treatment conducted on Yangmai 18, Pm21 was physically mapped into a similar interval. Comparative analysis revealed that the orthologous regions of the interval carrying Pm21 were narrowed to a 112.5 kb genomic region harboring 18 genes in Brachypodium, and a 23.2 kb region harboring two genes in rice, respectively. This study provides a high-density integrated map of the Pm21 locus, which will contribute to map-based cloning of Pm21.